Editing Microsatellite (section)

===Effects on proteins===
In mammals, 20–40% of proteins contain repeating sequences of [[amino acid]]s encoded by short sequence repeats.<ref name="Marcotte 1998">{{cite journal | vauthors = Marcotte EM, Pellegrini M, Yeates TO, Eisenberg D | title = A census of protein repeats | journal = Journal of Molecular Biology | volume = 293 | issue = 1 | pages = 151–60 | date = October 1999 | pmid = 10512723 | doi = 10.1006/jmbi.1999.3136 | s2cid = 11102561 }}</ref> Most of the short sequence repeats within protein-coding portions of the genome have a repeating unit of three nucleotides, since that length will not cause frame-shifts when mutating.<ref name="Sutherland 1995">{{cite journal | vauthors = Sutherland GR, Richards RI |author-link1=Grant Robert Sutherland | title = Simple tandem DNA repeats and human genetic disease | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 92 | issue = 9 | pages = 3636–41 | date = April 1995 | pmid = 7731957 | pmc = 42017 | doi = 10.1073/pnas.92.9.3636 | bibcode = 1995PNAS...92.3636S | doi-access = free }}</ref> Each trinucleotide repeating sequence is transcribed into a repeating series of the same amino acid. In yeasts, the most common repeated amino acids are glutamine, glutamic acid, asparagine, aspartic acid and serine.

Mutations in these repeating segments can affect the physical and chemical properties of proteins, with the potential for producing gradual and predictable changes in protein action.<ref name="Hancock 2005">{{cite journal | vauthors = Hancock JM, Simon M | title = Simple sequence repeats in proteins and their significance for network evolution | journal = Gene | volume = 345 | issue = 1 | pages = 113–8 | date = January 2005 | pmid = 15716087 | doi = 10.1016/j.gene.2004.11.023 }}</ref> For example, length changes in tandemly repeating regions in the [[Runx2]] gene lead to differences in facial length in domesticated dogs (''[[Canis familiaris]]''), with an association between longer sequence lengths and longer faces.<ref name="Fondon 2004">{{cite journal | vauthors = Fondon JW, Garner HR | title = Molecular origins of rapid and continuous morphological evolution | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 101 | issue = 52 | pages = 18058–63 | date = December 2004 | pmid = 15596718 | pmc = 539791 | doi = 10.1073/pnas.0408118101 | bibcode = 2004PNAS..10118058F | doi-access = free }}</ref> This association also applies to a wider range of Carnivora species.<ref name="Sears 2007">{{cite journal | vauthors = Sears KE, Goswami A, Flynn JJ, Niswander LA | title = The correlated evolution of Runx2 tandem repeats, transcriptional activity, and facial length in carnivora | journal = Evolution & Development | volume = 9 | issue = 6 | pages = 555–65 | year = 2007 | pmid = 17976052 | doi = 10.1111/j.1525-142X.2007.00196.x | s2cid = 26718314 }}</ref> Length changes in polyalanine tracts within the [[HOXA13]] gene are linked to [[hand-foot-genital syndrome]], a developmental disorder in humans.<ref name="Utsch 2002">{{cite journal | vauthors = Utsch B, Becker K, Brock D, Lentze MJ, Bidlingmaier F, Ludwig M | title = A novel stable polyalanine [poly(A)] expansion in the HOXA13 gene associated with hand-foot-genital syndrome: proper function of poly(A)-harbouring transcription factors depends on a critical repeat length? | journal = Human Genetics | volume = 110 | issue = 5 | pages = 488–94 | date = May 2002 | pmid = 12073020 | doi = 10.1007/s00439-002-0712-8 | s2cid = 22181414 }}</ref> Length changes in other triplet repeats are linked to more than 40 neurological diseases in humans, notably [[trinucleotide repeat disorder]]s such as [[fragile X syndrome]] and [[Huntington's disease]].<ref name="Pearson 2005">{{cite journal | vauthors = Pearson CE, Nichol Edamura K, Cleary JD | title = Repeat instability: mechanisms of dynamic mutations | journal = Nature Reviews. Genetics | volume = 6 | issue = 10 | pages = 729–42 | date = October 2005 | pmid = 16205713 | doi = 10.1038/nrg1689 | s2cid = 26672703 }}</ref> Evolutionary changes from replication slippage also occur in simpler organisms. For example, microsatellite length changes are common within surface membrane proteins in yeast, providing rapid evolution in cell properties.<ref name="Bowen 2006">{{cite journal | vauthors = Bowen S, Wheals AE | title = Ser/Thr-rich domains are associated with genetic variation and morphogenesis in Saccharomyces cerevisiae | journal = Yeast | volume = 23 | issue = 8 | pages = 633–40 | date = June 2006 | pmid = 16823884 | doi = 10.1002/yea.1381 | s2cid = 25142061 | doi-access = free }}</ref> Specifically, length changes in the FLO1 gene control the level of adhesion to substrates.<ref name="Verstrepen 2005">{{cite journal | vauthors = Verstrepen KJ, Jansen A, Lewitter F, Fink GR | title = Intragenic tandem repeats generate functional variability | journal = Nature Genetics | volume = 37 | issue = 9 | pages = 986–90 | date = September 2005 | pmid = 16086015 | pmc = 1462868 | doi = 10.1038/ng1618 }}</ref> Short sequence repeats also provide rapid evolutionary change to surface proteins in pathenogenic bacteria; this may allow them to keep up with immunological changes in their hosts.<ref name="Moxon 1994">{{cite journal | vauthors = Moxon ER, Rainey PB, Nowak MA, Lenski RE | title = Adaptive evolution of highly mutable loci in pathogenic bacteria | journal = Current Biology | volume = 4 | issue = 1 | pages = 24–33 | date = January 1994 | pmid = 7922307 | doi = 10.1016/S0960-9822(00)00005-1 | bibcode = 1994CBio....4...24M | s2cid = 11203457 }}</ref> Length changes in short sequence repeats in a fungus (''[[Neurospora crassa]]'') control the duration of its [[circadian clock]] cycles.<ref name="Michael 2007">{{cite journal | vauthors = Michael TP, Park S, Kim TS, Booth J, Byer A, Sun Q, Chory J, Lee K | display-authors = 6 | title = Simple sequence repeats provide a substrate for phenotypic variation in the Neurospora crassa circadian clock | journal = PLOS ONE | volume = 2 | issue = 8 | pages = e795 | date = August 2007 | pmid = 17726525 | pmc = 1949147 | doi = 10.1371/journal.pone.0000795 | bibcode = 2007PLoSO...2..795M | doi-access = free }}</ref>