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Flagellum

From Wikipedia, the free encyclopedia

A flagellum (/fldlm/; plural: flagella) is a lash-like


appendage that protrudes from the cell body of certain prokaryotic Flagellum
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.[1][2][3][4] Flagella are organelles
defined by function rather than structure. Large differences occur
between different types of flagella; the prokaryotic and eukaryotic
flagella differ greatly in protein composition, structure, and
mechanism of propulsion. However, both can be used for
swimming.

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.[5] An Structure of bacterial flagellum.
example of a eukaryotic flagellate cell is the mammalian sperm cell,
which uses its flagellum to propel itself through the female
reproductive tract.[6] Eukaryotic flagella are structurally identical to
eukaryotic cilia, although distinctions are sometimes made
according to function or length.[7] Fimbriae and pili are also thin
appendages, but have different functions and are usually smaller.

Contents
1 Types
1.1 Bacterial
1.1.1 Structure and composition
1.1.2 Motor SEM image of flagellated Chlamydomonas
1.1.3 Assembly sp. (10000)
1.1.4 Evolution
Identifiers
1.1.5 Flagella and the intelligent design debate
1.1.6 Flagellar arrangement schemes TH H1.00.01.1.01032
1.2 Archaeal FMA 67472
1.3 Eukaryotic
1.3.1 Terminology Anatomical terminology
1.3.2 Internal structure
1.3.3 Mechanism
1.3.4 Flagella vs cilia
1.3.5 Intraflagellar transport
1.3.6 Evolution and occurrence
1.3.7 Typology
2 See also
3 References
4 External links

Types
Three types of flagella have so far been distinguished: bacterial, archaeal, and eukaryotic.
The main differences among these three types are:

Bacterial flagella are helical filaments, each with a rotary motor


at its base which can turn clockwise or counterclockwise.[8][9][10]
They provide two of several kinds of bacterial motility.[11][12]
Archaeal flagella (archaella) are superficially similar to bacterial
flagella, but are different in many details and considered non-
homologous.[13][14][15] Prokaryotic flagella run in a rotary
Eukaryotic flagellathose of animal, plant, and protist cellsare movement, while eukaryotic flagella run
complex cellular projections that lash back and forth. Eukaryotic in a bending movement. The prokaryotic
flagella are classed along with eukaryotic motile cilia as flagella use a rotary motor, and the
undulipodia[16] to emphasize their distinctive wavy appendage eukaryotic flagella use a complex sliding
role in cellular function or motility. Primary cilia are immotile, filament system. Eukaryotic flagella are
and are not undulipodia; they have a structurally different 9+0 ATP driven, while prokaryote ones are
axoneme rather than the 9+2 axoneme found in both flagella and proton driven.
motile cilia undulipodia.

Bacterial

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.[17][18]

The flagellar filament is the long, helical screw that propels the
bacterium when rotated by the motor, through the hook. In most Physical model of a bacterial flagellum
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.[19]

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, bacterial flagella may have evolved from the type-three
secretion system; however, whether these pores are derived from the bacterial flagella or the bacterial secretory
system is not known for certain.

Motor

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[20]). 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.[21] The flagellum is
highly energy efficient and uses very little energy.[22] The exact mechanism for torque generation is still poorly
understood.[23] 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.[24]

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.[25]

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.[26]

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.[27]

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.[28]
In vitro, flagellar filaments assemble spontaneously in a solution containing purified flagellin as the sole
protein.[29]

Evolution

At least 10 protein components of the bacterial flagellum share homologous proteins with the type-three
secretion system (TTSS),[30] 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[31] 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,[31] but the TTSS evolving from the flagellum can be seen as 'reductive evolution', and receives
no topological support from the phylogenetic trees.[32] 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.[33]

Flagella and the intelligent design debate

Some authors have argued that flagella cannot have evolved because they can only function properly when all
proteins are in place.[34] In other words, the flagellar apparatus is "irreducibly complex". This has long been
debunked, because many proteins can be deleted or mutated and the flagellum still works, though sometimes at
reduced efficiency.[35] In addition, the composition of flagella is surprisingly diverse across bacteria, with many
proteins only found in some species, but not others.[36] 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.[37] Additional evidence for the evolution of bacterial
have been found that increase the motility of E. coli.[37] 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.[38] 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.[39]

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 bacterial 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
bacterium 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 Examples of bacterial flagella
are organized outside the cell body, helically twining about each other arrangement schemes: (A) monotrichous;
to form a thick structure (easily visible with the light microscope) called (B) lophotrichous; (C) amphitrichous;
a "fascicle". (D) peritrichous.

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". Even if all flagella
would rotate clockwise, they likely will not form a bundle, due to geometrical, as well as hydrodynamic
reasons.[40][41] 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[42]) and related proteobacteria such as Aeromonas,
In some Vibrio spp. (particularly Vibrio parahemolyticus[42]) 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.[43][44][45][46][47][48] These provide swarming motility
on surfaces or in viscous fluids.

Archaeal

The archaellum possessed by some members of domain Archea is superficially similar to the bacterial
flagellum; in the 1980s, they were thought to be homologous on the basis of gross morphology and
behavior.[49] 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.[4]

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 rotates 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, 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.[50]
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).[50]

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),[51] archaella have only recently begun to garner
scientific attention.

