Editing Centriole (section)

==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>