A flagellum (; ) is a hairlike appendage that protrudes from certain
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
and animal
sperm cell
Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
s, and from a wide range of
microorganism
A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s to provide
motility
Motility is the ability of an organism to move independently, using metabolic energy.
Definitions
Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
.
Many
protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s with flagella are termed as
flagellate
A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
s.
A microorganism may have from one to many flagella. A
gram-negative bacterium
Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
''
Helicobacter pylori
''Helicobacter pylori'', previously known as ''Campylobacter pylori'', is a gram-negative, microaerophilic, spiral (helical) bacterium usually found in the stomach. Its helical shape (from which the genus name, helicobacter, derives) is though ...
'' for example uses its multiple flagella to propel itself through the
mucus lining to reach the stomach
epithelium
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...
, where it may cause a
gastric ulcer
Peptic ulcer disease (PUD) is a break in the inner lining of the stomach, the first part of the small intestine, or sometimes the lower esophagus. An ulcer in the stomach is called a gastric ulcer, while one in the first part of the intestines i ...
to develop.
In some bacteria the flagellum can also function as a sensory
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
, being sensitive to wetness outside the cell.
Across the
three domains of
Bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
,
Archaea
Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
, and
Eukaryota
Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the ...
the flagellum has a different structure, protein composition, and mechanism of propulsion but shares the same function of providing motility. The
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word means "
whip
A whip is a tool or weapon designed to strike humans or other animals to exert control through pain compliance or fear of pain. They can also be used without inflicting pain, for audiovisual cues, such as in equestrianism. They are generally e ...
" to describe its lash-like swimming motion. The flagellum in archaea is called the
archaellum
The archaellum (; formerly archaeal flagellum) is a unique structure on the cell surface of many archaea, that allows for swimming motility. The archaellum consists of a rigid helical filament that is attached to the cell membrane by a molecular ...
to note its difference from the bacterial flagellum.
Eukaryotic flagella and
cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
are identical in structure but have different lengths and functions.
Prokaryotic
A prokaryote () is a Unicellular organism, single-celled organism that lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek language, Greek wikt:πρό#Ancient Greek, πρό (, 'before') a ...
fimbriae and
pili are smaller, and thinner appendages, with different functions.
Types
![Difference Between Prokaryote and Eukaryote Flagella](https://upload.wikimedia.org/wikipedia/commons/4/4c/Difference_Between_Prokaryote_and_Eukaryote_Flagella.svg)
The three types of flagella are bacterial, archaeal, and eukaryotic.
The flagella in eukaryotes have
dynein and
microtubule
Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nm and have an inner diameter between 11 an ...
s that move with a bending mechanism. Bacteria and archaea do not have dynein or microtubules in their flagella, and they move using a rotary mechanism.
Other differences among these three types are:
*Bacterial flagella are helical filaments, each with a
rotary motor
An electric motor is an electrical machine that converts electrical energy into mechanical energy. Most electric motors operate through the interaction between the motor's magnetic field and electric current in a wire winding to generate for ...
at its base which can turn clockwise or counterclockwise.
They provide two of several kinds of
bacterial motility
Bacterial motility is the ability of bacteria to move independently using metabolic energy. Most motility mechanisms which evolved among bacteria also evolved in parallel among the archaea. Most rod-shaped bacteria can move using their own ...
.
*Archaeal flagella (
archaella) are superficially similar to bacterial flagella in that it also has a rotary motor, 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 and
motile cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
are identical in structure, but have different lengths, waveforms, and functions.
Primary cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
are immotile, and have a
structurally different 9+0 axoneme
An axoneme, also called an axial filament is the microtubule-based cytoskeletal structure that forms the core of a cilium or flagellum. Cilia and flagella are found on many cells, organisms, and microorganisms, to provide motility. The axoneme ...
rather than the
9+2 axoneme
9 (nine) is the natural number following and preceding .
Evolution of the Arabic digit
In the beginning, various Indians wrote a digit 9 similar in shape to the modern closing question mark without the bottom dot. The Kshatrapa, Andhra and ...
found in both flagella and motile cilia.
Bacterial
Structure and composition
The bacterial flagellum is made up of
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
subunits of
flagellin
Flagellin is a globular protein that arranges itself in a hollow cylinder to form the filament in a bacterial flagellum. It has a mass of about 30,000 to 60,000 daltons. Flagellin is the principal component of bacterial flagella, and is present ...
