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Gene duplication
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=== Rate of gene duplication === Comparisons of genomes demonstrate that gene duplications are common in most species investigated. This is indicated by variable copy numbers ([[copy number variation]]) in the genome of humans<ref>{{cite journal | vauthors = Sebat J, Lakshmi B, Troge J, Alexander J, Young J, Lundin P, MΓ₯nΓ©r S, Massa H, Walker M, Chi M, Navin N, Lucito R, Healy J, Hicks J, Ye K, Reiner A, Gilliam TC, Trask B, Patterson N, Zetterberg A, Wigler M | display-authors = 6 | title = Large-scale copy number polymorphism in the human genome | journal = Science | volume = 305 | issue = 5683 | pages = 525β8 | date = July 2004 | pmid = 15273396 | doi = 10.1126/science.1098918 | bibcode = 2004Sci...305..525S | s2cid = 20357402 }}</ref><ref>{{cite journal | vauthors = Iafrate AJ, Feuk L, Rivera MN, Listewnik ML, Donahoe PK, Qi Y, Scherer SW, Lee C | display-authors = 6 | title = Detection of large-scale variation in the human genome | journal = Nature Genetics | volume = 36 | issue = 9 | pages = 949β51 | date = September 2004 | pmid = 15286789 | doi = 10.1038/ng1416 | doi-access = free }}</ref> or fruit flies.<ref>{{cite journal | vauthors = Emerson JJ, Cardoso-Moreira M, Borevitz JO, Long M | title = Natural selection shapes genome-wide patterns of copy-number polymorphism in Drosophila melanogaster | journal = Science | volume = 320 | issue = 5883 | pages = 1629β31 | date = June 2008 | pmid = 18535209 | doi = 10.1126/science.1158078 | bibcode = 2008Sci...320.1629E | s2cid = 206512885 }}</ref> However, it has been difficult to measure the rate at which such duplications occur. Recent studies yielded a first direct estimate of the genome-wide rate of gene duplication in ''[[Caenorhabditis elegans|C. elegans]]'', the first multicellular eukaryote for which such as estimate became available. The gene duplication rate in ''C. elegans'' is on the order of 10<sup>β7</sup> duplications/gene/generation, that is, in a population of 10 million worms, one will have a gene duplication per generation. This rate is two orders of magnitude greater than the spontaneous rate of point mutation per nucleotide site in this species.<ref>{{cite journal | vauthors = Lipinski KJ, Farslow JC, Fitzpatrick KA, Lynch M, Katju V, Bergthorsson U | title = High spontaneous rate of gene duplication in Caenorhabditis elegans | journal = Current Biology | volume = 21 | issue = 4 | pages = 306β10 | date = February 2011 | pmid = 21295484 | pmc = 3056611 | doi = 10.1016/j.cub.2011.01.026 | bibcode = 2011CBio...21..306L }}</ref> Older (indirect) studies reported locus-specific duplication rates in bacteria, ''Drosophila'', and humans ranging from 10<sup>β3</sup> to 10<sup>β7</sup>/gene/generation.<ref>{{cite journal | vauthors = Anderson P, Roth J | title = Spontaneous tandem genetic duplications in Salmonella typhimurium arise by unequal recombination between rRNA (rrn) cistrons | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 78 | issue = 5 | pages = 3113β7 | date = May 1981 | pmid = 6789329 | pmc = 319510 | doi = 10.1073/pnas.78.5.3113 | bibcode = 1981PNAS...78.3113A | doi-access = free }}</ref><ref>{{cite journal | vauthors = Watanabe Y, Takahashi A, Itoh M, Takano-Shimizu T | title = Molecular spectrum of spontaneous de novo mutations in male and female germline cells of Drosophila melanogaster | journal = Genetics | volume = 181 | issue = 3 | pages = 1035β43 | date = March 2009 | pmid = 19114461 | pmc = 2651040 | doi = 10.1534/genetics.108.093385 }}</ref><ref>{{cite journal | vauthors = Turner DJ, Miretti M, Rajan D, Fiegler H, Carter NP, Blayney ML, Beck S, Hurles ME | display-authors = 6 | title = Germline rates of de novo meiotic deletions and duplications causing several genomic disorders | journal = Nature Genetics | volume = 40 | issue = 1 | pages = 90β5 | date = January 2008 | pmid = 18059269 | pmc = 2669897 | doi = 10.1038/ng.2007.40 }}</ref>
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