Structure of bacterial flagellum.
SEM image of flagellated Chlamydomonas sp. (10000×)
A flagellum (//; plural: flagella) is a lash-like appendage that protrudes from the cell body of certain prokaryotic and eukaryotic cells. The word flagellum in Latin means whip. The primary role of the flagellum is locomotion but it also often has function as a sensory organelle, being sensitive to chemicals and temperatures outside the cell. Flagella are organelles defined by function rather than structure. There are large differences between different types of flagella; the prokaryotic and eukaryotic flagella differ greatly in protein composition, structure, and mechanism of propulsion. However, both are used for swimming.
An example of a flagellate bacterium is the ulcer-causing Helicobacter pylori, which uses multiple flagella to propel itself through the mucus lining to reach the stomach epithelium. An example of a eukaryotic flagellate cell is the mammalian sperm cell, which uses its flagellum to propel itself through the female reproductive tract. Eukaryotic flagella are structurally identical to eukaryotic cilia, although distinctions are sometimes made according to function and/or length.
Three types of flagella have so far been distinguished; bacterial, archaeal and eukaryotic.
The main differences among these three types are summarized below:
- Bacterial flagella are helical filaments, each with a rotary motor at its base which can turn clockwise or counterclockwise. They provide two of several kinds of bacterial motility.
- Archaeal flagella (Archaella) are superficially similar to bacterial flagella, but are different in many details and considered non-homologous.
- Eukaryotic flagella - those of animal, plant, and protist cells - are complex cellular projections that lash back and forth. Eukaryotic flagella are classed along with eukaryotic motile cilia as undulipodia to emphasize their distinctive wavy appendage role in cellular function or motility. Primary cilia are immotile, and are not undulipodia; they have a structurally different 9+0 axoneme rather than the 9+2 axoneme found in both flagella and motile cilia undulipodia.
Structure and composition. The bacterial flagellum is made up of the protein flagellin. 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 plasma membrane. The filament ends with a capping protein.
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 eleven protofilaments approximately parallel to the filament axis. Each protofilament is a series of tandem protein chains. However in Campylobacter jejuni, there are seven protofilaments.
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. Given the structural similarities between bacterial flagella and bacterial secretory systems, it is thought that bacterial flagella may have evolved from the type three secretion system; however, it is not known for certain whether these pores are derived from the bacterial flagella or the bacterial secretory system.
Motor. The bacterial flagellum is driven by a rotary engine (the 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 (in Vibrio species there are two kinds of flagella, lateral and polar, and some are driven by a sodium ion pump rather than a proton pump). The rotor transports protons across the membrane, and is turned in the process. The rotor alone can operate at 6,000 to 17,000 rpm, but with the flagellar filament attached usually only reaches 200 to 1000 rpm. The direction of rotation can be switched almost instantaneously, caused by a slight change in the position of a protein, FliG, in the rotor. The flagellum is highly energy efficient and uses very little energy.
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.
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 / second. Although at such a speed it would take a bacterium about 245 days to cover a kilometre, and 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 of body lengths per second. A cheetah for example, only achieves about 25 body lengths / sec.
Through use of their flagella, E. coli are able to move rapidly towards attractants and away from repellents. They do this by means of a biased random walk, with 'runs' and 'tumbles' brought about by rotating the flagellum counterclockwise and clockwise respectively.
Assembly. During flagellar assembly, components of the flagellum pass through the hollow cores of the basal body and the nascent filament. During assembly, protein components are added at the flagellar tip rather than at the base. In vitro, flagellar filaments assemble spontaneously in a solution containing purified flagellin as the sole protein.
Evolution. The evolution of bacterial flagella has been used as an argument against evolution by creationists. They argue that complex structures like flagella cannot evolve from simple structures. In other words, flagella are "irreducibly complex" and need all of their protein components to function. However, it has been shown by numerous studies that a large number of proteins can be deleted without (complete) loss of function. Moreover, it is generally accepted now that bacterial flagella have evolved from much simpler secretion systems, such as the Type III secretion system.
Flagellar arrangement schemes
Different species of bacteria have different numbers and arrangements of flagella.
