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Gene duplication

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206: 479: 142:, can occasionally act on cellular mRNA. Transcripts are reverse transcribed to DNA and inserted into random place in the genome, creating retrogenes. Resulting sequence usually lack introns and often contain poly(A) sequences that are also integrated into the genome. Many retrogenes display changes in gene regulation in comparison to their parental gene sequences, which sometimes results in novel functions. Retrogenes can move between different chromosomes to shape chromosomal evolution. 107: 353:. Such detrimental mutations are likely to be lost from the population and will not be preserved or develop novel functions. However, many duplications are, in fact, not detrimental or beneficial, and these neutral sequences may be lost or may spread through the population through random fluctuations via 393:
if a homolog to a human gene can be found in the genome of that species, but only if the homolog is orthologous. If they are paralogs and resulted from a gene duplication event, their functions are likely to be too different. One or more copies of duplicated genes that constitute a gene family may be
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Polyploidy is also a well known source of speciation, as offspring, which have different numbers of chromosomes compared to parent species, are often unable to interbreed with non-polyploid organisms. Whole genome duplications are thought to be less detrimental than aneuploidy as the relative dosage
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After a whole genome duplication, there is a relatively short period of genome instability, extensive gene loss, elevated levels of nucleotide substitution and regulatory network rewiring. In addition, gene dosage effects play a significant role. Thus, most duplicates are lost within a short period,
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begins to copy the DNA. At some point during the replication process, the polymerase dissociates from the DNA and replication stalls. When the polymerase reattaches to the DNA strand, it aligns the replicating strand to an incorrect position and incidentally copies the same section more than once.
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that occurs during meiosis between misaligned homologous chromosomes. The chance of it happening is a function of the degree of sharing of repetitive elements between two chromosomes. The products of this recombination are a duplication at the site of the exchange and a reciprocal deletion. Ectopic
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of it have no deleterious effects to its host organism. If one copy of a gene experiences a mutation that affects its original function, the second copy can serve as a 'spare part' and continue to function correctly. Thus, duplicate genes accumulate mutations faster than a functional single-copy
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is on the order of 10 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. Older (indirect) studies
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Gene duplications can also be identified through the use of next-generation sequencing platforms. The simplest means to identify duplications in genomic resequencing data is through the use of paired-end sequencing reads. Tandem duplications are indicated by sequencing read pairs which map in
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Whole-genome duplications are also frequent in cancers, detected in 30% to 36% of tumors from the most common cancer types. Their exact role in carcinogenesis is unclear, but they in some cases lead to loss of chromatin segregation leading to chromatin conformation changes that in turn lead to
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Rapid evolution and functional divergence have been observed at the level of the transcription of duplicated genes, usually by point mutations in short transcription factor binding motifs. Furthermore, rapid evolution of protein phosphorylation motifs, usually embedded within rapidly evolving
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Paralogs can be identified in single genomes through a sequence comparison of all annotated gene models to one another. Such a comparison can be performed on translated amino acid sequences (e.g. BLASTp, tBLASTx) to identify ancient duplications or on DNA nucleotide sequences (e.g. BLASTn,
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and performs two functions, often neither one of these two functions can be changed without affecting the other function. In this way, partitioning the ancestral functions into two separate genes can allow for adaptive specialization of subfunctions, thereby providing an adaptive benefit.
328:) or DDC (duplication-degeneration-complementation) model, in which the functionality of the original gene is distributed among the two copies. Neither gene can be lost, as both now perform important non-redundant functions, but ultimately neither is able to achieve novel functionality. 3096:
Quinton, Ryan J.; DiDomizio, Amanda; Vittoria, Marc A.; Kotýnková, Kristýna; Ticas, Carlos J.; Patel, Sheena; Koga, Yusuke; Vakhshoorzadeh, Jasmine; Hermance, Nicole; Kuroda, Taruho S.; Parulekar, Neha; Taylor, Alison M.; Manning, Amity L.; Campbell, Joshua D.; Ganem, Neil J. (2021).
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occurs when nondisjunction at a single chromosome results in an abnormal number of chromosomes. Aneuploidy is often harmful and in mammals regularly leads to spontaneous abortions (miscarriages). Some aneuploid individuals are viable, for example trisomy 21 in humans, which leads to
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Bielski, Craig M.; Zehir, Ahmet; Penson, Alexander V.; Donoghue, Mark T. A.; Chatila, Walid; Armenia, Joshua; Chang, Matthew T.; Schram, Alison M.; Jonsson, Philip; Bandlamudi, Chaitanya; Razavi, Pedram; Iyer, Gopa; Robson, Mark E.; Stadler, Zsofia K.; Schultz, Nikolaus (2018).
603:. In such cases the genetic duplication occurs in a somatic cell and affects only the genome of the cancer cells themselves, not the entire organism, much less any subsequent offspring. Recent comprehensive patient-level classification and quantification of driver events in 193:
intrinsically disordered regions is another contributing factor for survival and rapid adaptation/neofunctionalization of duplicate genes. Thus, a link seems to exist between gene regulation (at least at the post-translational level) and genome evolution.
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Subfunctionalization can occur through neutral processes in which mutations accumulate with no detrimental or beneficial effects. However, in some cases subfunctionalization can occur with clear adaptive benefits. If an ancestral gene is
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gene, over generations of organisms, and it is possible for one of the two copies to develop a new and different function. Some examples of such neofunctionalization is the apparent mutation of a duplicated digestive gene in a family of
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megablast) to identify more recent duplications. Most studies to identify gene duplications require reciprocal-best-hits or fuzzy reciprocal-best-hits, where each paralog must be the other's single best match in a sequence comparison.
