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{{short description|Preferential transmission of alleles over others during meiosis}}
'''Meiotic drive''' is a type of [[intragenomic conflict]], whereby one or more [[locus (genetics)|loci]] within a [[genome]] will affect a manipulation of the [[meiosis|meiotic]] process in such a way as to favor the transmission of one or more [[alleles]] over another, regardless of its phenotypic expression. More simply, meiotic drive is when one copy of a gene is passed on to offspring more than the expected 50% of the time. According to Buckler et al., "Meiotic drive is the subversion of meiosis so that particular genes are preferentially transmitted to the progeny. Meiotic drive generally causes the preferential segregation of small regions of the genome".<ref>{{cite journal | vauthors = Buckler ES, Phelps-Durr TL, Buckler CS, Dawe RK, Doebley JF, Holtsford TP | title = Meiotic drive of chromosomal knobs reshaped the maize genome | journal = Genetics | volume = 153 | issue = 1 | pages = 415–26 | date = September 1999 | doi = 10.1093/genetics/153.1.415 | pmid = 10471723 | pmc = 1460728 }}</ref>

==Meiotic drive in plants==

The first report of meiotic drive came from [[Marcus Morton Rhoades|Marcus Rhoades]] who in 1942 observed a violation of [[Mendelian_inheritance|Mendelian segregation]] ratios for the R locus - a [[gene]] controlling the production of the purple pigment [[anthocyanin]] in [[maize]] kernels - in a maize line carrying abnormal chromosome 10 (Ab10).<ref>{{cite journal | vauthors = Rhoades MM | title = Preferential Segregation in Maize | journal = Genetics | volume = 27 | issue = 4 | pages = 395–407 | date = July 1942 | doi = 10.1093/genetics/27.4.395 | pmid = 17247049 | pmc = 1209167 }}</ref> Ab10 differs from the normal chromosome 10 by the presence of a 150-base pair [[Heterochromatin|heterochromatic]] region called 'knob', which functions as a [[centromere]] during division (hence called 'neocentromere') and moves to the spindle poles faster than the centromeres during [[meiosis]] I and II.<ref>{{Cite journal |last1=Rhoades |first1=M.M. |last2=Vilkomerson |title=On the anaphase movement of chromosomes |journal=Proc. Natl. Acad. Sci. |year=1942 |publication-date=1942 |volume=28 |issue=10 |pages=433–436|doi=10.1073/pnas.28.10.433 |pmid=16588574 |pmc=1078510 |bibcode=1942PNAS...28..433R |doi-access=free }}</ref> The mechanism for this was later found to involve the activity of a [[kinesin]]-14 gene called Kinesin driver (''Kindr''). Kindr protein is a functional [[Minus-end-directed kinesin ATPase|minus-end directed]] motor, displaying quicker minus-end directed motility than an endogenous kinesin-14, such as Kin11. As a result ''Kindr'' outperforms the endogenous kinesins, pulling the 150 bp knobs to the poles faster than the centromeres and causing Ab10 to be preferentially inherited during meiosis <ref>{{cite journal | vauthors = Dawe RK, Lowry EG, Gent JI, Stitzer MC, Swentowsky KW, Higgins DM, Ross-Ibarra J, Wallace JG, Kanizay LB, Alabady M, Qiu W, Tseng KF, Wang N, Gao Z, Birchler JA, Harkess AE, Hodges AL, Hiatt EN | title = A Kinesin-14 Motor Activates Neocentromeres to Promote Meiotic Drive in Maize | journal = Cell | volume = 173 | issue = 4 | pages = 839–850.e18 | date = May 2018 | pmid = 29628142 | doi = 10.1016/j.cell.2018.03.009 | doi-access = free }}</ref>

==Meiotic drive in animals==

The unequal inheritance of gametes has been observed since the 1950s,<ref>{{cite journal |doi=10.1086/281969 |title=Meiotic Drive as an Evolutionary Force |journal=The American Naturalist |volume=91 |issue=857 |pages=105–110 |year=1957 | vauthors = Sandler L, Novitski E |s2cid=85014310 }}</ref> in contrast to [[Gregor Mendel]]'s First and Second Laws (the [[law of segregation]] and the [[law of independent assortment]]), which dictate that there is a random chance of each allele being passed on to offspring. Examples of selfish drive genes in animals have primarily been found in rodents and flies. These drive systems could play important roles in the process of [[speciation]]. For instance, the proposal that hybrid sterility ([[Haldane's rule]]) may arise from the divergent evolution of sex chromosome drivers and their suppressors.<ref>{{cite journal | vauthors = Helleu Q, Gérard PR, Montchamp-Moreau C | title = Sex chromosome drive | journal = Cold Spring Harbor Perspectives in Biology | volume = 7 | issue = 2 | article-number = a017616 | date = December 2014 | pmid = 25524548 | pmc = 4315933 | doi = 10.1101/cshperspect.a017616 }}</ref>