- Monotrichous bacteria have a single flagellum (e.g., Vibrio cholerae).
- Lophotrichous bacteria have multiple flagella located at the same spot on the bacteria's surfaces which act in concert to drive the bacteria in a single direction. In many cases, the bases of multiple flagella are surrounded by a specialized region of the cell membrane, the so-called polar organelle.
- Amphitrichous bacteria have a single flagellum on each of two opposite ends (only one flagellum operates at a time, allowing the bacteria to reverse course rapidly by switching which flagellum is active).
- Peritrichous bacteria have flagella projecting in all directions (e.g., E. coli).
In certain large forms of Selenomonas, more than 30 individual flagella are organized outside the cell body, helically twining about each other to form a thick structure (easily visible with the light microscope) called a "fascicle". Other bacteria, such as most Spirochetes, have two or more specialized flagella (endoflagella) arising from opposite poles of the cell, which together constitute the so-called "axial filament" that is located within the periplasmic space between the flexible cell wall and an outer sheath. The rotation of the axial filament relative to the cell body causes the entire bacterium to move forward in a corkscrew-like motion, even through material viscous enough to prevent the passage of normally flagellated bacteria.
Counterclockwise rotation of a monotrichous polar flagellum pushes the cell forward with the flagellum trailing behind, much like a corkscrew moving inside cork. Indeed water on the microscopic scale is highly viscous, very different from our daily experience of water.
Flagella are left-handed helices, and bundle and rotate together only when rotating counterclockwise. When some of the rotors reverse direction, the flagella unwind and the cell starts "tumbling". It has also been suggested that even if all flagella would rotate clockwise, they will not form a bundle, due to geometrical as well as hydrodynamic reasons. Such "tumbling" may happen occasionally, leading to the cell seemingly thrashing about in place, resulting in the reorientation of the cell. The clockwise rotation of a flagellum is suppressed by chemical compounds favorable to the cell (e.g. food), but the motor is highly adaptive to this. Therefore, when moving in a favorable direction, the concentration of the chemical attractant increases and "tumbles" are continually suppressed; however, when the cell's direction of motion is unfavorable (e.g., away from a chemical attractant), tumbles are no longer suppressed and occur much more often, with the chance that the cell will be thus reoriented in the correct direction.
In some Vibrio spp. (particularly Vibrio parahemolyticus) and related proteobacteria such as Aeromonas, two flagellar systems co-exist, using different sets of genes and different ion gradients for energy. The polar flagella are constitutively expressed and provide motility in bulk fluid, while the lateral flagella are expressed when the polar flagella meet too much resistance to turn. These provide swarming motility on surfaces or in viscous fluids.
The archaeal flagellum (Archaellum) is superficially similar to the bacterial (or eubacterial) flagellum; in the 1980s they were thought to be homologous on the basis of gross morphology and behavior. Both flagella and archaella consist of filaments extending outside the cell, and rotate to propel the cell. Archaeal flagella have a unique structure which lacks a central channel. Similar to bacterial type IV pilins, the archaeal flagellins (archaellins) are made with class 3 signal peptides and they are processed by a type IV prepilin peptidase-like enzyme. The archaellins are typically modified by the addition of N-linked glycans which are necessary for proper assembly and/or function.
Discoveries in the 1990s revealed numerous detailed differences between the archaeal and bacterial flagella; these include:
- Bacterial flagella are motorized by a flow of H+ ions (or occasionally Na+ ions); archaeal flagella are almost certainly powered by ATP. The torque-generating motor that powers rotation of the archaeal flagellum has not been identified.
- While bacterial cells often have many flagellar filaments, each of which rotates independently, the archaeal flagellum is composed of a bundle of many filaments that rotate as a single assembly.
- Bacterial flagella grow by the addition of flagellin subunits at the tip; archaeal flagella grow by the addition of subunits to the base.
- Bacterial flagella are thicker than archaella [Singular Archaellum], and the bacterial filament has a large enough hollow "tube" inside that the flagellin subunits can flow up the inside of the filament and get added at the tip; the archaellum is too thin (12-15 nm) to allow this.
