Flagellum
Overview
 
A flagellum is a tail-like projection that protrudes from the cell body of certain prokaryotic and eukaryotic cells, and plays the dual role of locomotion
Animal locomotion
Animal locomotion, which is the act of self-propulsion by an animal, has many manifestations, including running, swimming, jumping and flying. Animals move for a variety of reasons, such as to find food, a mate, or a suitable microhabitat, and to escape predators...

 and sense organ, being sensitive to chemicals and temperatures outside the cell. There are some notable differences between prokaryotic and eukaryotic flagella, such as protein composition, structure, and mechanism of propulsion. An example of a flagellated bacterium is the ulcer-causing Helicobacter pylori
Helicobacter pylori
Helicobacter pylori , previously named Campylobacter pyloridis, is a Gram-negative, microaerophilic bacterium found in the stomach. It was identified in 1982 by Barry Marshall and Robin Warren, who found that it was present in patients with chronic gastritis and gastric ulcers, conditions that were...

, which uses multiple flagella to propel itself through the mucus lining to reach the stomach epithelium
Epithelium
Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective...

.

An example of a eukaryotic flagellated cell is the sperm
Spermatozoon
A spermatozoon is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote...

 cell, which uses its flagellum to propel itself through the female reproductive tract.
Encyclopedia
A flagellum is a tail-like projection that protrudes from the cell body of certain prokaryotic and eukaryotic cells, and plays the dual role of locomotion
Animal locomotion
Animal locomotion, which is the act of self-propulsion by an animal, has many manifestations, including running, swimming, jumping and flying. Animals move for a variety of reasons, such as to find food, a mate, or a suitable microhabitat, and to escape predators...

 and sense organ, being sensitive to chemicals and temperatures outside the cell. There are some notable differences between prokaryotic and eukaryotic flagella, such as protein composition, structure, and mechanism of propulsion. An example of a flagellated bacterium is the ulcer-causing Helicobacter pylori
Helicobacter pylori
Helicobacter pylori , previously named Campylobacter pyloridis, is a Gram-negative, microaerophilic bacterium found in the stomach. It was identified in 1982 by Barry Marshall and Robin Warren, who found that it was present in patients with chronic gastritis and gastric ulcers, conditions that were...

, which uses multiple flagella to propel itself through the mucus lining to reach the stomach epithelium
Epithelium
Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective...

.

An example of a eukaryotic flagellated cell is the sperm
Spermatozoon
A spermatozoon is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote...

 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. The word flagellum is the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 word for whip
Whip
A whip is a tool traditionally used by humans to exert control over animals or other people, through pain compliance or fear of pain, although in some activities whips can be used without use of pain, such as an additional pressure aid in dressage...

.

Types

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 that rotate like screws. They provide two of several kinds of bacterial motility.
  • Archaea
    Archaea
    The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

    l flagella are superficially similar to bacterial flagella, but are different in many details and considered non-homologous
    Homology (biology)
    Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...

    .
  • Eukaryotic
    Eukaryote
    A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

     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
    Motility
    Motility is a biological term which refers to the ability to move spontaneously and actively, consuming energy in the process. Most animals are motile but the term applies to single-celled and simple multicellular organisms, as well as to some mechanisms of fluid flow in multicellular organs, in...

    . 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 undulopodia.

Bacterial

The bacterial flagellum is made up of the protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 flagellin
Flagellin
Flagellin is a protein that arranges itself in a hollow cylinder to form the filament in bacterial flagellum. It has a mass of about 30,000 to 60,000 daltons...

. Its shape is a 20 nanometer-thick hollow tube. It is helical
Helix
A helix is a type of smooth space curve, i.e. a curve in three-dimensional space. It has the property that the tangent line at any point makes a constant angle with a fixed line called the axis. Examples of helixes are coil springs and the handrails of spiral staircases. A "filled-in" helix – for...

 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
Basal body
A basal body is an organelle formed from a centriole, and a short cylindrical array of microtubules. It is found at the base of a eukaryotic undulipodium and serves as a nucleation site for the growth of the axoneme microtubules...