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Another possible fate for duplicate genes is that both copies are equally free to accumulate degenerative mutations, so long as any defects are complemented by the other copy. This leads to a neutral "
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Gene duplications are an essential source of genetic novelty that can lead to evolutionary innovation. Duplication creates genetic redundancy, where the second copy of the gene is often free from
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however, a considerable fraction of duplicates survive. Interestingly, genes involved in regulation are preferentially retained. Furthermore, retention of regulatory genes, most notably the
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during meiosis which results in additional copies of the entire genome. Polyploidy is common in plants, but it has also occurred in animals, with two rounds of whole genome duplication (
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Gene duplication does not necessarily constitute a lasting change in a species' genome. In fact, such changes often don't last past the initial host organism. From the perspective of
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abnormal orientations. Through a combination of increased sequence coverage and abnormal mapping orientation, it is possible to identify duplications in genomic sequencing data.
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It is important (but often difficult) to differentiate between paralogs and orthologs in biological research. Experiments on human gene function can often be carried out on other
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hybridization (array CGH), are used to detect chromosomal abnormalities, such as microduplications, in a high throughput fashion from genomic DNA samples. In particular, DNA
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Lambuta, Ruxandra A.; Nanni, Luca; Liu, Yuanlong; Diaz-Miyar, Juan; Iyer, Arvind; Tavernari, Daniele; Katanayeva, Natalya; Ciriello, Giovanni; Oricchio, Elisa (2023-03-15).
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Emerson JJ, Cardoso-Moreira M, Borevitz JO, Long M (June 2008). "Natural selection shapes genome-wide patterns of copy-number polymorphism in Drosophila melanogaster".
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elements offer one source of repetitive DNA that can facilitate recombination, and they are often found at duplication breakpoints in plants and mammals.
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recombination is typically mediated by sequence similarity at the duplicate breakpoints, which form direct repeats. Repetitive genetic elements such as
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Kellis M, Birren BW, Lander ES (April 2004). "Proof and evolutionary analysis of ancient genome duplication in the yeast Saccharomyces cerevisiae".
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regions of a chromosome. Many LCRs, due to their size (>1Kb), similarity, and orientation, are highly susceptible to duplications and deletions.
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Sebat J, Lakshmi B, Troge J, Alexander J, Young J, Lundin P, et al. (July 2004). "Large-scale copy number polymorphism in the human genome".
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Comparisons of genomes demonstrate that gene duplications are common in most species investigated. This is indicated by variable copy numbers (
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with annotated bands and sub-bands as used for the nomenclature of chromosome abnormalities. It shows dark and white regions as seen on
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into an antifreeze gene and duplication leading to a novel snake venom gene and the synthesis of 1 beta-hydroxytestosterone in pigs.
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levels of thousands of genes across many treatments or experimental conditions, greatly facilitating the evolutionary studies of
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Amoutzias, Grigoris D.; He, Ying; Gordon, Jonathan; Mossialos, Dimitris; Oliver, Stephen G.; Van de Peer, Yves (2010-02-16).
1738:"Nonrandom divergence of gene expression following gene and genome duplications in the flowering plant Arabidopsis thaliana" 2612:
Des Marais DL, Rausher MD (August 2008). "Escape from adaptive conflict after duplication in an anthocyanin pathway gene".
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machinery as well as through fortuitous capture by selfish genetic elements. Common sources of gene duplications include
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cohorts revealed that there are on average 12 driver events per tumor, of which 1.5 are amplifications of oncogenes.
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Replication slippage is also often facilitated by repetitive sequences, but requires only a few bases of similarity.
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Mao R, Pevsner J (2005). "The use of genomic microarrays to study chromosomal abnormalities in mental retardation".
3261: 2398: 880: 844: 840: 350: 2165:"Molecular spectrum of spontaneous de novo mutations in male and female germline cells of Drosophila melanogaster" 3421: 1170:"Retrogene Duplication and Expression Patterns Shaped by the Evolution of Sex Chromosomes in Malaria Mosquitoes" 1168:
Miller, Duncan; Chen, Jianhai; Liang, Jiangtao; Betrán, Esther; Long, Manyuan; Sharakhov, Igor V. (2022-05-28).
4583: 4178: 325: 279:(1970). Ohno argued that gene duplication is the most important evolutionary force since the emergence of the 1642:"Gene-balanced duplications, like tetraploidy, provide predictable drive to increase morphological complexity" 579:, or it can occur naturally, as described above. If it's a natural duplication, it can still take place in a 228:, the first multicellular eukaryote for which such as estimate became available. The gene duplication rate in 4719: 3655: 3501: 3488: 1106: 382:
genes present in different species which are each originally derived from the same ancestral sequence. (See
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is an error in DNA replication that can produce duplications of short genetic sequences. During replication
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Conant GC, Wolfe KH (December 2008). "Turning a hobby into a job: how duplicated genes find new functions".
4762: 4367: 4048: 506: 181:) in the vertebrate lineage leading to humans. It has also occurred in the hemiascomycete yeasts ~100 mya. 4053: 4287: 4043: 3254: 2263:"Inventing an arsenal: adaptive evolution and neofunctionalization of snake venom phospholipase A2 genes" 383: 398:
that causes significant variation between them in their sequence and finally may become responsible for
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Often the resulting genomic variation leads to gene dosage dependent neurological disorders such as
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Vyatkin, Alexey D.; Otnyukov, Danila V.; Leonov, Sergey V.; Belikov, Aleksey V. (14 January 2022).