.
Its shape is a 20-
nanometer
330px, Different lengths as in respect to the molecular scale.
The nanometre (international spelling as used by the International Bureau of Weights and Measures; SI symbol: nm) or nanometer (American and British English spelling differences#-re ...
-thick hollow tube. It is
helical
Helical may refer to:
* Helix, the mathematical concept for the shape
* Helical engine, a proposed spacecraft propulsion drive
* Helical spring, a coilspring
* Helical plc, a British property company, once a maker of steel bar stock
* Helicoil
A t ...
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
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). The basal body was named by Theodor W ...
, passing through protein rings in the cell's membrane that act as bearings.
Gram-positive
In bacteriology, gram-positive bacteria are bacteria that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of cell wall.
Gram-positive bacte ...
organisms have two of these basal body rings, one in the
peptidoglycan
Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...
layer and one in the
plasma membrane
The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
.
Gram-negative
Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
organisms have four such rings: the
L ring associates with the
lipopolysaccharides
Lipopolysaccharides (LPS) are large molecules consisting of a lipid and a polysaccharide that are bacterial toxins. They are composed of an O-antigen, an outer core, and an inner core all joined by a covalent bond, and are found in the outer me ...
, the
P ring
The P ring forms part of the basal body of the bacterial appendage known as the flagellum. It is known to be embedded in the peptidoglycan cell wall. Together with the L ring, it has the function of anchoring the flagellum to the cell surface.
Re ...
associates with
peptidoglycan
Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...
layer, the M ring is embedded in the
plasma membrane
The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
, 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
''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus ''Escher ...
,
Salmonella typhimurium
''Salmonella enterica'' subsp. ''enterica'' is a subspecies of ''Salmonella enterica'', the rod-shaped, flagellated, aerobic, Gram-negative bacterium. Many of the pathogenic serovars of the ''S. enterica'' species are in this subspecies, includi ...
,
Caulobacter crescentus
''Caulobacter crescentus'' is a Gram-negative, oligotrophic bacterium widely distributed in fresh water lakes and streams. The taxon is more properly known as ''Caulobacter vibrioides'' (Henrici and Johnson 1935).
''C. crescentus'' is an importa ...
'', and ''
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 pufferfish, where it is responsible for the production of ...
'', 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
''Campylobacter jejuni'' () is a species of pathogenic bacteria, one of the most common causes of food poisoning in Europe and in the US. The vast majority of cases occur as isolated events, not as part of recognized outbreaks. Active surveillan ...
'' 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.
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
Chemiosmosis is the movement of ions across a semipermeable membrane bound structure, down their electrochemical gradient. An important example is the formation of adenosine triphosphate, adenosine triphosphate (ATP) by the movement of hydrogen ion ...
, i.e., by the flow of protons (hydrogen ions) across the bacterial cell membrane due to a
concentration gradient
Molecular diffusion, often simply called diffusion, is the thermal motion of all (liquid or gas) particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size (mass) o ...
set up by the cell's metabolism (''
Vibrio
''Vibrio'' is a genus of Gram-negative bacteria, possessing a curved-rod (comma) shape, several species of which can cause foodborne infection, usually associated with eating undercooked seafood. Being highly salt tolerant and unable to survive ...
'' species have two kinds of flagella, lateral and polar, and some are driven by a sodium
ion pump
An ion pump (also referred to as a sputter ion pump) is a type of vacuum pump which operates by sputtering a metal getter. Under ideal conditions, ion pumps are capable of reaching pressures as low as 10−11 mbar. An ion pump first ionizes ga ...
rather than a
proton pump
A proton pump is an integral membrane protein pump that builds up a proton gradient across a biological membrane
A biological membrane, biomembrane or cell membrane is a selectively permeable membrane that separates the interior of a cell f ...
). 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 (abbreviated rpm, RPM, rev/min, r/min, or with the notation min−1) is a unit of rotational speed or rotational frequency for rotating machines.
Standards
ISO 80000-3:2019 defines a unit of rotation as the dimensionl ...
, 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 molecular biology, the flagellar motor switch protein (Flig) is one of three proteins in certain bacteria coded for by the gene ''fliG''. The other two proteins are FliN coded for by ''fliN'', and FliM coded for by ''fliM''. The protein comp ...
