A flagellum (/fləˈdʒɛləm/; plural: flagella) is a lash-like appendage that protrudes from the cell body of certain bacteria and eukaryotic cells termed as flagellates. A flagellate can have one or several flagella. The primary function of a flagellum is that of locomotion, but it also often functions as a sensory organelle, being sensitive to chemicals and temperatures outside the cell. The similar structure in the archaea functions in the same way but is structurally different and has been termed the archaellum.
Flagella are organelles defined by function rather than structure. Flagella vary greatly. Both prokaryotic and eukaryotic flagella can be used for swimming but they differ greatly in protein composition, structure, and mechanism of propulsion. The word flagellum in Latin means whip.
An example of a flagellated 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 or length. Fimbriae and pili are also thin appendages, but have different functions and are usually smaller.
Structure of bacterial flagellum.
Three types of flagella have so far been distinguished: bacterial, archaeal, and eukaryotic.
The main differences among these three types are:
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 11 protofilaments approximately parallel to the filament axis. Each protofilament is a series of tandem protein chains. However, Campylobacter jejuni has 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. 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.
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). 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 changed by the flagellar motor switch 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 exact mechanism for torque generation is still poorly understood. 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.
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 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 of body lengths per second. A cheetah, for example, only achieves about 25 body lengths per second.
Through use of their flagella, E. coli is able to move rapidly towards attractants and away from repellents, by means of a biased random walk, with 'runs' and 'tumbles' brought about by rotating its flagellum counterclockwise and clockwise, respectively. The two directions of rotation are not identical (with respect to flagellum movement) and are selected by a molecular switch.
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.
At least 10 protein components of the bacterial flagellum share homologous proteins with the type three secretion system (TTSS), hence one likely evolved from the other. Because the TTSS has a similar number of components as a flagellar apparatus (about 25 proteins), which one evolved first is difficult to determine. However, the flagellar system appears to involve more proteins overall, including various regulators and chaperones, hence it has been argued that flagella evolved from a TTSS. However, it has also been suggested that the flagellum may have evolved first or the two structures evolved in parallel. Early single-cell organisms' need for motility (mobility) support that the more mobile flagella would be selected by evolution first, but the TTSS evolving from the flagellum can be seen as 'reductive evolution', and receives no topological support from the phylogenetic trees. The hypothesis that the two structures evolved separately from a common ancestor accounts for the protein similarities between the two structures, as well as their functional diversity.
Some authors have argued that flagella cannot have evolved, assuming that they can only function properly when all proteins are in place In other words, the flagellar apparatus is "irreducibly complex". However, many proteins can be deleted or mutated and the flagellum still works, though sometimes at reduced efficiency. In addition, the composition of flagella is surprisingly diverse across bacteria, with many proteins only found in some species, but not others. Hence, the flagellar apparatus is clearly very flexible in evolutionary terms and perfectly able to lose or gain protein components. For instance, a number of mutations have been found that increase the motility of E. coli. Additional evidence for the evolution of bacterial flagella includes the existence of vestigial flagella, intermediate forms of flagella and patterns of similarities among flagellar protein sequences, including the observation that almost all of the core flagellar proteins have known homologies with non-flagellar proteins. Furthermore, several processes have been identified as playing important roles in flagellar evolution, including self-assembly of simple repeating subunits, gene duplication with subsequent divergence, recruitment of elements from other systems (‘molecular bricolage’) and recombination.
Different species of bacteria have different numbers and arrangements of flagella.
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".
Spirochetes, in contrast, have flagella arising from opposite poles of the cell, and are located within the periplasmic space as shown by breaking the outer-membrane and more recently by electron cryotomography microscopy. The rotation of the filaments 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". Even if all flagella would rotate clockwise, they likely 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 parahaemolyticus) 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 archaellum possessed by some archeae is superficially similar to the bacterial 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 or function.
Discoveries in the 1990s revealed numerous detailed differences between the archaeal and bacterial flagella. These include:
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 begun to garner scientific attention.
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 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.:63–84 For surface structures, see below.
Each of the outer 9 doublet microtubules extends a pair of dynein arms (an "inner" and an "outer" arm) to the adjacent microtubule; these 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.
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, 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.
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., the choanocytes of sponges, or 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 lifecycle 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), and 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), the gymnosperms except cycads and Ginkgo, angiosperms, pennate diatoms, some apicomplexans, some amoebozoans, in the sperm of some metazoans, and in fungi (except chytrids).