Eukaryotic

Terminology

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)[52] or "cilia" for both (e.g., Hlsmann, 1992;[53] Adl et al., 2012;[54] 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;[55]
Leadbeater et al., 2000).[56]

Internal structure
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 Cross section of an axoneme
500 nanometers long. Basal bodies
Eukaryotic flagella. 1axoneme, 2cell
are structurally identical to
membrane, 3IFT (IntraFlagellar centrioles. The flagellum is encased within the cell's plasma membrane,
Transport), 4Basal body, 5Cross so that the interior of the flagellum is accessible to the cell's cytoplasm.
section of flagella, 6Triplets of
microtubules of basal body 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.[57] For The "9+2" structure is visible in this
Longitudinal section through the flagella surface structures, see below. cross-section micrograph of
area in Chlamydomonas reinhardtii. In axoneme.
the cell apex is the basal body that is the Mechanism
anchoring site for a flagellum. Basal
bodies originate from and have a
Each of the outer 9 doublet microtubules extends a pair of dynein arms
substructure similar to that of centrioles,
(an "inner" and an "outer" arm) to the adjacent microtubule; these
with nine peripheral microtubule triplets
produce force through ATP hydrolysis. The flagellar axoneme also
(see structure at bottom center of image).
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
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.[58]

Evolution and occurrence

Eukaryotic flagella or cilia, probably an ancestral


characteristic,[59] 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).[60][61][62] Difference of beating pattern of flagellum and cilia

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),[63] 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,[64] and in fungi (except chytrids).

Typology

A number of terms related to flagella or cilia are used to characterize eukaryotes.[65][66][67][68][69] According to
surface structures present, flagella may be:

whiplash flagella (= smooth, acronematic flagella): without hairs, e.g., in Opisthokonta


hairy flagella (= tinsel, flimmer, pleuronematic flagella): with hairs (= mastigonemes sensu lato), divided
in:
with fine hairs (= non tubular, or simple hairs): occurs in Euglenophyceae, Dinoflagellata, some
Haptophyceae (Pavlovales)
with stiff hairs (= tubular hairs, retronemes, mastigonemes sensu stricto), divided in:
bipartite hairs: with two regions. Occurs in Cryptophyceae, Prasinophyceae, and some
Heterokonta
tripartite (= straminipilous) hairs: with three regions (a base, a tubular shaft, and one or more
terminal hairs). Occurs in most Heterokonta
stichonematic flagella: with a single row of hairs
pantonematic flagella: with two rows of hairs
acronematic: flagella with a single, terminal mastigoneme or flagellar hair (e.g., bodonids);[70] some
authors use the term as synonym of whiplash
with scales: e.g., Prasinophyceae
with spines: e.g., some brown algae
with undulating membrane: e.g., some kinetoplastids, some parabasalids
with proboscis (trunk-like protrusion of the cell): e.g., apusomonads, some bodonids[71]

According to the number of flagella, cells may be (remembering that some authors use "ciliated" instead of
"flagellated":[62][72]

uniflagellated: e.g., most Opisthokonta


biflagellated: e.g., all Dinoflagellata, the gametes of Charophyceae, of most bryophytes and of some
metazoans[73]
triflagellated: e.g., the gametes of some Foraminifera
quadriflagellated: e.g., some Prasinophyceae, Collodictyonidae
octoflagellated: e.g., some Diplomonada, some Prasinophyceae
multiflagellated: e.g., Opalinata, Ciliophora, Stephanopogon, Parabasalida, Hemimastigophora,
Caryoblastea, Multicilia, the gametes (or zoids) of Oedogoniales (Chlorophyta), some pteridophytes and
some gymnosperms

According to the place of insertion of the flagella:[74]

opisthokont: cells with flagella inserted posteriorlly, e.g., in Opisthokonta (Vischer, 1945). In
Haptophyceae, flagella are laterally to terminally inserted, but are directed posteriorly during rapid
swimming.[75]
akrokont: cells with flagella inserted apically
subakrokont: cells with flagella inserted subapically
pleurokont: cells with flagella inserted laterally

According to the beating pattern:

gliding: a flagellum that trails on the substrate[71]


heterodynamic: flagella with different beating patterns (usually with one flagellum functioning in food
capture and the other functioning in gliding, anchorage, propulsion or steering)[76]
isodynamic: flagella beating with the same patterns

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
stephanokont: cells with a crown of flagella near its anterior end, e.g., the gametes and spores of
Oedogoniales, the spores of some Bryopsidales. Term introduced by Blackman & Tansley (1902) to refer
to the Oedogoniales
akont: cells without flagella. It was also used to refer to taxonomic groups, as Aconta or Akonta: the
Zygnematophyceae and Bacillariophyceae (Oltmanns, 1904), or the Rhodophyceae (Christensen, 1962)

See also
Archaellum
Cilium
Evolution of flagella
Ciliopathy
Rotating locomotion in living systems
Undulipodium

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This article incorporates text from a publication now in the public domain: Chambers, Ephraim, ed. (1728).
"article name needed". Cyclopdia, or an Universal Dictionary of Arts and Sciences (first ed.). James and John
Knapton, et al.

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