, 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
In fluid mechanics, the Reynolds number () is a dimensionless quantity that helps predict fluid flow patterns in different situations by measuring the ratio between inertial and viscous forces. At low Reynolds numbers, flows tend to be domi ...
, 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.
[ Although according to ]
Through use of their flagella, bacteria are 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
Two-dimensional rotation can occur in two possible directions. Clockwise motion (abbreviated CW) 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 up to the top. The opposite ...
and
clockwise
Two-dimensional rotation can occur in two possible directions. Clockwise motion (abbreviated CW) 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 up to the top. The opposite ...
, respectively. The two directions of rotation are not identical (with respect to flagellum movement) and are selected by a molecular switch.
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
At least 10 protein components of the bacterial flagellum share homologous proteins with the
type three secretion system
Type three secretion system (often written Type III secretion system and abbreviated TTSS or T3SS, also called Injectisome) is a protein appendage found in several Gram-negative bacteria.
In pathogenic bacteria, the needle-like structure is u ...
(T3SS) found in many gram-negative bacteria,
hence one likely evolved from the other. Because the T3SS 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 T3SS. 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
Motility is the ability of an organism to move independently, using metabolic energy.
Definitions
Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
(mobility) support that the more mobile flagella would be selected by evolution first,
but the T3SS evolving from the flagellum can be seen as 'reductive evolution', and receives no topological support from the
phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
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.
Flagella and the intelligent design debate
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. Moreover, with many proteins unique to some number across species, diversity of bacterial flagella composition was higher than expected. 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.
Flagellar arrangements
Different species of bacteria have different numbers and arrangements of flagella,
named using the term ''tricho'', from the Greek ''trichos'' meaning ''hair''.
*Monotrichous bacteria such as ''
Vibrio cholerae
''Vibrio cholerae'' is a species of Gram-negative, facultative anaerobe and comma-shaped bacteria. The bacteria naturally live in brackish or saltwater where they attach themselves easily to the chitin-containing shells of crabs, shrimps, and oth ...
'' have a single polar flagellum.
*Amphitrichous bacteria have a single flagellum on each of two opposite ends (e.g., ''
Alcaligenes faecalis
''Alcaligenes faecalis'' is a species of Gram-negative, rod-shaped bacteria commonly found in the environment. It was originally named for its first discovery in feces, but was later found to be common in soil, water, and environments in associa ...
'')—only one flagellum operates at a time, allowing the bacterium to reverse course rapidly by switching which flagellum is active.
*Lophotrichous bacteria (''lopho'' Greek combining term meaning ''crest'' or ''tuft'')
have multiple flagella located at the same spot on the bacterial surface such as ''
Helicobacter pylori
''Helicobacter pylori'', previously known as ''Campylobacter pylori'', is a gram-negative, microaerophilic, spiral (helical) bacterium usually found in the stomach. Its helical shape (from which the genus name, helicobacter, derives) is though ...
'', 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, called the
polar organelle
A polar organelle is a structure at a specialised region of the bacterial polar membrane that is associated with the flagellar apparatus. This flagellum-associated structure can easily be distinguished from the other membrane regions in ultrathi ...
.
*Peritrichous bacteria have flagella projecting in all directions (e.g., ''E. coli'').
Counterclockwise rotation of a monotrichous polar flagellum pushes the cell forward with the flagellum trailing behind, much like a corkscrew moving inside cork. Water on the microscopic scale is highly
viscous
The viscosity of a fluid is a measure of its resistance to deformation at a given rate. For liquids, it corresponds to the informal concept of "thickness": for example, syrup has a higher viscosity than water.
Viscosity quantifies the inter ...
, unlike usual
water
Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a ...
.
Spirochete
A spirochaete () or spirochete is a member of the phylum Spirochaetota (), (synonym Spirochaetes) which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or s ...
s, in contrast, have flagella called
endoflagella
A spirochaete () or spirochete is a member of the phylum Spirochaetota (), (synonym Spirochaetes) which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or s ...
arising from opposite poles of the cell, and are located within the
periplasmic space
The periplasm is a concentrated gel-like matrix in the space between the inner cytoplasmic membrane and the bacterial outer membrane called the ''periplasmic space'' in gram-negative bacteria. Using cryo-electron microscopy it has been found tha ...
as shown by breaking the outer-membrane and also 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.