, passing through protein rings in the cell's membrane that act as bearings. Gram-positive
Gram-positive
Gram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...

 organisms have 2 of these basal body rings, one in the peptidoglycan
Peptidoglycan
Peptidoglycan, also known as murein, is a polymer consisting of sugars and amino acids that forms a mesh-like layer outside the plasma membrane of bacteria , forming the cell wall. The sugar component consists of alternating residues of β- linked N-acetylglucosamine and N-acetylmuramic acid...

 layer and one in the plasma membrane. Gram-negative
Gram-negative
Gram-negative bacteria are bacteria that do not retain crystal violet dye in the Gram staining protocol. In a Gram stain test, a counterstain is added after the crystal violet, coloring all Gram-negative bacteria with a red or pink color...

 organisms have 4 such rings: the L ring
L ring
The L-ring of the bacterial flagellum is the ring in the lipid outer cell membrane through which the axial filament passes....

 associates with the lipopolysaccharides, the P ring
P ring
The P ring is the part of the bacterial flagellum, which is known to be embedded in the peptidoglycan cell wall....

 associates with peptidoglycan
Peptidoglycan
Peptidoglycan, also known as murein, is a polymer consisting of sugars and amino acids that forms a mesh-like layer outside the plasma membrane of bacteria , forming the cell wall. The sugar component consists of alternating residues of β- linked N-acetylglucosamine and N-acetylmuramic acid...

 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 bacterial flagellum is driven by a rotary engine (the Mot complex) made up of protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

, located at the flagellum's anchor point on the inner cell membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

. The engine is powered by proton motive force, i.e., by the flow of proton
Proton
The proton is a subatomic particle with the symbol or and a positive electric charge of 1 elementary charge. One or more protons are present in the nucleus of each atom, along with neutrons. The number of protons in each atom is its atomic number....

s (hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 ion
Ion
An ion is an atom or molecule in which the total number of electrons is not equal to the total number of protons, giving it a net positive or negative electrical charge. The name was given by physicist Michael Faraday for the substances that allow a current to pass between electrodes in a...

s) across the bacterial cell membrane due to a concentration gradient set up by the cell's metabolism (in Vibrio
Vibrio
Vibrio is a genus of Gram-negative bacteria possessing a curved rod shape, several species of which can cause foodborne infection, usually associated with eating undercooked seafood. Typically found in saltwater, Vibrio are facultative anaerobes that test positive for oxidase and do not form...

species there are two kinds of flagella, lateral and polar, and some are driven by a sodium
Sodium
Sodium is a chemical element with the symbol Na and atomic number 11. It is a soft, silvery-white, highly reactive metal and is a member of the alkali metals; its only stable isotope is 23Na. It is an abundant element that exists in numerous minerals, most commonly as sodium chloride...

 ion pump rather than a proton pump
Proton pump
A proton pump is an integral membrane protein that is capable of moving protons across a cell membrane, mitochondrion, or other organelle. Mechanisms are based on conformational changes of the protein structure or on the Q cycle.-Function:...

). 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
Revolutions per minute
Revolutions per minute is a measure of the frequency of a rotation. It annotates the number of full rotations completed in one minute around a fixed axis...

, 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 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.

Flagella do not rotate at a constant speed but instead can increase or decrease their rotational speed in relation to the strength of the proton motive force. Flagellar rotation can move bacteria through liquid media at speeds of up to 60 cell lengths/second (sec). Although this is only about 0.00017 km/h (0.000105633102764853 mph), when comparing this speed with that of higher organisms in terms of number of lengths moved per second, it is extremely fast. By comparison, the cheetah
Cheetah
The cheetah is a large-sized feline inhabiting most of Africa and parts of the Middle East. The cheetah is the only extant member of the genus Acinonyx, most notable for modifications in the species' paws...

, the fastest land animal, can sprint at 110 km/h (68.4 mph), which is approximately 25 body lengths/sec.

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.

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
Escherichia coli
Escherichia coli is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms . Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls...

, Salmonella typhimurium, Caulobacter crescentus
Caulobacter crescentus
Caulobacter crescentus is a Gram-negative, oligotrophic bacterium widely distributed in fresh water lakes and streams.Caulobacter is an important model organism for studying the regulation of the cell cycle, asymmetric cell division, and cellular differentiation. Caulobacter daughter cells have...

, and Vibrio alginolyticus
Vibrio alginolyticus
Vibrio alginolyticus is a Gram-negative marine bacterium. It is medically important since it causes otitis and wound infection. It is also present in the bodies of animals such as Puffer fish, where it is responsible for the production of the potent neurotoxin, tetrodotoxin....

, 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
Campylobacter jejuni
Campylobacter jejuni is a species of curved, helical-shaped, non-spore forming, Gram-negative, microaerophilic bacteria commonly found in animal feces. It is one of the most common causes of human gastroenteritis in the world. Food poisoning caused by Campylobacter species can be severely...

, 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
Type three secretion system
Type three secretion system is a protein appendage found in several Gram-negative bacteria....

; however, it is not known for certain whether these pores are derived from the bacterial flagella or the bacterial secretory system.