According to the place of insertion of the flagella:
According to the beating pattern:
Other terms related to the flagellar type:
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.Aeromonadales
The Aeromonadales are an order of Proteobacteria, with 10 genera in two families. The species are anaerobic. The cells are rod-shaped. Some species of this order are motile by a single polar flagellum; others are not motile.Alteromonadaceae
The Alteromonadaceae are a family of Proteobacteria. They are now one of several families in the order Alteromonadales, including Alteromonas and its closest relatives. Species of this family are mostly rod-like shaped and motile by using one polar flagellum.Alteromonadales
The Alteromonadales are an order of Proteobacteria. Although they have been treated as a single family, the Alteromonadaceae, they were divided into eight by Ivanova et al. in 2004. The cells are straight or curved rods. They are motile by the use of a single flagellum. Most of the species are marine.Ancyromonadida
Ancyromonadida or Planomonadida is a small group of biflagellated protists found in the soil and in aquatic habitats, where they feed on bacteria. Includes freshwater or marine organisms, benthic, dorsoventrally compressed and with two unequal flagellae, each emerging from a separate pocket. The apical anterior flagellum can be very thin or end in the cell membrane, while the posterior flagellum is long and is inserted ventrally or laterally. The cell membrane is supported by a thin single layer teak and the mitochondrial crests are discoidal / flat.The group's placement is doubtful, as it seems to fall outside the five supergroups of Eukarya. Cavalier-Smith considers that they constitute a basal group to Amoebozoa and Opisthokonta and places it together with other related groups in Sulcozoa. However, they appear more basal than Malawimonas, placing them in Loukouzoa, possibly as stem podiates, and depending on the placement of the root position of the Eukaryotes.Archaellum
An archaellum (plural: archaella, formerly archaeal flagellum) is a unique whip-like structure on the cell surface of many archaea. The name was proposed in 2012 following studies that showed it to be evolutionarily and structurally different from the bacterial and eukaryotic flagella. The archaellum is functionally the same – it can be rotated and is used to swim in liquid environments. The archaellum was found to be structurally similar to the type IV pilus.Attila
Attila (; fl. c. 406–453), frequently called Attila the Hun, was the ruler of the Huns from 434 until his death in March 453. He was also the leader of a tribal empire consisting of Huns, Ostrogoths, and Alans among others, in Central and Eastern Europe.
During his reign, he was one of the most feared enemies of the Western and Eastern Roman Empires. He crossed the Danube twice and plundered the Balkans, but was unable to take Constantinople. His unsuccessful campaign in Persia was followed in 441 by an invasion of the Eastern Roman (Byzantine) Empire, the success of which emboldened Attila to invade the West. He also attempted to conquer Roman Gaul (modern France), crossing the Rhine in 451 and marching as far as Aurelianum (Orléans) before being defeated at the Battle of the Catalaunian Plains.
He subsequently invaded Italy, devastating the northern provinces, but was unable to take Rome. He planned for further campaigns against the Romans, but died in 453. After Attila's death, his close adviser, Ardaric of the Gepids, led a Germanic revolt against Hunnic rule, after which the Hunnic Empire quickly collapsed.Basal body
A basal body (synonymous with basal granule, kinetosome, and in older cytological literature with blepharoplast) is a protein structure found at the base of a eukaryotic undulipodium (cilium or flagellum). It is formed from a centriole and several additional protein structures, and is, essentially, a modified centriole. The basal body serves as a nucleation site for the growth of the axoneme microtubules. Centrioles, from which basal bodies are derived, act as anchoring sites for proteins that in turn anchor microtubules, and are known as the microtubule organizing center (MTOC). These microtubules provide structure and facilitate movement of vesicles and organelles within many eukaryotic cells.Choanoflagellate
The choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of the animals. Choanoflagellates are collared flagellates having a funnel shaped collar of interconnected microvilli at the base of a flagellum. Choanoflagellates are capable of both asexual and sexual reproduction. They have a distinctive cell morphology characterized by an ovoid or spherical cell body 3–10 µm in diameter with a single apical flagellum surrounded by a collar of 30–40 microvilli (see figure). Movement of the flagellum creates water currents that can propel free-swimming choanoflagellates through the water column and trap bacteria and detritus against the collar of microvilli, where these foodstuffs are engulfed. This feeding provides a critical link within the global carbon cycle, linking trophic levels. In addition to their critical ecological roles, choanoflagellates are of particular interest to evolutionary biologists studying the origins of multicellularity in animals. As the closest living relatives of animals, choanoflagellates serve as a useful model for reconstructions of the last unicellular ancestor of animals.Chromerida
Chromerida is a phylum of unicellular alveolates, which includes photosynthetic species Chromera velia and Vitrella brassicaformis. General features of the phylum include spherical cells each with a thick cell wall, chloroplast present with chlorophyll a only (no chlorophyll b or c), and an internal developing flagellum at some lifestages.