In certain large forms of ''
Selenomonas
Members of the genus ''Selenomonas'' (motile crescent-shaped bacteria in general) are referred to trivially as selenomonads. The genus ''Selenomonas'' constitutes a group of motile crescent-shaped bacteria and includes species living in the gast ...
'', 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
Fascicle or ''fasciculus'' may refer to:
Anatomy and histology
* Muscle fascicle, a bundle of skeletal muscle fibers
* Nerve fascicle, a bundle of axons (nerve fibers)
** Superior longitudinal fasciculus
*** Arcuate fasciculus
** Gracile fas ...
".
In some ''Vibrio'' spp. (particularly ''
Vibrio parahaemolyticus
''Vibrio parahaemolyticus'' (V. parahaemolyticus) is a curved, rod-shaped, Gram-negative bacterium found in the sea and in estuaries which, when ingested, may cause gastrointestinal illness in humans. ''V. parahaemolyticus'' is oxidase positiv ...
''
) and related
bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
such as ''
Aeromonas
''Aeromonas'' is a genus of Gram-negative, facultative anaerobic, rod-shaped bacteria that morphologically resemble members of the family Enterobacteriaceae. Most of the 14 described species have been associated with human diseases. The most imp ...
'', 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.
= Bundling
=
Bundling is an event that can happen in multi-flagellated cells, bundling the flagella together and causing them to rotate in a coordinated manner.
Flagella are left-handed helices, and when rotated counter-clockwise by their rotors, they can bundle and rotate together. When the rotors reverse direction, thus rotating clockwise, the flagellum unwinds from the bundle. This may cause the cell to stop its forward motion and instead start twitching in place, referred to as "tumbling". Tumbling results in a stochastic reorientation of the cell, causing it to change the direction of its forward swimming.
It is not known which stimuli drive the switch between bundling and tumbling, but the motor is highly adaptive to different signals. In the model describing
chemotaxis
Chemotaxis (from '' chemo-'' + ''taxis'') is the movement of an organism or entity in response to a chemical stimulus. Somatic cells, bacteria, and other single-cell or multicellular organisms direct their movements according to certain chemica ...
("movement on purpose") the clockwise rotation of a flagellum is suppressed by chemical compounds favorable to the cell (e.g. food). When moving in a favorable direction, the concentration of such chemical attractants increases and therefore tumbles are continually suppressed, allowing forward motion; likewise, 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.
Even if all flagella would rotate clockwise, however, they often cannot form a bundle due to geometrical and hydrodynamic reasons.
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
An undulipodium or undulopodium (a Greek word meaning "swinging foot"), or a 9+2 organelle is a motile filamentous extracellular projection of eukaryotic cells. It is basically synonymous to flagella and cilia which are differing terms for simi ...
" (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
Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow (21 October 1942 – 19 March 2021), was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford.
His research has led to disc ...
), preserving "flagella" for the bacterial structure. However, the discriminative usage of the terms "cilia" and "flagella" for eukaryotes adopted in this article (see below) is still common (e.g., Andersen et al., 1991; Leadbeater et al., 2000).
Internal structure
The core of a eukaryotic flagellum, known as the
axoneme
An axoneme, also called an axial filament is the microtubule-based cytoskeletal structure that forms the core of a cilium or flagellum. Cilia and flagella are found on many cells, organisms, and microorganisms, to provide motility. The axonem ...
is a bundle of nine fused pairs of
microtubule
Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nm and have an inner diameter between 11 an ...
s known as ''doublets'' surrounding two central single microtubules (''singlets''). This
9+2 axoneme
9 (nine) is the natural number following and preceding .
Evolution of the Arabic digit
In the beginning, various Indians wrote a digit 9 similar in shape to the modern closing question mark without the bottom dot. The Kshatrapa, Andhra and ...
is characteristic of the eukaryotic flagellum. At the base of a eukaryotic flagellum is a
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). The basal body was named by Theodor W ...
, "blepharoplast" or kinetosome, which is the
microtubule organizing center The microtubule-organizing center (MTOC) 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 spind ...
for flagellar microtubules and is about 500 nanometers long. Basal bodies are structurally identical to
centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and a ...
s. The flagellum is encased within the cell's
plasma membrane
The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
, so that the interior of the flagellum is accessible to the cell's
cytoplasm
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
.
