Editing Centriole

{{Short description|Organelle in eukaryotic cells}}
{{See also|Centrosome}}
[[File:Blausen 0214 Centrioles.png|thumb|280px|3D rendering of centrioles showing the triplets]]

In [[cell biology]] a '''centriole''' is a cylindrical [[organelle]] composed mainly of a protein called [[tubulin]].<ref name="Eddé-1990">{{Cite journal|doi=10.1126/science.1967194|pmid=1967194|year=1990|last1=Eddé|first1=B|last2=Rossier|first2=J|last3=Le Caer|first3=JP|last4=Desbruyères|first4=E|last5=Gros|first5=F|last6=Denoulet|first6=P|title=Posttranslational glutamylation of alpha-tubulin|volume=247|issue=4938|pages=83–5|journal=Science|bibcode=1990Sci...247...83E}}</ref> Centrioles are found in most [[eukaryotic]] [[Cell (biology)|cell]]s, but are not present in conifers ([[Pinophyta]]), flowering plants ([[Flowering plant|angiosperms]]) and most [[fungi]], and are only present in the male gametes of [[charophytes]], [[bryophyte]]s, seedless [[vascular plant]]s, [[cycad]]s, and ''[[Ginkgo]]''.<ref>{{Cite journal|pmid=15928206|year=2005|last1=Quarmby|first1=LM|last2=Parker|first2=JD|title=Cilia and the cell cycle?|volume=169|issue=5|pages=707–10|doi=10.1083/jcb.200503053|pmc=2171619|journal=The Journal of Cell Biology}}</ref><ref>{{Cite journal|last1=Silflow|first1=CD|last2=Lefebvre|first2=PA|title=Assembly and motility of eukaryotic cilia and flagella. Lessons from Chlamydomonas reinhardtii|journal=Plant Physiology|year=2001|volume=127|issue=4|pages=1500–1507|doi=10.1104/pp.010807|pmid=11743094|pmc=1540183}}</ref> A bound pair of centrioles, surrounded by a highly ordered mass of dense material, called the [[pericentriolar material]] (PCM),<ref>{{Cite journal|last1=Lawo|first1=Steffen|last2=Hasegan|first2=Monica|last3=Gupta|first3=Gagan D.|last4=Pelletier|first4=Laurence|date=November 2012|title=Subdiffraction imaging of centrosomes reveals higher-order organizational features of pericentriolar material|url=https://pubmed.ncbi.nlm.nih.gov/23086237/|journal=Nature Cell Biology|volume=14|issue=11|pages=1148–1158|doi=10.1038/ncb2591|issn=1476-4679|pmid=23086237|s2cid=11286303}}</ref> makes up a structure called a [[centrosome]].<ref name="Eddé-1990"/>

Centrioles are typically made up of nine sets of [[microtubule|short microtubule]] triplets, arranged in a cylinder. Deviations from this structure include [[crabs]] and ''[[Drosophila melanogaster]]'' embryos, with nine doublets, and ''[[Caenorhabditis elegans]]'' [[sperm cells]] and early embryos, with nine singlets.<ref>{{Cite journal|pmid=15075224|year=2004|last1=Delattre|first1=M|last2=Gönczy|first2=P|title=The arithmetic of centrosome biogenesis|volume=117|issue=Pt 9|pages=1619–30|doi=10.1242/jcs.01128|journal=Journal of Cell Science|s2cid=7046196|url=https://infoscience.epfl.ch/record/182433/files/1619.full.pdf |archive-url=https://web.archive.org/web/20170818212233/http://infoscience.epfl.ch/record/182433/files/1619.full.pdf |archive-date=2017-08-18 |url-status=live|doi-access=free}}</ref><ref>{{Cite journal |pmid=15665853|year=2005 |last1=Leidel|first1=S. |last2=Delattre |first2=M. |last3=Cerutti |first3=L. |last4=Baumer |first4=K. |last5=Gönczy |first5=P|title=SAS-6 defines a protein family required for centrosome duplication in ''C. elegans'' and in human cells |volume=7 |issue=2 |pages=115–25 |doi=10.1038/ncb1220 |journal=Nature Cell Biology|s2cid=4634352 }}</ref> Additional proteins include [[centrin]], [[cenexin]] and [[tektin]].<ref name="Rieder-2001">{{Cite journal 
| pmid = 11567874 
| date=Oct 2001 | first2 = S. | last3=Khodjakov | first3 = A. 
| title = The centrosome in vertebrates: more than a microtubule-organizing center 
| volume = 11 
| issue = 10 
| pages = 413–419 
| issn = 0962-8924 
| journal = Trends in Cell Biology 
| doi = 10.1016/S0962-8924(01)02085-2
| last1=Rieder | first1=C. L. | last2=Faruki }}</ref>

