An axoneme, also called an axial filament is the
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 ...
-based
cytoskeletal
The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compo ...
structure that forms the core of a
cilium
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 ...
or
flagellum
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have f ...
.
Cilia and flagella are found on many
cells,
organism
In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
s, and
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. The axoneme serves as the "skeleton" of these
organelles
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 ...
, both giving support to the structure and, in some cases, the ability to bend. Though distinctions of function and length may be made between cilia and flagella, the internal structure of the axoneme is common to both.
Structure
Inside a cilium and a flagellum is a
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 ...
-based
cytoskeleton
The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...
called the axoneme. The axoneme of a
primary cilium
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 ...
typically has a ring of nine outer microtubule doublets (called a 9+0 axoneme), and the axoneme of a
motile cilium
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 ...
has two central microtubules in addition to the nine outer doublets (called a 9+2 axoneme). The axonemal cytoskeleton acts as a scaffolding for various
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 ...
complexes and provides binding sites for
molecular motor
Molecular motors are natural (biological) or artificial molecular machines that are the essential agents of movement in living organisms. In general terms, a motor is a device that consumes energy in one form and converts it into motion or mech ...
proteins such as
kinesin-2, that help carry proteins up and down the microtubules.
[
]
Primary cilia
The axoneme structure in non-motile
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 ...
is of an outer nine microtubule doublets with no central microtubule singlets, and no
dynein arms
Dyneins are a family of cytoskeletal motor proteins that move along microtubules in cells. They convert the chemical energy stored in ATP to mechanical work. Dynein transports various cellular cargos, provides forces and displacements important ...
on the outer doublets. This arrangement is known as the 9+0 axoneme. Primary cilia appear to serve sensory functions.
Motile cilia
The building-block of the axoneme is the
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 ...
; each axoneme is composed of several microtubules aligned in a characteristic pattern known as the 9+2 axoneme as shown in the image at right. Nine sets of doublet microtubules (a specialized structure consisting of two linked microtubules) form a ring around a central pair of single microtubules.
Besides the microtubules, the axoneme contains many
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 ...
s and protein complexes necessary for its function. The
dynein arms, for example, are motor complexes that produce the force needed for bending. Each dynein arm is anchored to a doublet microtubule; by "walking" along an adjacent microtubule, the
dynein motors can cause the microtubules to slide against each other. When this is carried out in a synchronized fashion, with the microtubules on one side of the axoneme being pulled 'down' and those on the other side pulled 'up,' the axoneme as a whole can bend back and forth. This process is responsible for ciliary/flagellar beating, as in the well-known example of the
human sperm
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, ...
.
The
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 ...
is another protein complex of the axoneme. Thought to be important in regulating the motion of the axoneme, this "T"-shape complex projects from each set of outer doublets toward the central microtubules. The inter-doublet connections between adjacent microtubule pairs are termed
nexin linkages.
History of discovery
The first investigation of
sperm
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 ...
flagellar morphology was begun in 1888, by German cytologist Ballowitz, who observed using light microscopy and mordant stains that a rooster sperm flagellum could be splayed into as many as 11, longitudinal fibrils. About 60 years later, Grigg and Hodge in 1949 and a year later Manton and Clarke observed these 11 fibers in splayed flagella by electron microscopy (EM); these investigators proposed that two thinner fibers were surrounded by nine thicker outer fibers. In 1952, using advancements in fixation, embedding, and ultramicrotomy, Fawcett and
Porter
Porter may refer to:
Companies
* Porter Airlines, Canadian regional airline based in Toronto
* Porter Chemical Company, a defunct U.S. toy manufacturer of chemistry sets
* Porter Motor Company, defunct U.S. car manufacturer
* H.K. Porter, Inc., ...
proved by EM thin sections that the core of epithelial cilia within the ciliary membrane consisted of nine doublet microtubules surrounding two central, singlet microtubules (i.e., the “central pair microtubule apparatus”), and hence the term, the “9 + 2” axoneme. Because of the high degree of evolutionary conservation between cilia and flagella from most species, our understanding of sperm flagella has been aided by studies of both organelles and from species ranging from protists to mammals. Cilia are typically short (5–10 μm) and beat in an oar-like fashion with an effective stroke followed by a recovery stroke. Flagella beat with a snake-like motion and are typically longer (generally 50–150 μm, but ranging from 12 μm to several mm in some species), with flagellar length in the protist ''Chlamydomonas'' being regulated by several genes encoding kinases. It was recognized first by Manton and Clarke that the 9 + 2 axoneme was possibly ubiquitous among species, and indeed, the nine doublet microtubules are evolutionary conserved structures that evolved in early eukaryotes nearly a billion years ago; however, there is wide variation among species with regard to the detailed structure of sperm flagella and their accessory structures. Axonemal doublet microtubules assemble from the ends of nine centriolar/basal body triplet microtubules, whose ninefold symmetry and clockwise pinwheel pattern (looking from inside the cell to the flagellar tip) is organized by the conserved protein of the SAS6 gene, and which is introduced into some eggs to establish the first mitotic spindle. The nine doublet microtubules are then connected around the axoneme by nexin links. Currently, the molecular structure of the axoneme is known to an extraordinary resolution of <4 nm through the use of cryo-electron tomography, as initially pioneered by Nicastro. Sperm flagellar (and ciliary) motility has been effectively analyzed in simple systems (e.g., protist flagella and sea urchin sperm), whose flagella contain several hundred polypeptides by proteomic analysis.
[ ]
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Creative Commons Attribution 4.0 International License
Clinical significance
Mutations or defects in primary cilia have been found to play a role in human diseases. These
ciliopathies include
polycystic kidney disease
Polycystic kidney disease (PKD or PCKD, also known as polycystic kidney syndrome) is a genetic disorder in which the renal tubules become structurally abnormal, resulting in the development and growth of multiple cysts within the kidney. These cy ...
(PKD),
retinitis pigmentosa
Retinitis pigmentosa (RP) is a genetic disorder of the eyes that causes loss of vision. Symptoms include trouble seeing at night and decreasing peripheral vision (side and upper or lower visual field). As peripheral vision worsens, people may ...
,
Bardet–Biedl syndrome
Bardet–Biedl syndrome (BBS) is a ciliopathic human genetic disorder that produces many effects and affects many body systems. It is characterized by rod/cone dystrophy, polydactyly, central obesity, hypogonadism, and kidney dysfunction in so ...
, and other developmental defects.
References
Further reading
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Organelles