- Many components of bacterial flagella share sequence similarity to components of the type III secretion systems, but the components of bacterial flagella and archaella share no sequence similarity. Instead, some components of archaella share sequence and morphological similarity with components of type IV pili, which are assembled through the action of type II secretion systems (the nomenclature of pili and protein secretion systems is not consistent).
These differences could mean that the bacterial flagella and archaella could be a classic case of biological analogy, or convergent evolution, rather than homology. However, in comparison to the decades of well-publicized study of bacterial flagella (e.g. by Howard Berg), archaella have only recently[when?] begun to get serious scientific attention. Therefore, many[who?] assume erroneously that there is only one basic kind of prokaryotic flagellum, and that archaella are homologous to it. For example, Cavalier-Smith (2002) is aware of the differences between bacterial flagellins and archaeal flagellins (archaellins).
Aiming to emphasize the distinction between the bacterial flagella and the eukaryotic cilia and flagella, some authors attempted to replace the name of these two eukaryotic structures with "undulipodia" (e.g., all papers by Margulis since the 1970s) or "cilia" for both (e.g., Hülsmann, 1992; Adl et al., 2012; most papers of Cavalier-Smith), preserving "flagella" for the bacterial structure. However, the discriminative usage of the terms "cilia" and "flagella" for eukaryotes adopted in this article is still common (e.g., Andersen et al., 1991; Leadbeater et al., 2000).
A eukaryotic flagellum is a bundle of nine fused pairs of microtubule doublets surrounding two central single microtubules. The so-called "9 + 2" structure is characteristic of the core of the eukaryotic flagellum called an axoneme. At the base of a eukaryotic flagellum is a basal body, "blepharoplast" or kinetosome, which is the microtubule organizing center (MTOC) for flagellar microtubules and is about 500 nanometers long. Basal bodies are structurally identical to centrioles. The flagellum is encased within the cell's plasma membrane, so that the interior of the flagellum is accessible to the cell's cytoplasm.
Besides the axoneme and basal body, relatively constant in morphology, other internal structures of the flagellar apparatus are the transition zone (where the axoneme and basal body meet) and the root system (microtubular or fibrilar structures which extends from the basal bodies into the cytoplasm), more variable and useful as indicators of phylogenetic relationships of eukaryotes. Other structures, more uncommon, are the paraflagellar (or paraxial, paraxonemal) rod, the R fiber and the S fiber. For surface structures, see below.
Each of the outer nine doublet microtubules extends a pair of dynein arms (an "inner" and an "outer" arm) to the adjacent microtubule; these dynein arms are responsible for flagellar beating, as the force produced by the arms causes the microtubule doublets to slide against each other and the flagellum as a whole to bend. These dynein arms produce force through ATP hydrolysis. The flagellar axoneme also contains radial spokes, polypeptide complexes extending from each of the outer nine microtubule doublets towards the central pair, with the "head" of the spoke facing inwards. The radial spoke is thought to be involved in the regulation of flagellar motion, although its exact function and method of action are not yet understood.
Flagella vs cilia
The regular beat patterns of eukaryotic cilia and flagella generate motion on a cellular level. Examples range from the propulsion of single cells such as the swimming of spermatozoa to the transport of fluid along a stationary layer of cells such as in the respiratory tract. Though eukaryotic flagella and motile cilia are ultrastructurally identical, the beating pattern of the two organelles can be different. In the case of flagella, the motion is often planar and wave-like, whereas the motile cilia often perform a more complicated three-dimensional motion with a power and recovery stroke.
Intraflagellar transport (IFT), the process by which axonemal subunits, transmembrane receptors, and other proteins are moved up and down the length of the flagellum, is essential for proper functioning of the flagellum, in both motility and signal transduction.
Evolution and occurrence
Eukaryotic flagella or cilia, probably an ancestral characteristic, are widespread in almost all groups of eukaryotes, as a relatively perennial condition, or as a flagellated life cycle stage (e.g., zoids, gametes, zoospores, which may be produced continually or not).