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Taylor JS, Raes J (2004). "Duplication and divergence: the evolution of new genes and old ideas".
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Turner DJ, Miretti M, Rajan D, Fiegler H, Carter NP, Blayney ML, et al. (January 2008).
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Iafrate AJ, Feuk L, Rivera MN, Listewnik ML, Donahoe PK, Qi Y, et al. (September 2004).
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Casneuf, Tineke; De Bodt, Stefanie; Raes, Jeroen; Maere, Steven; Van de Peer, Yves (2006).
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between the homologs of gene duplicates due to less or no similarity in their sequences.
399: 252: 42: 3206: 3179: 3163: 3114: 2815: 2625: 2564: 2464: 2121: 2017: 1925: 1862: 1602: 1539: 1360: 1295: 379: 371: 4680: 3819: 3727: 3546: 3139: 3098: 3073: 3040: 2931: 2904: 2800:"Rapid evolution of expression and regulatory divergences after yeast gene duplication" 2740: 2713: 2694: 2645: 2594: 2528: 2503: 2484: 2424: 2337: 2289: 2262: 2238: 2213: 2189: 2164: 2081: 2056: 2037: 1945: 1889: 1846: 1772: 1737: 1567: 1499: 1464: 1445: 1388: 1325: 1255: 1228: 1204: 1169: 1029: 551: 547: 426: 418: 271:; this stance has been held by members of the scientific community for over 100 years. 3907: 2834: 2799: 2140: 2105: 1125: 4767: 4675: 4379: 3359: 3285: 3211: 3193: 3144: 3126: 3078: 3060: 3018: 2936: 2839: 2780: 2745: 2686: 2637: 2586: 2533: 2476: 2431: 2402: 2372: 2329: 2294: 2243: 2194: 2145: 2086: 2041: 2029: 1986: 1937: 1894: 1876: 1824: 1816: 1777: 1759: 1718: 1710: 1671: 1663: 1622: 1614: 1559: 1551: 1504: 1486: 1437: 1429: 1380: 1372: 1317: 1309: 1260: 1209: 1191: 760: 725: 2698: 1949: 1449: 4305: 3938: 3881: 3344: 3201: 3183: 3134: 3118: 3068: 3052: 2926: 2916: 2829: 2819: 2772: 2735: 2725: 2676: 2629: 2598: 2578: 2551:
Stoltzfus A (August 1999). "On the possibility of constructive neutral evolution".
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Force A, Lynch M, Pickett FB, Amores A, Yan YL, Postlethwait J (April 1999).
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for duplications of parts of a chromosome. For example, dup(17p12) causes
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Schematic of a region of a chromosome before and after a duplication event
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chromosome pairs, both the female (XX) and male (XY) versions of the two
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was one of the most famous developers of this theory in his classic book
272: 53:. Gene duplications can arise as products of several types of errors in 3041:"Genome doubling shapes the evolution and prognosis of advanced cancers" 2633: 2472: 1547: 1368: 562:. Genetic amplification can occur artificially, as with the use of the 4617: 4612: 4565: 4558: 4553: 4538: 4533: 4374: 4309: 4058: 4000: 3848: 3665: 2776: 2582: 2229: 1658: 1641: 1074: 1059: 750: 491: 458: 446: 333: 284: 166: 150: 74: 70: 58: 3246: 3164:"Whole-genome doubling drives oncogenic loss of chromatin segregation" 4622: 4607: 4594: 4346: 4134: 3965: 3767: 3458: 2057:"High spontaneous rate of gene duplication in Caenorhabditis elegans" 669: 596: 512: 498: 487: 483: 303: 268: 2325: 1229:"Two rounds of whole genome duplication in the ancestral vertebrate" 4653: 4599: 4501: 4445: 4440: 4331: 4292: 4248: 4150: 3955: 3891: 3777: 3541: 3437: 2162: 1981: 1964: 1847:"Posttranslational regulation impacts the fate of duplicated genes" 804: 800: 587:
cell (which would be necessary for a lasting evolutionary change).
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The two genes that exist after a gene duplication event are called
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Mental Retardation and Developmental Disabilities Research Reviews
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A brief overview of mutation, gene duplication and translocation
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events can be quite common. It is believed that the entire
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oncogenic epigenetic and transcriptional modifications.
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with a similar function and/or structure. By contrast,
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are the most prolific genome duplicators. For example,
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International System for Human Cytogenomic Nomenclature
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reported locus-specific duplication rates in bacteria,
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Li, Wen-Hsiung; Yang, Jing; Gu, Xun (November 2005).