The main function of centrioles is to produce [[Cilium|cilia]] during [[interphase]] and the [[Aster (cell biology)|aster]] and the [[spindle apparatus|spindle]] during cell division.

==History==
The centrosome was discovered jointly by [[Walther Flemming]] in 1875<ref>Flemming, W. (1875). Studien uber die Entwicklungsgeschichte der Najaden. Sitzungsgeber. Akad. Wiss. Wien 71, 81–147</ref><ref name="Bloodgood-2009">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.</ref> and [[Edouard Van Beneden]] in 1876.<ref>Van Beneden, E. (1876). Contribution a l’histoire de la vesiculaire germinative et du premier noyau embryonnaire. Bull. Acad. R. Belg (2me series) 42, 35–97.</ref><ref name="Bloodgood-2009"/> [[Edouard Van Beneden]] made the first observation of [[centrosomes]] as composed of two orthogonal centrioles in 1883.<ref>{{Cite journal | year = 2002 | doi = 10.1007/s00109-002-0374-y| pmid =  12226736 | issue =  9 | pages = 545–548| volume = 80| title = JMM - Past and Present| last1 = Wunderlich| journal = Journal of Molecular Medicine| first1 = V.| doi-access = free}}</ref> [[Theodor Boveri]] introduced the term "centrosome" in 1888<ref>Boveri, T. (1888). Zellen-Studien II. Die Befruchtung und Teilung des Eies von Ascaris megalocephala.
Jena. Z. Naturwiss. 22, 685–882.</ref><ref name="Bloodgood-2009"/><ref>Boveri, T. ''Ueber das Verhalten der Centrosomen bei der Befruchtung des Seeigel-Eies nebst allgemeinen Bemerkungen über Centrosomen und Verwandtes''. Verh. d. Phys.-Med. Ges. zu Würzburg, N. F., Bd. XXIX, 1895. [https://archive.org/details/bub_gb_UYcPAQAAMAAJ link].</ref><ref>Boveri, T. (1901). ''Zellen-Studien: Uber die Natur der Centrosomen. IV''. Fischer, Jena. [https://archive.org/details/zellenstudienbe00bovegoog link].</ref> and the term "centriole" in 1895.<ref>Boveri, T. (1895). Ueber die Befruchtungs und Entwickelungsfahigkeit kernloser Seeigeleier und uber die Moglichkeit ihrer Bastardierung. Arch. Entwicklungsmech. Org. (Wilhelm Roux) 2, 394–443.</ref><ref name="Bloodgood-2009"/> The [[basal body]] was named by [[Theodor Wilhelm Engelmann]] in 1880.<ref>Engelmann, T. W. (1880). Zur Anatomie und Physiologie der Flimmerzellen. Pflugers Arch. 23, 505–535.</ref><ref name="Bloodgood-2009"/> The pattern of centriole duplication was first worked out independently by [[Étienne de Harven]] and [[Joseph G. Gall]] c. 1950.<ref>{{cite book | last = Wolfe | first = Stephen L. | author-link = Stephen L. Wolfe | title = Biology: the foundations | publisher = Wadsworth | year = 1977|edition=First | url =https://archive.org/details/biologyfoundatio00wolf| url-access = registration | isbn = 9780534004903 }}</ref><ref>{{Cite book|volume=106 |year=1987 |pages=227–293|doi=10.1016/S0074-7696(08)61714-3 |title=The Centrosome and Its Role in the Organization of Microtubules |first1=I. A. |last1=Vorobjev |first2=E. S. |last2=Nadezhdina |series=International Review of Cytology|isbn=978-0-12-364506-7 |pmid=3294718}}. See also de Harven's own recollections of this work: {{Cite journal|title=Early observations of centrioles and mitotic spindle fibers by transmission electron microscopy|first=Etienne|last=de Harven |journal=Biology of the Cell |year=1994 |volume=80 |pages=107–109 |doi=10.1111/j.1768-322X.1994.tb00916.x|pmid=8087058|issue=2–3|s2cid=84594630|df=dmy-all}}</ref>