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 that extend 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.
Mechanism
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 spoke
Radial is a geometric term of location which may refer to:
Mathematics and Direction
* Vector (geometric), a line
* Radius, adjective form of
* Radial distance, a directional coordinate in a polar coordinate system
* Radial set
* A bearing fro ...
s, 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 versus cilia
The regular beat patterns of eukaryotic cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
and flagella generate motion on a cellular level. Examples range from the propulsion of single cells such as the swimming of spermatozoa
A spermatozoon (; also spelled spermatozoön; ; ) 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. (A zygote is a single cell, with a complete set of chromosomes, ...
to the transport of fluid along a stationary layer of cells such as in the respiratory tract
The respiratory tract is the subdivision of the respiratory system involved with the process of respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa.
Air is breathed in through the nose to th ...
.[
Although eukaryotic ]cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
and flagella are ultimately the same, they are sometimes classed by their pattern of movement, a tradition from before their structures have been known. 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. Yet another traditional form of distinction is by the number of 9+2 organelles on the cell.
Intraflagellar transport
Intraflagellar transport
Intraflagellar transport (IFT) is a bidirectional motility along axoneme microtubules that is essential for the formation (ciliogenesis) and maintenance of most eukaryotic cilia and flagella. It is thought to be required to build all cilia that ass ...
, the process by which axonemal subunits, transmembrane receptors
Cell surface receptors (membrane receptors, transmembrane receptors) are receptors that are embedded in the plasma membrane of cells. They act in cell signaling by receiving (binding to) extracellular molecules. They are specialized integral m ...
, 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., zoid
In botany, a zoid or zoïd is a reproductive cell that possesses one or more flagella, and is capable of independent movement. Zoid can refer to either an asexually reproductive spore or a sexually reproductive gamete. In sexually reproductive ...
s, gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s, 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.
Diversity Flagella types
Zoospores may possess one or mo ...
s, which may be produced continually or not).
The first situation is found either in specialized cells of multicellular organisms (e.g., the choanocyte
Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a th ...
s of sponges
Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
, or the ciliated epithelia
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
of metazoan
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s), as in ciliate
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
s and many eukaryotes with a "flagellate condition" (or "monadoid level of organization An integrative level, or level of organization, is a set of phenomena emerging from pre-existing phenomena of a lower level. The levels concept is an intellectual framework for structuring reality. It arranges all entities, structures, and processe ...
", see Flagellata, an artificial group).
Flagellated lifecycle stages are found in many groups, e.g., many green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
(zoospores and male gametes), bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s (male gametes), pteridophyte
A pteridophyte is a vascular plant (with xylem and phloem) that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, ...
s (male gametes), some gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s (cycad
Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s and ''Ginkgo
''Ginkgo'' is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and is now the only living genus within ...
'', as male gametes), centric diatom
A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s (male gametes), brown algae
Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and po ...
(zoospores and gametes), oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s (assexual zoospores and gametes), hyphochytrid
Hyphochytrids are eukaryotic organisms in the group of Stramenopiles (Heterokonta).
Characteristics
They are distinguished by an anterior tinsel flagellum on their zoospores. Also they have a rhizoidal or hypha-like vegetative system (hence the ...
s (zoospores), labyrinthulomycetes
The Labyrinthulomycetes (ICBN) or Labyrinthulea (ICZN) are a class of protists that produce a network of filaments or tubes, which serve as tracks for the cells to glide along and absorb nutrients for them. The two main groups are the labyrint ...
(zoospores), some apicomplexans (gametes), some radiolarian
The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm. The elab ...
s (probably gametes), foraminiferan
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an ...
s (gametes), plasmodiophoromycetes (zoospores and gametes), myxogastrids (zoospores), metazoan
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s (male gametes), and chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. 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
Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
, some green algae (Zygnematophyceae
Zygnematophyceae (or Conjugatophyceae) is a class of green algae in the paraphylum streptophyte algae, also referred to as Charophyta, consisting of more than 4000 described species. It contains five orders: the Spirogloeales, the Serritaeniales ...
), the gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s except cycads and ''Ginkgo'', angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, pennate diatom
A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s, some apicomplexans, some amoebozoan
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported classi ...
s, in the sperm of some metazoan
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s, and in fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
(except chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
s).