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
Biased random walk (biochemistry)
In cell biology, a biased random walk enables bacteria to search for food and flee from harm. Bacteria propel themselves with the aid of flagella in a process called chemotaxis, and a typical bacteria trajectory has many characteristics of a random walk. They move forward for a certain distance,...

, with 'runs' and 'tumbles' brought about by rotating the flagellum counter-clockwise and clockwise
Clockwise
Circular motion can occur in two possible directions. A clockwise motion is one that proceeds in the same direction as a clock's hands: from the top to the right, then down and then to the left, and back to the top...

 respectively.

Flagella arrangement schemes

Different species of bacteria have different numbers and arrangements of flagella. Monotrichous bacteria have a single flagellum (e.g., Vibrio cholerae
Vibrio cholerae
Vibrio cholerae is a Gram-negative, comma-shaped bacterium. Some strains of V. cholerae cause the disease cholera. V. cholerae is facultatively anaerobic and has a flagella at one cell pole. V...

). 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 membrane
Polar membrane
In the scientific literature polar membrane has acquired two separate meanings.1) A lipid biomembrane expressing polarity in the electrical sense...

. 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 some bacteria, such as the larger forms of Selenomonas
Selenomonad
The genus Selenomonas constitutes a group of motile crescent-shaped bacteria within the Veillonellaceae family and includes species living in the gastrointestinal tracts of animals, in particular, the Ruminants....

, the individual flagella are organized outside the cell body, helically twining about each other to form a thick structure called a "fascicle".
Other bacteria, such as Spirochetes, have a specialized type of flagellum called an "axial filament" that is located in the periplasmic space
Periplasmic space
The periplasmic space or periplasm is a space between the peptidoglycan cell wall and inner membrane of Gram-negative bacteria or the equivalent space outside the inner membrane of Gram-positive bacteria. It may constitute up to 40% of the total cell volume in Gram-negative species, and is...

, the rotation of which causes the entire bacterium to move forward in a corkscrew-like motion.

Counterclockwise rotation of monotrichous polar flagella pushes the cell forward with the flagella trailing behind, much like a corkscrew moving inside cork. Indeed water in the microscopic scale is highly viscous, very different from our daily experience of water. The 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 hydrodynamical 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
Vibrio
Vibrio is a genus of Gram-negative bacteria possessing a curved rod shape, several species of which can cause foodborne infection, usually associated with eating undercooked seafood. Typically found in saltwater, Vibrio are facultative anaerobes that test positive for oxidase and do not form...

spp. (particularly Vibrio parahemolyticus) and related proteobacteria
Proteobacteria
The Proteobacteria are a major group of bacteria. They include a wide variety of pathogens, such as Escherichia, Salmonella, Vibrio, Helicobacter, and many other notable genera....

 such as Aeromonas
Aeromonas
Aeromonas is a gram-negative, facultative anaerobic rod that morphologically resembles members of the family Enterobacteriaceae. Fourteen species of Aeromonas have been described, most of which have been associated with human diseases. The most important pathogens are A. hydrophila, A. caviae, and...

, 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.

Archaeal

The archaeal flagellum 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 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 pilin
Pilin
Pilin refers to a class of fibrous proteins that are found in pilus structures in bacteria. Bacterial pili are used in the exchange of genetic material during bacterial conjugation, and a short pilus called a fimbrium is used as a cell adhesion mechanism. Although not all bacteria have pili or...

s, the component flagellins are made with class 3 signal peptides and they are processed by a type IV prepilin peptidase-like enzyme. The archaeal flagellins 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+
    Hydrogen
    Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

     ion
    Ion
    An ion is an atom or molecule in which the total number of electrons is not equal to the total number of protons, giving it a net positive or negative electrical charge. The name was given by physicist Michael Faraday for the substances that allow a current to pass between electrodes in a...

    s (or occasionally Na+
    Sodium
    Sodium is a chemical element with the symbol Na and atomic number 11. It is a soft, silvery-white, highly reactive metal and is a member of the alkali metals; its only stable isotope is 23Na. It is an abundant element that exists in numerous minerals, most commonly as sodium chloride...

     ions); archaeal flagella are almost certainly powered by ATP
    Adenosine triphosphate
    Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

    . The torque
    Torque
    Torque, moment or moment of force , is the tendency of a force to rotate an object about an axis, fulcrum, or pivot. Just as a force is a push or a pull, a torque can be thought of as a twist....

    -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 archaeal flagella, 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 archaeal flagellum is too thin to allow this.
  • Many components of bacterial flagella share sequence similarity to components of the type III secretion systems
    Type three secretion system
    Type three secretion system is a protein appendage found in several Gram-negative bacteria....