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Gene duplication is believed to play a major role in
2714:"Gene family evolution across 12 Drosophila genomes" 294:underwent duplication about 100 million years ago. 237:, and humans ranging from 10 to 10/gene/generation. 1689:Davis, Jerel C.; Petrov, Dmitri A. (October 2005). 402:. This may also render the chances and the rate of 365: 3006: 3004: 3002: 3000: 2998: 2996: 2994: 2992: 2990: 2988: 2986: 2984: 2982: 2980: 2978: 2976: 2974: 2972: 2970: 2450: 2423: 1463:Conant, Gavin C.; Wolfe, Kenneth H. (April 2006). 1342: 3010: 2968: 2966: 2964: 2962: 2960: 2958: 2956: 2954: 2952: 2950: 2611: 1640:Freeling, Michael; Thomas, Brian C. (July 2006). 306:), meaning that it has six copies of its genome. 4739: 2880: 2348: 611:Common oncogene amplifications in human cancers 1797:"Expression divergence between duplicate genes" 1220: 2947: 2544: 1639: 27:Duplication of a gene sequence within a genome 3923: 3422: 3262: 3234:A bibliography on gene and genome duplication 1408:"The evolutionary consequences of polyploidy" 464: 361:Identifying duplications in sequenced genomes 2797: 2103: 1521: 1277: 1040:Gene amplification in Paramecium tetraurelia 84: 2444: 2311: 1688: 1584: 1462: 1278:Wolfe, K. H.; Shields, D. C. (1997-06-12). 213: 200: 3930: 3916: 3429: 3415: 3269: 3255: 2762: 2711: 2495: 2354: 1226: 1107:"Evolution by gene duplication: an update" 449:technology can simultaneously monitor the 3205: 3187: 3138: 3072: 2930: 2920: 2833: 2823: 2739: 2729: 2712:Hahn MW, Han MV, Han SG (November 2007). 2680: 2662: 2605: 2572: 2550: 2527: 2415: 2383: 2288: 2278: 2237: 2188: 2139: 2129: 2080: 1980: 1888: 1870: 1771: 1753: 1657: 1498: 1480: 1423: 1303: 1254: 1244: 1203: 1185: 1146:. MedicineNet. 2012-03-19. Archived from 1098: 1794: 523:(ISCN) is an international standard for 477: 204: 197:of individual genes should be the same. 105: 94:Duplications arise from an event termed 89: 3276: 2798:Gu X, Zhang Z, Huang W (January 2005). 1585:Lynch, M.; Conery, J. S. (2000-11-10). 1104: 433:Genomic microarrays detect duplications 309: 240: 114: 14: 4740: 2862:Coriell Institute for Medical Research 2369:10.1146/annurev.genet.38.072902.092831 2305: 2254: 566:technique to amplify short strands of 3911: 3410: 3250: 2260: 1840: 1838: 130: 2858:"ISCN Symbols and Abbreviated Terms" 2705: 2426:Sex Chromosomes and Sex-linked Genes 2421: 2389: 1405: 599:are a common cause of many types of 490:. Each row is vertically aligned at 209:Evolutionary fate of duplicate genes 4758:Modification of genetic information 541: 24: 3498:Short tandem repeat/Microsatellite 2663:Lee JA, Lupski JR (October 2006). 1835: 1227:Dehal P, Boore JL (October 2005). 189:, has led to adaptive innovation. 25: 4779: 3340:Models of nucleotide substitution 3226: 3014:The genetic basis of human cancer 3011:Kinzler KW, Vogelstein B (2002). 1114:Trends in Ecology & Evolution 590: 384:Homology of sequences in genetics 18:Amplification (molecular biology) 1140:"Definition of Gene duplication" 441:, also called array comparative 413:Most gene duplications exist as 366:Criteria and single genome scans 3155: 3089: 3031: 2896: 2874: 2850: 2791: 2756: 2656: 2205: 2156: 2104:Anderson P, Roth J (May 1981). 2097: 2048: 1997: 1956: 1905: 1788: 1729: 1682: 1633: 1578: 554:is one of many ways in which a 473: 4584:Last universal common ancestor 4179:Defective interfering particle 3502:Trinucleotide repeat disorders 2553:Journal of Molecular Evolution 1515: 1456: 1399: 1336: 1271: 1161: 1132: 326:constructive neutral evolution 13: 1: 4748:Evolutionary biology concepts 4720:Clonally transmissible cancer 4156:Satellite-like nucleic acids 3489:Variable number tandem repeat 2395:Evolution by gene duplication 1611:10.1126/science.290.5494.1151 1406:Otto, Sarah P. 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McGraw-Hill. p. 116. 