Typology
A number of terms related to flagella or cilia are used to characterize eukaryotes. 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 (= mastigoneme
Mastigonemes are lateral "hairs" that attach to protistan flagella. Flimsy hairs attach to the flagella of euglenid flagellates, while stiff hairs occur in stramenopile and cryptophyte protists.Hoek, C. van den, Mann, D. G. and Jahns, H. ...
s ''sensu lato''), divided in:
**with fine hairs (= non-tubular, or simple hairs): occurs in Euglenophyceae, Dinoflagellata, some Haptophyceae
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
(Pavlovales
:
Pavlovaceae is a family of haptophyte
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used in ...
)
**with stiff hairs (= tubular hairs, retronemes, mastigonemes ''sensu stricto''), divided in:
***bipartite hairs: with two regions. Occurs in Cryptophyceae
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape ...
, Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
, and some Heterokonta
Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...
***tripartite (= straminipilous) hairs: with three regions (a base, a tubular shaft, and one or more terminal hairs). Occurs in most Heterokonta
Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...
*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); some authors use the term as synonym of whiplash
*with scales: e.g., Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
*with spines: e.g., some brown algae
Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and po ...
*with undulating membrane: e.g., some kinetoplastid
Kinetoplastida (or Kinetoplastea, as a class) is a group of flagellated protists belonging to the phylum Euglenozoa, and characterised by the presence of an organelle with a large massed DNA called kinetoplast (hence the name). The organisms are ...
s, some parabasalid
The parabasalids are a group of flagellated protists within the supergroup Excavata. Most of these eukaryotic organisms form a symbiotic relationship in animals. These include a variety of forms found in the intestines of termites and cockroaches ...
s
*with proboscis (trunk-like protrusion of the cell): e.g., apusomonads, some bodonids
According to the number of flagella, cells may be: (remembering that some authors use "ciliated" instead of "flagellated")
*uniflagellated: e.g., most Opisthokonta
*biflagellated: e.g., all Dinoflagellata, the gametes of Charophyceae
Charophyceae is a class of charophyte green algae. AlgaeBase places it in division Charophyta. Extant (living) species are placed in a single order Charales, commonly known as "stoneworts" and "brittleworts". Fossil members of the class may be ...
, of most bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s and of some metazoan
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s
*triflagellated: e.g., the gametes of some Foraminifera
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
*quadriflagellated: e.g., some Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
, Collodictyonidae
*octoflagellated: e.g., some Diplomonada, some Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
*multiflagellated: e.g., Opalinata
Opalinata is a superclass of non-phagotrophic heterokonts that unites the classes Opalinea and Blastocystea, and is the sister group to Opalomonadea. Description
When Opalinata was first erected as a taxon in 1926, it was placed as the sole class ...
, Ciliophora
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different ...
, ''Stephanopogon
''Stephanopogon'' is a genus of flagellated marine protist that superficially resembles a ciliate.
Characteristics
''Stephanopogon'' closely resembles certain ciliates and was originally classified with them (, but is now considered related to h ...
'', Parabasalid
The parabasalids are a group of flagellated protists within the supergroup Excavata. Most of these eukaryotic organisms form a symbiotic relationship in animals. These include a variety of forms found in the intestines of termites and cockroaches ...
a, Hemimastigophora
Hemimastigophora is a group of single-celled eukaryotic organisms with a single family, Spironemidae, first identified in 1988. Over the next 30 years, different authors proposed placing these organisms in various branches of the eukaryotes ...
, Caryoblastea, '' Multicilia'', the gametes (or zoid
In botany, a zoid or zoïd is a reproductive cell that possesses one or more flagella, and is capable of independent movement. Zoid can refer to either an asexually reproductive spore or a sexually reproductive gamete. In sexually reproductive ...
s) of Oedogoniales (Chlorophyta
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to ...
), some pteridophyte
A pteridophyte is a vascular plant (with xylem and phloem) that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, ...
s and some gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s
According to the place of insertion of the flagella:
*opisthokont: cells with flagella inserted posteriorly, e.g., in Opisthokonta (Vischer, 1945). In Haptophyceae
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
, flagella are laterally to terminally inserted, but are directed posteriorly during rapid swimming.
*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
*heterodynamic: flagella with different beating patterns (usually with one flagellum functioning in food capture and the other functioning in gliding, anchorage, propulsion or "steering")
*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
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to ...
