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{{Short description|Process by which genes shift in function}}
'''Functional divergence''' is the process by which genes, after [[gene duplication]], shift in function from an ancestral function. Functional divergence can result in either [[subfunctionalization]], where a paralog specializes one of several ancestral functions, or [[neofunctionalization]], where a totally new functional capability evolves. It is thought that this process of gene duplication and functional divergence is a major originator of molecular novelty and has produced the many large [[protein families]] that exist today.<ref>{{cite book | last1 = Gu | first1 = X | chapter = Functional divergence in protein (Family) sequence evolution | date = Jul 2003 | title = Origin and Evolution of New Gene Functions | journal = Genetica | volume = 118 | issue = 2–3| pages = 133–41 | doi=10.1007/978-94-010-0229-5_4| pmid = 12868604 | series = Contemporary Issues in Genetics and Evolution | isbn = 978-94-010-3982-6 }}</ref><ref>{{cite journal | last1 = Fay | first1 = JC | last2 = Wu | first2 = CI | year = 2003 | title = Sequence divergence, functional constraint, and selection in protein evolution | journal = Annu Rev Genom Hum Genet | volume = 4 | pages = 213–35 | doi = 10.1146/annurev.genom.4.020303.162528 | pmid = 14527302 | doi-access = free }}</ref>

Functional divergence is just one possible outcome of [[gene duplication]] events. Other fates include [[nonfunctionalization]] where one of the [[paralogs]] acquires [[deleterious]] [[mutations]] and becomes a [[pseudogene]] and [[superfunctionalization]] (reinforcement),<ref>{{cite journal | last1 = Dvornyk | first1 = V | last2 = Vinogradova | first2 = ON | last3 = Nevo | first3 = E | year = 2002 | title = Long-term microclimatic stress causes rapid adaptive radiation of kaiABC clock gene family in a cyanobacterium, Nostoc linckia, from "Evolution Canyons" I and II, Israel | journal = Proc Natl Acad Sci USA | volume = 99 | issue = 4| pages = 2082–2087 | doi=10.1073/pnas.261699498 | pmid=11842226 | pmc=123721| bibcode = 2002PNAS...99.2082D | doi-access = free }}</ref> where both paralogs maintain original function. While gene, chromosome, or whole genome duplication events are considered the canonical sources of functional divergence of [[paralogs]], [[orthologs]] (genes descended from speciation events) can also undergo functional divergence <ref>{{cite journal | last1 = Studer | first1 = RA | last2 = Robinson-Rechavi | first2 = M | year = 2009 | title = How confident can we be that orthologs are similar, but paralogs differ? | url = https://serval.unil.ch/notice/serval:BIB_39F8106EE698| journal = Trends in Genetics | volume = 25 | issue = 5| pages = 210–6 | doi = 10.1016/j.tig.2009.03.004 | pmid=19368988}}</ref><ref>{{cite journal | last1 = Studer | last2 = Robinson-Rechavi | first2 = M | year = 2010 | title = Large-scale analysis of orthologs and paralogs under covarion-like and constant-but-different models of amino acid evolution | journal = Molecular Biology and Evolution | volume = 27 | issue = 11| pages = 2618–2627 | doi = 10.1093/molbev/msq149 | pmid=20551039 | pmc=2955734}}</ref><ref>{{cite journal | last1 = Gharib | first1 = WH | last2 = Robinson-Rechavi | first2 = M | year = 2011 | title = When orthologs diverge between human and mouse | journal = Briefings in Bioinformatics | volume = 12 | issue = 5| pages = 436–441 | doi = 10.1093/bib/bbr031 | pmid = 21677033 | pmc = 3178054 }}</ref><ref>{{cite journal | last1 = Nehrt | first1 = NL | last2 = Clark | first2 = WT | last3 = Radivojac | first3 = P | last4 = Hahn | first4 = MW | year = 2011 | title = Testing the ortholog conjecture with comparative functional genomic data from mammals | journal = PLOS Computational Biology | volume = 7 | issue = 6| article-number = e1002073 | doi = 10.1371/journal.pcbi.1002073 | pmid=21695233 | pmc=3111532| bibcode = 2011PLSCB...7E2073N | doi-access = free }}</ref> and [[horizontal gene transfer]] can also result in multiple copies of a gene in a genome, providing the opportunity for functional divergence.

Many well known protein families are the result of this process, such as the ancient gene duplication event that led to the divergence of [[hemoglobin]] and [[myoglobin]], the more recent duplication events that led to the various subunit expansions (alpha and beta) of vertebrate [[hemoglobin]]s,<ref>{{cite journal | last1 = Storz | first1 = Jay F. | last2 = Hoffmann | first2 = Federico G. | last3 = Opazo | first3 = Juan C. | last4 = Moriyama | first4 = Hideaki | date = March 2008 | title = Adaptive Functional Divergence Among Triplicated α-Globin Genes in Rodents | journal = Genetics | volume = 178 | issue = 3| pages = 1623–1638 | doi = 10.1534/genetics.107.080903 | pmid = 18245844 | pmc = 2278084 }}</ref> or the expansion of G-protein alpha subunits <ref>{{cite journal | last1 = Zheng | first1 = Y | last2 = Xu | first2 = D | last3 = Gu | first3 = X | year = 2007 | title = Functional divergence after gene duplication and sequence-structure relationship: a case study of G-protein alpha subunits | url = https://lib.dr.iastate.edu/rtd/15063| journal = Journal of Experimental Zoology Part B: Molecular and Developmental Evolution | volume = 308 | issue = 1| pages = 85–96 | doi = 10.1002/jez.b.21140 | pmid = 17094082 | doi-access = free }}</ref>

==See also==
*[[Heterotachy]]
*[[Subfunctionalization]]
*[[Neofunctionalization]]

==References==
{{Reflist}}

{{Refbegin}}
* {{cite journal | last1 = Gu | first1 = X | date = Oct 2006 | title = A simple statistical method for estimating type-II (cluster-specific) functional divergence of protein sequences | journal = Mol Biol Evol | volume = 23 | issue = 10| pages = 1937–45 | doi=10.1093/molbev/msl056| pmid = 16864604 | doi-access = free }}
{{Refend}}

{{DEFAULTSORT:Functional Divergence}}
[[Category:Genetics]]
[[Category:Evolution]]

Functional divergence - История изменений

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