==Role in cell division==
[[File:Centriole-en.svg|thumb|280px|A mother and daughter centriole, attached [[orthogonally]]]]
Centrioles are involved in the organization of the [[spindle apparatus|mitotic spindle]] and in the completion of [[cytokinesis]].<ref name="Salisbury-2002">{{Cite journal|doi=10.1016/S0960-9822(02)01019-9|pmid=12176356|year=2002|last1=Salisbury|first1=JL|last2=Suino|first2=KM|last3=Busby|first3=R|last4=Springett|first4=M|title=Centrin-2 is required for centriole duplication in mammalian cells|volume=12|issue=15|pages=1287–92|journal=Current Biology|s2cid=1415623|doi-access=free}}</ref> 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 G<sub>1</sub> stage of [[interphase]] before centrioles can be synthesized later in a de novo fashion.<ref name="La Terra-2005">{{Cite journal|pmid=15738265|year=2005|last1=La Terra|first1=S|last2=English|first2=CN|last3=Hergert|first3=P|last4=McEwen|first4=BF|last5=Sluder|first5=G|last6=Khodjakov|first6=A|title=The de novo centriole assembly pathway in HeLa cells: cell cycle progression and centriole assembly/maturation|volume=168|issue=5|pages=713–22|doi=10.1083/jcb.200411126|pmc=2171814|journal=The Journal of Cell Biology}}</ref> Additionally, mutant flies lacking centrioles develop normally, although the adult flies' cells lack [[flagella]] and [[cilia]] and as a result, they die shortly after birth.<ref name="Basto-2006">{{Cite journal|pmid=16814722|year=2006|last1=Basto|first1=R|last2=Lau|first2=J|last3=Vinogradova|first3=T|last4=Gardiol|first4=A|last5=Woods|first5=CG|last6=Khodjakov|first6=A|last7=Raff|first7=JW|title=Flies without centrioles|volume=125|issue=7|pages=1375–86|doi=10.1016/j.cell.2006.05.025|journal=Cell|s2cid=2080684|doi-access=free}}</ref>
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]].<ref name="Feldman-2007">{{Cite journal|pmid=17518519|year=2007|last1=Feldman|first1=JL|last2=Geimer|first2=S|last3=Marshall|first3=WF|title=The mother centriole plays an instructive role in defining cell geometry|volume=5|issue=6|pages=e149|doi=10.1371/journal.pbio.0050149|pmc=1872036|journal=PLOS Biology |doi-access=free }}</ref><ref>{{Cite journal|doi=10.1016/S0955-0674(02)00017-0|pmid=12517710|year=2003|last1=Beisson|first1=J|last2=Wright|first2=M|title=Basal body/centriole assembly and continuity|volume=15|issue=1|pages=96–104|journal=Current Opinion in Cell Biology}}</ref> The position of the centriole determines the position of the nucleus and plays a crucial role in the spatial arrangement of the cell.

==Fertility==
[[Sperm]] centrioles are important for 2 functions:<ref>Avidor-Reiss, T., Khire, A., Fishman, E. L., & Jo, K. H. (2015). Atypical centrioles during sexual reproduction. Frontiers in cell and developmental biology, 3, 21.
Chicago	</ref> (1) to form the sperm [[flagellum]] and sperm movement and (2) for the development of the embryo after fertilization. The [[sperm]] supplies the centriole that creates the centrosome and microtubule system of the zygote.<ref>{{Cite book|chapter=The biology of fertilization in humans|editor=Patrizio, Pasquale|title=A color atlas for human assisted reproduction: laboratory and clinical insights|publisher=Lippincott Williams & Wilkins|year=2003|isbn=978-0-7817-3769-2|page=3|chapter-url=https://books.google.com/books?id=2SBoQ8H-KMIC&pg=PA3|author1=Hewitson, Laura  |author2=Schatten, Gerald P. |name-list-style=amp |access-date=2013-11-09|display-editors=etal}}</ref>