2922:10.1371/journal.pgen.1009996 2731:10.1371/journal.pgen.0030197 2682:10.1016/j.neuron.2006.09.027 1482:10.1371/journal.pbio.0040109 1246:10.1371/journal.pbio.0030314 351:Pelizaeus–Merzbacher disease 7: 2520:10.1093/genetics/151.4.1531 2181:10.1534/genetics.108.093385 1144:medterms medical dictionary 1011: 536:Charcot–Marie–Tooth disease 10: 4784: 4276:Class II or DNA transposon 4271:Class I or retrotransposon 3396:Nonsynonymous substitution 3189:10.1038/s41586-023-05794-2 3123:10.1038/s41586-020-03133-3 1425:10.1016/j.cell.2007.10.022 510: 465:Next generation sequencing 457:after gene duplication or 313: 244: 4646: 4589:Earliest known life forms 4574: 4487: 4463:Repeated sequences in DNA 4355: 4237: 4226: 4199: 4171: 4121: 4110: 4083: 4072: 4017: 3946: 3857: 3810: 3678: 3646: 3623: 3600: 3591: 3582: 3557: 3517: 3474: 3465: 3456: 3368: 3327: 3284: 3057:10.1038/s41588-018-0165-1 2357:Annual Review of Genetics 2261:Lynch VJ (January 2007). 2073:10.1016/j.cub.2011.01.026 1813:10.1016/j.tig.2005.08.006 1707:10.1016/j.tig.2005.07.008 964: 929: 864: 834: 794: 759: 724: 699: 634: 564:polymerase chain reaction 394:affected by insertion of 281:universal common ancestor 85:Mechanisms of duplication 4436:Endogenous viral element 4254:Horizontal gene transfer 2893:Updated : 4/23/2014 2314:Nature Reviews. Genetics 2267:BMC Evolutionary Biology 1050:Horizontal gene transfer 302:is hexaploid (a kind of 214:Rate of gene duplication 201:As an evolutionary event 171:whole genome duplication 4133:dsDNA satellite virus ( 3391:Synonymous substitution 3335:Models of DNA evolution 2825:10.1073/pnas.0409186102 2026:10.1126/science.1158078 1934:10.1126/science.1098918 1872:10.1073/pnas.0911603107 1755:10.1186/gb-2006-7-2-r13 1080:Tandem exon duplication 1065:Mobile genetic elements 340: 35:chromosomal duplication 4691:Helper dependent virus 4007:Biological dark matter 3897:Protein tandem repeats 3825:Tandemly arrayed genes 2881:Cassandra L. Kniffin. 2131:10.1073/pnas.78.5.3113 991:Small cell lung cancer 516: 210: 111: 4451:Endogenous retrovirus 4424:Origin of replication 4140:ssDNA satellite virus 4130:ssRNA satellite virus 3314:Stabilizing selection 3299:Directional selection 2280:10.1186/1471-2148-7-2 1187:10.3390/genes13060968 1085:Unequal crossing over 901:Hepatocellular cancer 511:Further information: 481: 396:transposable elements 374:and usually code for 220:copy number variation 208: 109: 96:unequal crossing-over 90:Ectopic recombination 63:ectopic recombination 4395:Secondary chromosome 4390:Extrachromosomal DNA 4266:Transposable element 3870:Pathogenicity island 3304:Disruptive selection 1019:Comparative genomics 866:Head and neck cancer 507:mitochondrial genome 322:subfunctionalization 316:Subfunctionalization 310:Subfunctionalization 247:Neofunctionalization 241:Neofunctionalization 120:Replication slippage 115:Replication slippage 79:replication slippage 4763:Molecular evolution 4631:Model lipid bilayer 4473:Interspersed repeat 3369:Molecular processes 3294:Balancing selection 3278:Molecular evolution 3180:2023Natur.615..925L 3115:2021Natur.590..492Q 2816:2005PNAS..102..707G 2634:10.1038/nature07092 2626:2008Natur.454..762D 2565:1999JMolE..49..169S 2473:10.1038/nature02424 2465:2004Natur.428..617K 2430:. Springer-Verlag. 2122:1981PNAS...78.3113A 2018:2008Sci...320.1629E 1926:2004Sci...305..525S 1863:2010PNAS..107.2967A 1603:2000Sci...290.1151L 1597:(5494): 1151–1155. 1548:10.1038/nature01771 1540:2003Natur.424..194P 1369:10.1038/nature02424 1361:2004Natur.428..617K 1296:1997Natur.387..708W 1070:Molecular evolution 612: 494:level. It shows 22 400:divergent evolution 43:molecular evolution 3941:organic structures 3820:Gene amplification 3309:Negative selection 2777:10.1002/mrdd.20082 2583:10.1007/PL00006540 2230:10.1038/ng.2007.40 1801:Trends in Genetics 1695:Trends in Genetics 1659:10.1101/gr.3681406 1030:De novo gene birth 610: 552:gene amplification 548:molecular genetics 517: 509:(at bottom left). 