    , but the components of bacterial and archaeal flagella share no sequence similarity. Instead, some components of archaeal flagella 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 and archaeal flagella could be a classic case of biological analogy
Analogy
Analogy is a cognitive process of transferring information or meaning from a particular subject to another particular subject , and a linguistic expression corresponding to such a process...

, or convergent evolution
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...

, rather than homology
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...

. However, in comparison to the decades of well-publicized study of bacterial flagella (e.g. by Berg), archaeal flagella have only recently begun to get serious scientific attention. Therefore, many assume erroneously that there is only one basic kind of prokaryotic flagellum, and that archaeal flagella are homologous to it. For example, Cavalier-Smith (2002) is aware of the differences between archaeal and bacterial flagellins, but retains the misconception that the basal bodies are homologous.

Eukaryotic

Along with cilia, flagella make up a group of organelles known as undulipodia.

Structure

A eukaryotic flagellum is a bundle of nine fused pairs of microtubule
Microtubule
Microtubules are a component of the cytoskeleton. These rope-like polymers of tubulin can grow as long as 25 micrometers and are highly dynamic. The outer diameter of microtubule is about 25 nm. Microtubules are important for maintaining cell structure, providing platforms for intracellular...

 doublets surrounding two central single microtubules. The so-called "9+2" structure is characteristic of the core of the eukaryotic flagellum called an axoneme
Axoneme
Numerous eukaryotic cells carry whip-like appendages whose inner core consists of a cytoskeletal structure called the axoneme....

. At the base of a eukaryotic flagellum is a basal body
Basal body
A basal body is an organelle formed from a centriole, and a short cylindrical array of microtubules. It is found at the base of a eukaryotic undulipodium and serves as a nucleation site for the growth of the axoneme microtubules...

, "blepharoplast" or kinetosome, which is the microtubule organizing center
Microtubule organizing center
The microtubule-organizing center is a structure found in eukaryotic cells from which microtubules emerge. MTOCs have two main functions: the organization of eukaryotic flagella and cilia and the organization of the mitotic and meiotic spindle apparatus, which separate the chromosomes during cell...

 (MTOC) for flagellar microtubules and is about 500 nanometers long. Basal bodies are structurally identical to centriole
Centriole
A Centriole is a barrel-shaped cell structure found in most animal eukaryotic cells, though it is absent in higher plants and most fungi. The walls of each centriole are usually composed of nine triplets of microtubules...

s. The flagellum is encased within the cell's plasma membrane, so that the interior of the flagellum is accessible to the cell's cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...

.

Mechanism

Each of the outer 9 doublet microtubules extends a pair of dynein
Dynein
Dynein is a motor protein in cells which converts the chemical energy contained in ATP into the mechanical energy of movement. Dynein transports various cellular cargo by "walking" along cytoskeletal microtubules towards the minus-end of the microtubule, which is usually oriented towards the cell...

 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
Hydrolysis
Hydrolysis is a chemical reaction during which molecules of water are split into hydrogen cations and hydroxide anions in the process of a chemical mechanism. It is the type of reaction that is used to break down certain polymers, especially those made by condensation polymerization...

. The flagellar axoneme also contains radial spoke
Radial spoke
The radial spoke is a multi-unit protein structure found in the axonemes of eukaryotic cilia and flagella. Although experiments have determined the importance of the radial spoke in the proper function of these organelles, its structure and mode of action remain poorly understood.- Cellular...

s, polypeptide complexes extending from each of the outer 9 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 3D motion with a power and recovery stroke.

Intraflagellar Transport

Intraflagellar transport
Intraflagellar transport
Intraflagellar transport or IFT is a bidirectional motility along axonemal microtubules that is essential for the formation and maintenance of most eukaryotic cilia and flagella. It is thought to be required to build all cilia that assemble within a membrane projection from the cell surface...

 (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.

For information on biologists' ideas about how the various flagella may have evolved, see evolution of flagella
Evolution of flagella
The evolution of flagella is of great interest to biologists because the three known varieties of flagella each represent a sophisticated cellular structure that requires the interaction of many different systems....

.

See also

  • Cilia
  • Evolution of flagella
    Evolution of flagella
    The evolution of flagella is of great interest to biologists because the three known varieties of flagella each represent a sophisticated cellular structure that requires the interaction of many different systems....

  • Genetic ciliopathy
    Ciliopathy
    A ciliopathy is a genetic disorder of the cellular cilia or the cilia anchoring structures, the basal bodies, or of ciliary function.Although ciliopathies are usually considered to involve proteins that localize to the primary cilia or centrosomes, it is possible for ciliopathies to be associated...

  • Rotating locomotion in living systems
  • Undulipodia
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