==Ciliogenesis==
In [[flagellate]]s and [[ciliate]]s, 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]].<ref name="Cui-2011">{{Cite journal|doi=10.1242/dmm.006262|pmc=3008963|pmid= 21045211|year=2011|author1=Cui, Cheng |author2=Chatterjee, Bishwanath |author3=Francis, Deanne |author4=Yu, Qing |author5=SanAgustin, Jovenal T. |author6=Francis, Richard |author7=Tansey, Terry |author8=Henry, Charisse |author9=Wang, Baolin |author10=Lemley, Bethan |author11=Pazour, Gregory J. |author12=Lo, Cecilia W. |title=Disruption of Mks1 localization to the mother centriole causes cilia defects and developmental malformations in Meckel-Gruber syndrome|volume=4|issue=1|pages=43–56|journal=Dis. Models Mech.}}</ref>

==Animal development==
[[File:Spindle centriole - embryonic brain mouse - TEM.jpg|thumb|Electron micrograph of a centriole from a mouse embryo.]]
Proper orientation of cilia via centriole positioning toward the posterior of embryonic node cells is critical for establishing [[left-right asymmetry]], during mammalian development.<ref>{{Cite journal|last1=Babu|first1=Deepak|last2=Roy|first2=Sudipto|date=2013-05-01|title=Left–right asymmetry: cilia stir up new surprises in the node|journal=Open Biology|language=en|volume=3|issue=5|pages=130052|doi=10.1098/rsob.130052|issn=2046-2441|pmc=3866868|pmid=23720541}}</ref>

==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]]ly until [[mitosis]]. At that point the mother and daughter centrioles separate dependently on an [[enzyme]] called [[separase]].<ref>{{Cite journal|pmid=16862117|year=2006|last1=Tsou|first1=MF|last2=Stearns|first2=T|title=Mechanism limiting centrosome duplication to once per cell cycle|volume=442|issue=7105|pages=947–51|doi=10.1038/nature04985|journal=Nature|bibcode = 2006Natur.442..947T |s2cid=4413248}}</ref>

The two centrioles in the centrosome are tied to one another. The mother centriole has radiating appendages at the [[Anatomical terms of location#Proximal and distal|distal]] end of its long axis and is attached to its daughter at the [[Anatomical terms of location#Proximal and distal|proximal]] end. Each daughter cell formed after cell division will inherit one of these pairs. Centrioles start duplicating when DNA replicates.<ref name="Salisbury-2002" />

==Origin==
[[Eukaryogenesis|LECA]], the last common ancestor of all [[eukaryote]]s was a [[cilia]]ted cell with centrioles.{{Citation needed|date=April 2024}} 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 [[Pinophyta|conifers]] and [[angiosperm|flowering plants]], which do not have ciliate or flagellate gametes.<ref>{{cite journal | last1 = Marshall | first1 = W.F. | year = 2009 | title = Centriole Evolution | journal = Current Opinion in Cell Biology | volume = 21 | issue = 1| pages = 14–19 | doi = 10.1016/j.ceb.2009.01.008 | pmid=19196504 | pmc=2835302}}</ref>
It is unclear if the last common ancestor had one<ref name="Bornens-2007">{{Cite book| doi = 10.1007/978-0-387-74021-8_10| last2 = Azimzadeh| pmid = 17977464| isbn = 978-0-387-74020-1| year = 2007| pages = [https://archive.org/details/eukaryoticmembra00gasp/page/119 119–129]| series = Advances in Experimental Medicine and Biology| last1 = Bornens| first2 = J.| chapter = Origin and Evolution of the Centrosome| title = Eukaryotic Membranes and Cytoskeleton| volume = 607| first1 = M.| chapter-url-access = registration| chapter-url = https://archive.org/details/eukaryoticmembra00gasp| url = https://archive.org/details/eukaryoticmembra00gasp/page/119}}</ref> or two cilia.<ref>{{Cite journal | doi = 10.1093/gbe/evp011 | last1 = Rogozin | first1 = I. B. | last2 = Basu | first2 = M. K. | last3 = Csürös | first3 = M. | last4 = Koonin | first4 = E. V. | title = Analysis of Rare Genomic Changes Does Not Support the Unikont-Bikont Phylogeny and Suggests Cyanobacterial Symbiosis as the Point of Primary Radiation of Eukaryotes | journal = Genome Biology and Evolution | volume = 1 | pages = 99–113 | year = 2009 | pmid = 20333181 | pmc = 2817406}}</ref> Important genes such as those coding for [[centrins]], required for centriole growth, are only found in eukaryotes, and not in [[bacteria]] or [[archaea]].<ref name="Bornens-2007"/>