===Meiotic drive in mice===

Early observations of mouse t-haplotypes by Mary Lyon described numerous genetic loci on chromosome 17 that suppress X-chromosome sex ratio distortion.<ref>{{cite journal |doi=10.1016/0092-8674(84)90393-3 |pmid=6722884 |title=Transmission ratio distortion in mouse t-haplotypes is due to multiple distorter genes acting on a responder locus |journal=Cell |volume=37 |issue=2 |pages=621–628 |year=1984 |last1=Lyon |first1=Mary F. |s2cid=21065216 | name-list-style = vanc }}</ref><ref>{{cite journal |doi=10.1016/0092-8674(86)90770-1 |title=Male sterility of the mouse t-complex is due to homozygosity of the distorter genes |journal=Cell |volume=44 |issue=2 |pages=357–363 |year=1986 |last1=Lyon |first1=Mary F. |pmid=3943128 |s2cid=30795392 | name-list-style = vanc }}</ref> If a driver is left unchecked, this may lead to population extinction as the population would fix for the driver (e.g. a selfish X chromosome), removing the Y chromosome (and therefore males) from the population. The idea that meiotic drivers and their suppressors may govern speciation is supported by observations that mouse Y chromosomes lacking certain genetic loci produce female-biased offspring, implying these loci encode suppressors of drive.<ref>{{cite journal | vauthors = Cocquet J, Ellis PJ, Yamauchi Y, Mahadevaiah SK, Affara NA, Ward MA, Burgoyne PS | title = The multicopy gene Sly represses the sex chromosomes in the male mouse germline after meiosis | journal = PLOS Biology | volume = 7 | issue = 11 | article-number = e1000244 | date = November 2009 | pmid = 19918361 | pmc = 2770110 | doi = 10.1371/journal.pbio.1000244 | doi-access = free }}</ref> Moreover, matings of certain mouse strains used in research results in unequal offspring ratios. One gene responsible for sex ratio distortion in mice is r2d2 ([[R2d2 (mouse gene)|r2d2]] – responder to meiotic drive 2), which predicts which strains of mice can successfully breed without offspring sex ratio distortion.<ref>{{cite journal | vauthors = Didion JP, Morgan AP, Clayshulte AM, Mcmullan RC, Yadgary L, Petkov PM, Bell TA, Gatti DM, Crowley JJ, Hua K, Aylor DL, Bai L, Calaway M, Chesler EJ, French JE, Geiger TR, Gooch TJ, Garland T, Harrill AH, Hunter K, McMillan L, Holt M, Miller DR, O'Brien DA, Paigen K, Pan W, Rowe LB, Shaw GD, Simecek P, Sullivan PF, Svenson KL, Weinstock GM, Threadgill DW, Pomp D, Churchill GA, Pardo-Manuel de Villena F | title = A multi-megabase copy number gain causes maternal transmission ratio distortion on mouse chromosome 2 | journal = PLOS Genetics | volume = 11 | issue = 2 | article-number = e1004850 | date = February 2015 | pmid = 25679959 | pmc = 4334553 | doi = 10.1371/journal.pgen.1004850 | display-authors = 29 | doi-access = free }}
*{{cite web |author= |date=February 11, 2015 |title=R2d2 beats Mendel: Scientists find selfish gene that breaks long-held law of inheritance |website=Phys.org |url=http://phys.org/news/2015-02-r2d2-mendel-scientists-selfish-gene.html}}</ref>

===Meiotic drive in flies===
[[File:Stalk-eyed fly.jpg|thumb|right|A stalk-eyed fly]]
Selfish chromosomes of [[stalk-eyed flies]] have had ecological consequences. Driving X chromosomes lead to reductions in male fecundity and mating success, leading to [[frequency dependent selection]] maintaining both the driving alleles and wild-type alleles.<ref>{{cite journal | vauthors = Wilkinson GS, Johns PM, Kelleher ES, Muscedere ML, Lorsong A | title = Fitness effects of X chromosome drive in the stalk-eyed fly, Cyrtodiopsis dalmanni | journal = Journal of Evolutionary Biology | volume = 19 | issue = 6 | pages = 1851–60 | date = November 2006 | pmid = 17040382 | doi = 10.1111/j.1420-9101.2006.01169.x | url = https://www.life.umd.edu/faculty/wilkinson/Wilkinsonetal06.pdf | doi-access = free }}</ref>