The first situation is found either in specialized cells of multicellular organisms (e.g., in the ciliated epithelia of metazoans), as in ciliates and many eukaryotes with a "flagellate condition" (or "monadoid level of organization", see Flagellata, an artificial group).
Flagellated life cycle stages are found in many groups, e.g., many green algae (zoospores and male gametes), bryophytes (male gametes), pteridophytes (male gametes), some gymnosperms (cycads and Ginkgo, as male gametes), centric diatoms (male gametes), brown algae (zoospores and gametes), oomycetes (assexual zoospores and gametes), hyphochytrids (zoospores), labyrinthulomycetes (zoospores), some apicomplexans (gametes), some radiolarians (probably gametes), foraminiferans (gametes), plasmodiophoromycetes (zoospores and gametes), myxogastrids (zoospores), metazoans (male gametes), chytrid fungi (zoospores and gametes).
Flagella or cilia are completely absent in some groups, probably due to a loss rather than being a primitive condition. The loss of cilia occurred in red algae, some green algae (Zygnematophyceae), some gymnosperms, angiosperms, pennate diatoms, some apicomplexans, some amoebozoans, in the sperm of some metazoans, and in fungi (except chytrids).
- whiplash (= smooth, acronematic flagella): without hairs, e.g., in Opisthokonta
- hairy (= tinsel, flimmer, pleuronematic flagella): with hairs (= mastigonemes sensu lato), divided in:
- fine (= non tubular, or simple hairs): occurs in Euglenophyceae, Dinoflagellata, some Haptophyceae (Pavlovales)
- stiff (= tubular hairs, retronemes, mastigonemes sensu stricto), divided in:
- stichonematic: with single row of hairs
- pantonematic: with two rows of hairs
- with scales: e.g., Prasinophyceae
- with spines: e.g., some brown algae
- with undulating membrane: e.g., some kinetoplastids, some parabasalids
- uniflagellated: e.g., most Opisthokonta
- biflagellated: e.g., all Dinoflagellata, the gametes of Charophyceae, of most bryophytes and of some metazoans
- triflagellated: e.g., the gametes of some Foraminifera
- quadriflagellated: e.g., some Prasinophyceae
- octoflagellated: e.g., Diplomonada, some Prasinophyceae
- multiflagellated: e.g., Opalinata, Ciliophora, Stephanopogon, Parabasalida, Hemimastigophora, Caryoblastea, Multicilia, the gametes (or zoids) of some pteridophytes and some gymnosperms
Other terms related to the flagellar type:
- isokont: cells with flagella of equal length. It was also formerly used to refer to the Chlorophyta
- anisokont: cells with flagella of unequal length, e.g., some Euglenophyceae and Prasinophyceae
- heterokont: term introduced by Luther (1899) to refer to the Xanthophyceae, due to the pair of flagella of unequal length. It has taken on a specific meaning in referring to cells with an anterior straminipilous flagellum (with tripartite mastigonemes, in one or two rows) and a posterior usually smooth flagellum. It is also used to refer to the taxon Heterokonta
- opisthokont: cells with a single posterior whiplash flagella. Term introduced by Vischer (1945), also used to refer to the taxon Opisthokonta
- stephanokont: cells with a crown of flagella near its anterior end. Term introduced by Blackman & Tansley (1902) to refer to the Oedogoniales
- Aconta or Akonta: used to refer to the Zygnematophyceae and Bacillariophyceae (Oltmanns, 1904), or Rhodophyceae (Christensen, 1962), due to their absence of flagella
- Evolution of flagella
- Genetic ciliopathy
- Rotating locomotion in living systems
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- Berg, Howard C. (January 2000). "Motile Behavior of Bacteria". Physics Today 53 (1): 24. doi:10.1063/1.882934.
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- N. J. Matzke (2003-11-10). "Evolution in (Brownian) space: a model for the origin of the bacterial flagellum". www.talkdesign.org.
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This article incorporates text from a publication now in the public domain: Chambers, Ephraim, ed. (1728). "article name needed". Cyclopædia, or an Universal Dictionary of Arts and Sciences (first ed.). James and John Knapton, et al.