==Etymology and pronunciation==
The word ''centriole'' ({{IPAc-en|ˈ|s|ɛ|n|t|r|i|oʊ|l}}) uses [[classical compound|combining forms]] of ''centri-'' and ''[[wikt:-ole#Etymology 2|-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.<ref>{{cite journal |doi=10.1016/j.ceb.2012.10.016 |pmid=23199753 |pmc=3578074 |title=Building a centriole |journal=Current Opinion in Cell Biology |volume=25 |issue=1 |pages=72–7 |year=2013 |last1=Avidor-Reiss |first1=Tomer |last2=Gopalakrishnan |first2=Jayachandran }}</ref> 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 [[Caenorhabditis elegans|''C. elegans'']]. Atypical centrioles are centrioles that do not have microtubules, such as the [[Proximal Centriole-Like]] found in ''D. melanogaster'' sperm,<ref>{{cite journal |doi=10.1534/genetics.109.101709 |pmid=19293139 |pmc=2674812 |title=A Proximal Centriole-Like Structure is Present in Drosophila Spermatids and Can Serve as a Model to Study Centriole Duplication |journal=Genetics |volume=182 |issue=1 |pages=133–44 |year=2009 |last1=Blachon |first1=S |last2=Cai |first2=X |last3=Roberts |first3=K. A |last4=Yang |first4=K |last5=Polyanovsky |first5=A |last6=Church |first6=A |last7=Avidor-Reiss |first7=T }}</ref> or that have microtubules with no radial symmetry, such as in the distal centriole of human [[spermatozoon]].<ref>{{cite journal |doi=10.1038/s41467-018-04678-8 |pmid=29880810 |pmc=5992222 |title=A novel atypical sperm centriole is functional during human fertilization |journal=Nature Communications |volume=9 |issue=1 |pages=2210 |year=2018 |last1=Fishman |first1=Emily L |last2=Jo |first2=Kyoung |last3=Nguyen |first3=Quynh P. H |last4=Kong |first4=Dong |last5=Royfman |first5=Rachel |last6=Cekic |first6=Anthony R |last7=Khanal |first7=Sushil |last8=Miller |first8=Ann L |last9=Simerly |first9=Calvin |last10=Schatten |first10=Gerald |last11=Loncarek |first11=Jadranka |last12=Mennella |first12=Vito |last13=Avidor-Reiss |first13=Tomer |bibcode=2018NatCo...9.2210F }}</ref> Atypical centrioles may have evolved at least eight times independently during vertebrate evolution and may evolve in the sperm after [[internal fertilization]] evolves.<ref>Turner, K., N. Solanki, H.O. Salouha, and T. Avidor-Reiss. 2022. Atypical Centriolar Composition Correlates with Internal Fertilization in Fish. Cells. 11:758, https://www.mdpi.com/2073-4409/11/5/758</ref>

It wasn't clear why the centriole became 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 that couples 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.<ref>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</ref>

==References==
{{Reflist}}

{{Centrosome}}
{{Cellular structures}}

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{{Use dmy dates|date=April 2017}}

[[Category:Centrosome]]
[[Category:Protein complexes]]
[[Category:Cell movement]]