*anisokont: cells with flagella of unequal length, e.g., some Euglenophyceae and Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
*heterokont: term introduced by Luther (1899) to refer to the Xanthophyceae
Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamento ...
, 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
Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...
*stephanokont: cells with a crown of flagella near its anterior end, e.g., the gametes and spores of Oedogoniales, the spores of some Bryopsidales
Bryopsidales is an order of green algae, in the class Ulvophyceae.
Characteristics
The thallus is filamentous and much branched and may be packed into a mass. It is coenocytic, having multi-nucleate cells consisting of cytoplasm contained withi ...
. 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
Zygnematophyceae (or Conjugatophyceae) is a class of green algae in the paraphylum streptophyte algae, also referred to as Charophyta, consisting of more than 4000 described species. It contains five orders: the Spirogloeales, the Serritaeniales ...
and Bacillariophyceae
Bacillariophyceae is a group of pennate diatoms with a raphe (raphids). According to Ruggiero et al., 2015, the diatoms are treated as follows. This treatment largely reflects that used by Algaebase as at 2015, and is also reflected in the current ...
(Oltmanns, 1904), or the Rhodophyceae
Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
(Christensen, 1962)
Archaeal
The archaellum
The archaellum (; formerly archaeal flagellum) is a unique structure on the cell surface of many archaea, that allows for swimming motility. The archaellum consists of a rigid helical filament that is attached to the cell membrane by a molecular ...
possessed by some species of Archaea
Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
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 proteins (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 glycan
The terms glycans and polysaccharides are defined by IUPAC as synonyms meaning "compounds consisting of a large number of monosaccharides linked glycosidically". However, in practice the term glycan may also be used to refer to the carbohydrate p ...
s which are necessary for proper assembly or function.
Discoveries in the 1990s revealed numerous detailed differences between the archaeal and bacterial flagella. These include:
*Bacterial flagella rotation is powered by the proton motive force
Chemiosmosis is the movement of ions across a semipermeable membrane bound structure, down their electrochemical gradient. An important example is the formation of adenosine triphosphate (ATP) by the movement of hydrogen ions (H+) across a membra ...
– a flow of H+ ions or occasionally by the sodium-motive force
Members of the Solute:Sodium Symporter (SSS) FamilyTC# 2.A.21 catalyze solute:Na+ symport. The SSS family is within the APC Superfamily. The solutes transported may be sugars, amino acids, organo cations such as choline, nucleosides, inositols, v ...
– a flow of Na+ ions; archaeal flagella rotation is powered by ATP.
*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.
*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
A pilus (Latin for 'hair'; plural: ''pili'') is a hair-like appendage found on the surface of many bacteria and archaea. The terms ''pilus'' and '' fimbria'' (Latin for 'fringe'; plural: ''fimbriae'') can be used interchangeably, although some r ...
, 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
Analogy (from Greek ''analogia'', "proportion", from ''ana-'' "upon, according to" lso "against", "anew"+ ''logos'' "ratio" lso "word, speech, reckoning" is a cognitive process of transferring information or meaning from a particular subject ( ...
, or convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
, rather than homology
Homology may refer to:
Sciences
Biology
*Homology (biology), any characteristic of biological organisms that is derived from a common ancestor
* Sequence homology, biological homology between DNA, RNA, or protein sequences
*Homologous chrom ...
. 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.
Additional images
File:EMpylori.jpg, Multiple flagella in lophotrichous arrrangement on surface of ''Helicobacter pylori''
File:Physical model of a bacterial flagellum.jpg, Physical model of a bacterial flagellum
See also
*Ciliopathy
A ciliopathy is any genetic disorder that affects the cellular cilia or the cilia anchoring structures, the basal bodies, or ciliary function. Primary cilia are important in guiding the process of development, so abnormal ciliary function while ...
*RpoF
The gene ''rpoF'' (RNA polymerase, flagellum F) encodes the sigma factor ''sigma-28'' (σ28, or RpoF), a protein in ''Escherichia coli'' and other species of bacteria. Depending on the bacterial species, this gene may be referred to as ''sigD'' o ...
References
Further reading
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External links
Cell Image Library - Flagella
{{Authority control
Cell movement
Organelles
Protein complexes
Bacteria