cell biology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...

a centriole is a cylindrical

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 ...

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 are only present in the male gametes of

charophytes Charophyta () is a group of freshwater green algae, called charophytes (), sometimes treated as a phylum, division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged within Charophyta, possibly fro ...

, bryophytes, seedless vascular plants, cycads, and '' Ginkgo''. A bound pair of centrioles, surrounded by a highly ordered mass of dense material, called the pericentriolar material (PCM), makes up a structure called a centrosome. Centrioles are typically made up of nine sets of short microtubule triplets, arranged in a cylinder. Deviations from this structure include crabs and '' Drosophila melanogaster'' embryos, with nine doublets, and ''

Caenorhabditis elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (ro ...

'' sperm cells and early embryos, with nine singlets. Additional proteins include centrin,

cenexin Outer dense fiber protein 2, also known as cenexin, is a protein that in humans is encoded by the ''ODF2'' gene. The outer dense fibers are cytoskeletal structures that surround the axoneme in the middle piece and principal piece of the sper ...

and tektin. The main function of centrioles is to produce

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 ...

during interphase and the aster and the spindle during cell division.

History

The centrosome was discovered jointly by Walther Flemming in 1875 Bloodgood RA. From central to rudimentary to primary: the history of an underappreciated organelle whose time has come. The primary cilium. Methods Cell Biol. 2009;94:3-52. doi: 10.1016/S0091-679X(08)94001-2. Epub 2009 Dec 23. PMID 20362083. and

Edouard Van Beneden Édouard Joseph Louis Marie Van Beneden (5 March 1846 in Leuven – 28 April 1910 in Liège), son of Pierre-Joseph Van Beneden, was a Belgian embryologist, cytologist and marine biologist. He was professor of zoology at the University of Liège ...

in 1876.

made the first observation of centrosomes as composed of two orthogonal centrioles in 1883. Theodor Boveri introduced the term "centrosome" in 1888 and the term "centriole" in 1895. The basal body was named by Theodor Wilhelm Engelmann in 1880. The pattern of centriole duplication was first worked out independently by

Étienne de Harven Etienne de Harven (1928-2019) was a Belgian-born pathologist and electron microscopist. Born in Brussels, he did most of his work in New York City, Paris and Toronto. He did pioneering research on viruses, mostly related to murine leukemia. He ...

and

Joseph G. Gall Joseph Grafton Gall (born April 14, 1928) is an American cell biologist who is noted for studies revealing the details of chromosome structure and function. Gall's studies were greatly facilitated by his knowledge of many different organisms bec ...

c. 1950.

Role in cell division

Centrioles are involved in the organization of the mitotic spindle and in the completion of cytokinesis. Centrioles were previously thought to be required for the formation of a mitotic spindle in animal cells. However, more recent experiments have demonstrated that cells whose centrioles have been removed via laser ablation can still progress through the G₁ stage of interphase before centrioles can be synthesized later in a de novo fashion. Additionally, mutant flies lacking centrioles develop normally, although the adult flies' cells lack

flagella 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 ...

and

and as a result, they die shortly after birth. The centrioles can self replicate during cell division.

Cellular organization

Centrioles are a very important part of centrosomes, which are involved in organizing microtubules in the cytoplasm. The position of the centriole determines the position of the nucleus and plays a crucial role in the spatial arrangement of the cell. Blausen 0214 Centrioles

Fertility

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 ...

centrioles are important for 2 functions: (1) to form the sperm flagellum and sperm movement and (2) for the development of the embryo after fertilization. The

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 ...

supplies the centriole that creates the centrosome and microtubule system of the zygote.

Ciliogenesis

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 and ciliates, the position of the flagellum or cilium is determined by the mother centriole, which becomes the basal body. An inability of cells to use centrioles to make functional flagella and cilia has been linked to a number of genetic and developmental diseases. In particular, the inability of centrioles to properly migrate prior to ciliary assembly has recently been linked to Meckel–Gruber syndrome.

Animal development

Spindle centriole - embryonic brain mouse - TEM

Proper orientation of cilia via centriole positioning toward the posterior of embryonic node cells is critical for establishing left-right asymmetry, during mammalian development.

Centriole duplication

Before DNA replication, cells contain two centrioles, an older mother centriole, and a younger daughter centriole. During cell division, a new centriole grows at the proximal end of both mother and daughter centrioles. After duplication, the two centriole pairs (the freshly assembled centriole is now a daughter centriole in each pair) will remain attached to each other

orthogonal In mathematics, orthogonality is the generalization of the geometric notion of ''perpendicularity''. By extension, orthogonality is also used to refer to the separation of specific features of a system. The term also has specialized meanings in ...

ly until

mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...

. At that point the mother and daughter centrioles separate dependently on an enzyme called separase. The two centrioles in the centrosome are tied to one another. The mother centriole has radiating appendages at the

distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

end of its long axis and is attached to its daughter at the

proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...

end. Each daughter cell formed after cell division will inherit one of these pairs. Centrioles start duplicating when DNA replicates.

Origin

The last common ancestor of all

eukaryote Eukaryotes () are organisms whose cells have a 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 three domains of life. Bacte ...

s was a

ted cell with centrioles. Some lineages of eukaryotes, such as land plants, do not have centrioles except in their motile male gametes. Centrioles are completely absent from all cells of

conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...

and flowering plants, which do not have ciliate or flagellate gametes. It is unclear if the last common ancestor had one or two cilia. Important genes such as

centrins Centrins, also known as caltractins, are a family of calcium-binding phosphoproteins found in the centrosome of eukaryotes. Centrins are present in the centrioles and pericentriolar lattice. Human centrin genes are CETN1, CETN2 and CETN3. History ...

required for centriole growth, are only found in eukaryotes, and not in bacteria or

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 ...

Etymology and pronunciation

The word ''centriole'' () uses combining forms of ''centri-'' and '' -ole'', yielding "little central part", which describes a centriole's typical location near the center of the cell.

Atypical centrioles

Typical centrioles are made of 9 triplets of microtubules organized with radial symmetry. Centrioles can vary the number of microtubules and can be made of 9 doublets of microtubules (as in '' Drosophila melanogaster'') or 9 singlets of microtubules as in ''C. elegans''. Atypical centrioles are centrioles that do not have microtubules, such as the Proximal Centriole-Like found in ''D. melanogaster'' sperm, or that have microtubules with no radial symmetry, such as in the distal centriole of human

spermatozoon 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, t ...

. Atypical centrioles may have evolved at least eight times independently during vertebrate evolution and may evolve in the sperm after internal fertilization evolves. It wasn't clear why centriole become atypical until recently. The atypical distal centriole forms a dynamic basal complex (DBC) that, together with other structures in the sperm neck, facilitates a cascade of internal sliding, coupling tail beating with head kinking. The atypical distal centriole's properties suggest that it evolved into a transmission system that couples the sperm tail motors to the whole sperm, thereby enhancing sperm function.Khanal, S., M.R. Leung, A. Royfman, E.L. Fishman, B. Saltzman, H. Bloomfield-Gadelha, T. Zeev-Ben-Mordehai, and T. Avidor-Reiss. 2021. A dynamic basal complex modulates mammalian sperm movement. Nat Commun. 12:3808.. https://doi.org/10.1038/s41467-021-24011-0

References

{{Use dmy dates, date=April 2017 Centrosome Protein complexes Cell movement