347:Rett-like syndrome 285:genome duplication 253:selective pressure 211: 131:Retrotransposition 112: 67:retrotransposition 39:gene amplification 4753:Genetics concepts 4735: 4734: 4676:Non-cellular life 4483: 4482: 4222: 4221: 4195: 4194: 4149:ssRNA satellite ( 3905: 3904: 3806: 3805: 3674: 3673: 3578: 3577: 3467:Repeated sequence 3442:repeated sequence 3404: 3403: 3286:Natural selection 3174:(7954): 925–933. 3109:(7846): 492–497. 3024:978-0-07-137050-9 2437:978-91-554-5776-1 2422:Ohno, S. (1967). 2408:978-0-04-575015-3 2012:(5883): 1629–31. 1534:(6945): 194–197. 1355:(6983): 617–624. 1290:(6634): 708–713. 1004: 1003: 761:Esophageal cancer 726:Colorectal cancer 505:, as well as the 16:(Redirected from 4775: 4412:Gene duplication 4235: 4234: 4231:self-replication 4119: 4118: 4081: 4080: 3939:Self-replicating 3932: 3925: 3918: 3909: 3908: 3882:Low copy repeats 3875:Symbiosis island 3812:Gene duplication 3598: 3597: 3589: 3588: 3472: 3471: 3450:gene duplication 3431: 3424: 3417: 3408: 3407: 3381:Gene duplication 3345:Allele frequency 3271: 3264: 3257: 3248: 3247: 3220: 3219: 3209: 3191: 3159: 3153: 3152: 3142: 3093: 3087: 3086: 3076: 3051:(8): 1189–1195. 3035: 3029: 3028: 3008: 2945: 2944: 2934: 2924: 2900: 2894: 2892: 2878: 2872: 2871: 2869: 2868: 2854: 2848: 2847: 2837: 2827: 2795: 2789: 2788: 2760: 2754: 2753: 2743: 2733: 2709: 2703: 2702: 2684: 2660: 2654: 2653: 2609: 2603: 2602: 2576: 2548: 2542: 2541: 2531: 2499: 2493: 2492: 2459:(6983): 617–24. 2448: 2442: 2441: 2429: 2419: 2413: 2412: 2387: 2381: 2380: 2352: 2346: 2345: 2309: 2303: 2302: 2292: 2282: 2258: 2252: 2251: 2241: 2209: 2203: 2202: 2192: 2160: 2154: 2153: 2143: 2133: 2101: 2095: 2094: 2084: 2052: 2046: 2045: 2001: 1995: 1994: 1984: 1960: 1954: 1953: 1909: 1903: 1902: 1892: 1874: 1857:(7): 2967–2971. 1842: 1833: 1832: 1792: 1786: 1785: 1775: 1757: 1733: 1727: 1726: 1686: 1680: 1679: 1661: 1637: 1631: 1630: 1582: 1576: 1575: 1519: 1513: 1512: 1502: 1484: 1460: 1454: 1453: 1427: 1403: 1397: 1396: 1340: 1334: 1333: 1307: 1275: 1269: 1268: 1258: 1248: 1224: 1218: 1217: 1207: 1189: 1165: 1159: 1158: 1156: 1155: 1136: 1130: 1129: 1111: 1105:Zhang J (2003). 1102: 613: 609: 595:Duplications of 583:, rather than a 542:As amplification 525:human chromosome 415:low copy repeats 324:" (a process of 173:is a product of 136:Retrotransposons 49:that contains a 31:Gene duplication 21: 4783: 4782: 4778: 4777: 4776: 4774: 4773: 4772: 4738: 4737: 4736: 4731: 4681:Synthetic virus 4669:Artificial cell 4642: 4570: 4479: 4368:RNA replication 4363:DNA replication 4351: 4342:Group II intron 4240: 4230: 4218: 4209:Mammalian prion 4191: 4167: 4146:dsRNA satellite 4143:ssDNA satellite 4113: 4106: 4075: 4068: 4013: 3942: 3936: 3906: 3901: 3853: 3802: 3670: 3642: 3619: 3593:Retrotransposon 3574: 3565:Inverted repeat 3553: 3538:DNA transposon 3534:Retrotransposon 3529:Gene conversion 3520: 3513: 3510: 3461: 3452: 3435: 3405: 3400: 3386:Silent mutation 3376:Gene conversion 3364: 3323: 3319:Selective sweep 3280: 3275: 3229: 3224: 3223: 3160: 3156: 3094: 3090: 3045:Nature Genetics 3036: 3032: 3025: 3009: 2948: 2915:(1): e1009996. 2901: 2897: 2879: 2875: 2866: 2864: 2856: 2855: 2851: 2796: 2792: 2761: 2757: 2710: 2706: 2661: 2657: 2620:(7205): 762–5. 2610: 2606: 2574:10.1.1.466.5042 2549: 2545: 2500: 2496: 2449: 2445: 2438: 2420: 2416: 2409: 2399:Springer-Verlag 2388: 2384: 2353: 2349: 2326:10.1038/nrg2482 2310: 2306: 2259: 2255: 2218:Nature Genetics 2210: 2206: 2161: 2157: 2102: 2098: 2061:Current Biology 2053: 2049: 2002: 1998: 1969:Nature Genetics 1961: 1957: 1920:(5683): 525–8. 1910: 1906: 1843: 1836: 1807:(11): 602–607. 1793: 1789: 1734: 1730: 1701:(10): 548–551. 