Multiple species of fruit fly are known to have driving X chromosomes, of which the best-characterized are found in ''[[Drosophila simulans]]''. Three independent driving X chromosomes are known in ''D. simulans'', called Paris, Durham, and Winters. In Paris, the driving gene encodes a DNA modelling protein ("heterochromatin protein 1 D2" or ''HP1D2''), where the allele of the driving copy fails to prepare the male Y chromosome for meiosis.<ref>{{cite journal | vauthors = Helleu Q, Gérard PR, Dubruille R, Ogereau D, Prud'homme B, Loppin B, Montchamp-Moreau C | title = Rapid evolution of a Y-chromosome heterochromatin protein underlies sex chromosome meiotic drive | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 113 | issue = 15 | pages = 4110–5 | date = April 2016 | pmid = 26979956 | pmc = 4839453 | doi = 10.1073/pnas.1519332113 | bibcode = 2016PNAS..113.4110H | doi-access = free }}</ref> In Winters, the gene responsible ("Distorter on the X" or ''Dox'') has been identified, though the mechanism by which it acts is still unknown.<ref>{{cite journal | vauthors = Courret C, Gérard PR, Ogereau D, Falque M, Moreau L, Montchamp-Moreau C | title = X-chromosome meiotic drive in Drosophila simulans: a QTL approach reveals the complex polygenic determinism of Paris drive suppression | journal = Heredity | volume = 122 | issue = 6 | pages = 906–915 | date = December 2018 | pmid = 30518968 | pmc = 6781156 | doi = 10.1038/s41437-018-0163-1 }}</ref> The strong selective pressure imposed by these driving X chromosomes has given rise to suppressors of drive, of which the genes are somewhat known for Winters, Durham, and Paris. These suppressors encode hairpin RNAs which match the sequence of driver genes (such as ''Dox''), leading host [[RNA interference]] pathways to degrade ''Dox'' sequence.<ref>{{cite journal | vauthors = Lin CJ, Hu F, Dubruille R, Vedanayagam J, Wen J, Smibert P, Loppin B, Lai EC | title = The hpRNA/RNAi Pathway Is Essential to Resolve Intragenomic Conflict in the Drosophila Male Germline | journal = Developmental Cell | volume = 46 | issue = 3 | pages = 316–326.e5 | date = August 2018 | pmid = 30086302 | pmc = 6114144 | doi = 10.1016/j.devcel.2018.07.004 }}</ref> Autosomal suppressors of drive are known in ''[[Drosophila mediopunctata]]'',<ref name=pmid8276637>{{cite journal | vauthors = de Carvalho AB, Klaczko LB | title = Autosomal suppressors of sex-ratio in Drosophila mediopunctata | journal = Heredity | volume = 71 ( Pt 5) | issue = 5 | pages = 546–51 | date = November 1993 | pmid = 8276637 | doi = 10.1038/hdy.1993.174 | doi-access = free }}</ref> ''[[Drosophila paramelanica]]'',<ref>{{cite journal | vauthors = Stalker HD | title = The Genetic Systems Modifying Meiotic Drive in Drosophila Paramelanica | journal = Genetics | volume = 46 | issue = 2 | pages = 177–202 | date = February 1961 | doi = 10.1093/genetics/46.2.177 | pmid = 17248041 | pmc = 1210188 }}</ref> ''[[Drosophila quinaria]]'',<ref>{{cite journal | vauthors = Jaenike J | title = Suppression of Sex-Ratio Meiotic Drive and the Maintenance of Y-Chromosome Polymorphism in Drosophila | journal = Evolution; International Journal of Organic Evolution | volume = 53 | issue = 1 | pages = 164–174 | date = February 1999 | pmid = 28565182 | doi = 10.1111/j.1558-5646.1999.tb05342.x | doi-access =free }}</ref> and ''[[Drosophila testacea]]'',<ref>{{cite journal | vauthors = Keais GL, Hanson MA, Gowen BE, Perlman SJ | title = X chromosome drive in a widespread Palearctic woodland fly, Drosophila testacea | journal = Journal of Evolutionary Biology | volume = 30 | issue = 6 | pages = 1185–1194 | date = June 2017 | pmid = 28402000 | doi = 10.1111/jeb.13089 | doi-access = free }}</ref> emphasizing the importance of these drive systems in natural populations.

== See also==
* [[Fixed allele]]

== References ==
{{reflist}}

[[Category:Genetics]]

Meiotic drive - История изменений

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