They often live in close association with corals, and studies suggest their closest relatives is the parastic group Apicomplexa, which evolved from photosyntethic ancestors, making Chromerida the last remaining photosynthetic members of an otherwise parasitic branch within Alveolata.Carter Lab at University of Sydney has undertaken new experiments to isolate novel Chromerids, using the same methods that were used to isolate Chromera velia and Vitrella brassicaformis. These methods were agreed at the First Chromera Conference and Workshop held at the Heron Island Research Station, Queensland, Australia from November 21–25, 2011.Evolution of flagella
The evolution of flagella is of great interest to biologists because the three known varieties of flagella (eukaryotic, bacterial, and archaeal) each represent a sophisticated cellular structure that requires the interaction of many different systems.Masticophis flagellum
Masticophis flagellum is a species of nonvenomous colubrid snake, commonly referred to as the coachwhip or the whip snake, which is endemic to the United States and Mexico. Six subspecies are recognized, including the nominotypical subspecies.Opisthokont
The opisthokonts (Greek: ὀπίσθιος (opísthios) = "rear, posterior" + κοντός (kontós) = "pole" i.e. "flagellum") are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and Breviata comprise the larger clade Obazoa.Parvularcula
Parvularcula bermudensis is a marine bacterium which was identified in 2003 in the western Sargasso Sea in the Atlantic Ocean. It forms a deep branch in the Alpha Proteobacteria, distinct from the other orders.
Parvularcula isolates are Gram-negative, strictly aerobic, chemoheterotrophic, slightly motile short rods with a single flagellum. Colonies on marine agar are very small (0·3–0·8 mm in diameter), yellowish-brown and very hard. They are oxidase positive and catalase negative.Pedinellales
Pedinellales is a group of single-celled algae found in both marine environments and freshwater.These are found in both freshwater and marine environments, and most genera are sessile, attached by posterior stalks. The flagellum is at the anterior of the cell, and the tentacles surround it, often capturing small prey drawn in by its current. The colored genera are Pedinella, Apedinella, Pseudopedinella, and Mesopedinella. Several more genera have lost their chloroplasts and feed entirely by phagocytosis. These are Parapedinella, Actinomonas, and Pteridomonas.
It also appears that certain heliozoa are actually derived pedinellids. Ciliophrys alternates between a mobile flagellate stage and a heliozoan feeding stage, where the body is contracted with extended axopods all over its surface, and the flagellum is curled up into a tight figure eight. The actinophryids, Actinophrys and Actinosphaerium, exist only in a heliozoan form with no flagellum and with more elaborate bundles of microtubules supporting their axopods. Their inclusion was argued by Mikrjukov and Patterson, who coined the term actinodine to refer specifically to this extended group.
Pedinellids were classified as heliozoans by some authors. The colored pedinellids were originally treated as a family of golden algae in the order Ochromonadales, promoted to an order Pedinellales by Zimmerman in 1984. Their relationship to the silicoflagellates became apparent some time later, and Patterson defined this rankless group for the two in 1994. Moestrup treated it as the class Dictyochophyceae, previously restricted to the silicoflagellates, while Cavalier-Smith defined a new class Actinochrysophyceae for them.Per Flagellum Sanguemque, Tenebras Veneramus
Per Flagellum Sanguemque, Tenebras Veneramus (Latin for With Blood and Whip, We Worship the Dark) is the sixth full-length studio album by Gnaw Their Tongues, released on November 8, 2011 by Crucial Blast. The album saw a return to the more direct and aggressive sound of Gnaw Their Tongues' earlier work, which contrasts the more polished and lighter toned L'arrivée de la terne mort triomphante released the previous year.Sperm
Sperm is the male reproductive cell. In the types of sexual reproduction known as anisogamy and its subtype oogamy, there is a marked difference in the size of the gametes with the smaller one being termed the "male" or sperm cell. A uniflagellar sperm cell that is motile is referred to as a spermatozoon, whereas a non-motile sperm cell is referred to as a spermatium. Sperm cells cannot divide and have a limited life span, but after fusion with egg cells during fertilization, a new organism begins developing, starting as a totipotent zygote. The human sperm cell is haploid, so that its 23 chromosomes can join the 23 chromosomes of the female egg to form a diploid cell. In mammals, sperm develops in the testicles, is stored in the epididymis, and released from the penis.
The word "sperm" is derived from the Greek word (σπέρμα) sperma (meaning "seed").Syntrophobacterales
The Syntrophobacterales are an order of Proteobacteria, with two families. All genera are strictly anaerobic. Many of the family Syntrophobacteraceae are sulfate-reducing. Some species are motile by using one polar flagellum.Trypanosomatida
Trypanosomatida is a group of kinetoplastid excavates distinguished by having only a single flagellum. The name is derived from the Greek trypano (borer) and soma (body) because of the corkscrew-like motion of some trypanosomatid species. All members are exclusively parasitic, found primarily in insects. A few genera have life-cycles involving a secondary host, which may be a vertebrate, invertebrate or plant. These include several species that cause major diseases in humans.Zoospore
A zoospore is a motile asexual spore that uses a flagellum for locomotion. Also called a swarm spore, these spores are created by some protists, bacteria and fungi to propagate themselves.