1687: 1683: 1646:Genome Research 1638: 1634: 1583: 1579: 1520: 1516: 1461: 1457: 1404: 1400: 1341: 1337: 1276: 1272: 1225: 1221: 1166: 1162: 1153: 1151: 1138: 1137: 1133: 1109: 1103: 1099: 1094: 1089: 1014: 1006: 701:Cervical cancer 629: 627: 625: 620: 619:Associated gene 593: 544: 515: 503:sex chromosomes 476: 467: 455:gene regulation 435: 419:pericentronomic 404:gene conversion 368: 363: 343: 318: 312: 249: 243: 216: 203: 164: 148: 133: 117: 92: 87: 55:DNA replication 28: 23: 22: 15: 12: 11: 5: 4781: 4771: 4770: 4765: 4760: 4755: 4750: 4733: 4732: 4730: 4729: 4724: 4723: 4722: 4717: 4707: 4701: 4695: 4694: 4693: 4688: 4678: 4673: 4672: 4671: 4666: 4656: 4650: 4648: 4644: 4643: 4641: 4640: 4639: 4638: 4633: 4625: 4620: 4615: 4610: 4604: 4603: 4602: 4591: 4586: 4580: 4578: 4572: 4571: 4569: 4568: 4563: 4562: 4561: 4556: 4548: 4546:Kappa organism 4543: 4542: 4541: 4536: 4531: 4526: 4521: 4511: 4510: 4509: 4504: 4493: 4491: 4485: 4484: 4481: 4480: 4478: 4477: 4476: 4475: 4470: 4460: 4459: 4458: 4453: 4448: 4443: 4433: 4432: 4431: 4421: 4420: 4419: 4417:Non-coding DNA 4414: 4409: 4399: 4398: 4397: 4392: 4387: 4382: 4372: 4371: 4370: 4359: 4357: 4353: 4352: 4350: 4349: 4344: 4339: 4337:Group I intron 4334: 4329: 4328: 4327: 4317: 4316: 4315: 4312: 4303: 4300: 4295: 4290: 4280: 4279: 4278: 4273: 4263: 4262: 4261: 4259:Genomic island 4256: 4245: 4243: 4239:Mobile genetic 4232: 4224: 4223: 4220: 4219: 4217: 4216: 4211: 4205: 4203: 4197: 4196: 4193: 4192: 4190: 4189: 4188: 4187: 4184: 4175: 4173: 4169: 4168: 4166: 4165: 4164: 4163: 4160: 4154: 4147: 4144: 4141: 4138: 4131: 4127: 4125: 4116: 4108: 4107: 4105: 4104: 4097: 4089: 4087: 4078: 4070: 4069: 4067: 4066: 4064:dsDNA-RT virus 4061: 4059:ssRNA-RT virus 4056: 4054:(−)ssRNA virus 4051: 4049:(+)ssRNA virus 4046: 4041: 4036: 4035: 4034: 4023: 4021: 4015: 4014: 4012: 4011: 4010: 4009: 4004: 3994:Incertae sedis 3990: 3989: 3988: 3983: 3978: 3973: 3963: 3958: 3952: 3950: 3944: 3943: 3935: 3934: 3927: 3920: 3912: 3903: 3902: 3900: 3899: 3894: 3889: 3884: 3879: 3878: 3877: 3872: 3865:Genomic island 3861: 3859: 3855: 3854: 3852: 3851: 3846: 3845: 3844: 3834: 3833: 3832: 3822: 3816: 3814: 3808: 3807: 3804: 3803: 3801: 3800: 3795: 3790: 3785: 3780: 3775: 3770: 3765: 3760: 3755: 3750: 3745: 3740: 3735: 3730: 3725: 3720: 3715: 3710: 3705: 3700: 3695: 3690: 3684: 3682: 3680:DNA transposon 3676: 3675: 3672: 3671: 3669: 3668: 3663: 3658: 3652: 3650: 3644: 3643: 3641: 3640: 3635: 3629: 3627: 3621: 3620: 3618: 3617: 3612: 3606: 3604: 3595: 3586: 3580: 3579: 3576: 3575: 3573: 3572: 3567: 3561: 3559: 3555: 3554: 3552: 3551: 3550: 3549: 3544: 3536: 3531: 3525: 3523: 3515: 3514: 3512: 3511: 3508:Macrosatellite 3505: 3495: 3486: 3480: 3478: 3476:Tandem repeats 3469: 3463: 3462: 3457: 3454: 3453: 3434: 3433: 3426: 3419: 3411: 3402: 3401: 3399: 3398: 3393: 3388: 3383: 3378: 3372: 3370: 3366: 3365: 3363: 3362: 3360:Fay and Wu's H 3357: 3352: 3347: 3342: 3337: 3331: 3329: 3325: 3324: 3322: 3321: 3316: 3311: 3306: 3301: 3296: 3290: 3288: 3282: 3281: 3274: 3273: 3266: 3259: 3251: 3245: 3244: 3237: 3228: 3227:External links 3225: 3222: 3221: 3154: 3088: 3030: 3023: 2946: 2895: 2873: 2849: 2790: 2755: 2704: 2655: 2604: 2543: 2514:(4): 1531–45. 2494: 2443: 2436: 2414: 2407: 2382: 2347: 2320:(12): 938–50. 2304: 2253: 2204: 2175:(3): 1035–43. 2155: 2096: 2047: 1996: 1982:10.1038/ng1416 1955: 1904: 1834: 1787: 1742:Genome Biology 1728: 1681: 1652:(7): 805–814. 1632: 1577: 1514: 1455: 1418:(3): 452–462. 1398: 1335: 1270: 1219: 1160: 1131: 1096: 1095: 1093: 1090: 1088: 1087: 1082: 1077: 1072: 1067: 1062: 1057: 1052: 1047: 1042: 1037: 1035:Exon shuffling 1032: 1027: 1021: 1015: 1013: 1010: 1002: 1001: 998: 993: 987: 986: 983: 977: 976: 973: 968: 962: 961: 958: 952: 951: 948: 942: 941: 938: 933: 931:Ovarian cancer 927: 926: 923: 918: 912: 911: 908: 903: 897: 896: 893: 887: 886: 883: 877: 876: 873: 868: 862: 861: 858: 852: 851: 848: 838: 832: 831: 828: 822: 821: 818: 812: 811: 808: 798: 796:Gastric cancer 792: 791: 788: 782: 781: 778: 772: 771: 768: 763: 757: 756: 753: 747: 746: 743: 737: 736: 733: 728: 722: 721: 718: 712: 711: 708: 703: 697: 696: 693: 687: 686: 683: 677: 676: 673: 662: 661: 658: 647: 646: 643: 638: 632: 631: 628:in cancer type 626:amplification 624:Prevalence of 622: 621:amplifications 617: 592: 591:Role in cancer 589: 543: 540: 475: 472: 466: 463: 434: 431: 367: 364: 362: 359: 342: 339: 314:Main article: 311: 308: 245:Main article: 242: 239: 215: 212: 202: 199: 175:nondisjunction 163: 160: 147: 144: 132: 129: 124:DNA polymerase 116: 113: 91: 88: 86: 83: 26: 9: 6: 4: 3: 2: 4780: 4769: 4766: 4764: 4761: 4759: 4756: 4754: 4751: 4749: 4746: 4745: 4743: 4728: 4725: 4721: 4718: 4716: 4713: 4712: 4711: 4708: 4706: 4702: 4700: 4699:Nanobacterium 4696: 4692: 4689: 4687: 4684: 4683: 4682: 4679: 4677: 4674: 4670: 4667: 4665: 4664:Cell division 4662: 4661: 4660: 4657: 4655: 4652: 4651: 4649: 4645: 4637: 4634: 4632: 4629: 4628: 4626: 4624: 4621: 4619: 4616: 4614: 4611: 4609: 4605: 4601: 4598: 4597: 4596: 4592: 4590: 4587: 4585: 4582: 4581: 4579: 4577: 4573: 4567: 4564: 4560: 4557: 4555: 4552: 4551: 4549: 4547: 4544: 4540: 4537: 4535: 4532: 4530: 4527: 4525: 4522: 4520: 4517: 4516: 4515: 4512: 4508: 4507:Hydrogenosome 4505: 4503: 4500: 4499: 4498: 4497:Mitochondrion 4495: 4494: 4492: 4490: 4489:Endosymbiosis 4486: 4474: 4471: 4469: 4468:Tandem repeat 4466: 4465: 4464: 4461: 4457: 4454: 4452: 4449: 4447: 4444: 4442: 4439: 4438: 4437: 4434: 4430: 4427: 4426: 4425: 4422: 4418: 4415: 4413: 4410: 4408: 4405: 4404: 4403: 4400: 4396: 4393: 4391: 4388: 4386: 4383: 4381: 4378: 4377: 4376: 4373: 4369: 4366: 4365: 4364: 4361: 4360: 4358: 4356:Other aspects 4354: 4348: 4345: 4343: 4340: 4338: 4335: 4333: 4330: 4326: 4323: 4322: 4321: 4318: 4313: 4311: 4307: 4304: 4301: 4299: 4296: 4294: 4291: 4289: 4286: 4285: 4284: 4281: 4277: 4274: 4272: 4269: 4268: 4267: 4264: 4260: 4257: 4255: 4252: 4251: 4250: 4247: 4246: 4244: 4242: 4236: 4233: 4229: 4225: 4215: 4212: 4210: 4207: 4206: 4204: 4202: 4198: 4185: 4182: 4181: 4180: 4177: 4176: 4174: 4170: 4161: 4158: 4157: 4155: 4152: 4148: 4145: 4142: 4139: 4136: 4132: 4129: 4128: 4126: 4124: 4120: 4117: 4115: 4109: 4103: 4102: 4101:Avsunviroidae 4098: 4096: 4095: 4094:Pospiviroidae 4091: 4090: 4088: 4086: 4082: 4079: 4077: 4071: 4065: 4062: 4060: 4057: 4055: 4052: 4050: 4047: 4045: 4042: 4040: 4037: 4033: 4030: 4029: 4028: 4025: 4024: 4022: 4020: 4016: 4008: 4005: 4003: 4002: 3998: 3997: 3996: 3995: 3991: 3987: 3984: 3982: 3979: 3977: 3974: 3972: 3969: 3968: 3967: 3964: 3962: 3959: 3957: 3954: 3953: 3951: 3949: 3948:Cellular life 3945: 3940: 3933: 3928: 3926: 3921: 3919: 3914: 3913: 3910: 3898: 3895: 3893: 3890: 3888: 3885: 3883: 3880: 3876: 3873: 3871: 3868: 3867: 3866: 3863: 3862: 3860: 3856: 3850: 3847: 3843: 3840: 3839: 3838: 3835: 3831: 3830:Ribosomal DNA 3828: 3827: 3826: 3823: 3821: 3818: 3817: 3815: 3813: 3809: 3799: 3796: 3794: 3791: 3789: 3786: 3784: 3781: 3779: 3776: 3774: 3771: 3769: 3766: 3764: 3761: 3759: 3756: 3754: 3751: 3749: 3746: 3744: 3741: 3739: 3736: 3734: 3731: 3729: 3726: 3724: 3721: 3719: 3716: 3714: 3711: 3709: 3706: 3704: 3701: 3699: 3696: 3694: 3691: 3689: 3686: 3685: 3683: 3681: 3677: 3667: 3664: 3662: 3659: 3657: 3654: 3653: 3651: 3649: 3645: 3639: 3636: 3634: 3631: 3630: 3628: 3626: 3622: 3616: 3613: 3611: 3608: 3607: 3605: 3603: 3599: 3596: 3594: 3590: 3587: 3585: 3581: 3571: 3570:Direct repeat 3568: 3566: 3563: 3562: 3560: 3556: 3548: 3545: 3543: 3540: 3539: 3537: 3535: 3532: 3530: 3527: 3526: 3524: 3522: 3516: 3509: 3506: 3503: 3499: 3496: 3494: 3493:Minisatellite 3490: 3487: 3485: 3484:Satellite DNA 3482: 3481: 3479: 3477: 3473: 3470: 3468: 3464: 3460: 3455: 3451: 3447: 3443: 3439: 3432: 3427: 3425: 3420: 3418: 3413: 3412: 3409: 3397: 3394: 3392: 3389: 3387: 3384: 3382: 3379: 3377: 3374: 3373: 3371: 3367: 3361: 3358: 3356: 3353: 3351: 3348: 3346: 3343: 3341: 3338: 3336: 3333: 3332: 3330: 3326: 3320: 3317: 3315: 3312: 3310: 3307: 3305: 3302: 3300: 3297: 3295: 3292: 3291: 3289: 3287: 3283: 3279: 3272: 3267: 3265: 3260: 3258: 3253: 3252: 3249: 3243: 3242: 3238: 3236: 3235: 3231: 3230: 3217: 3213: 3208: 3203: 3199: 3195: 3190: 3185: 3181: 3177: 3173: 3169: 3165: 3158: 3150: 3146: 3141: 3136: 3132: 3128: 3124: 3120: 3116: 3112: 3108: 3104: 3100: 3092: 3084: 3080: 3075: 3070: 3066: 3062: 3058: 3054: 3050: 3046: 3042: 3034: 3026: 3020: 3016: 3015: 3007: 3005: 3003: 3001: 2999: 2997: 2995: 2993: 2991: 2989: 2987: 2985: 2983: 2981: 2979: 2977: 2975: 2973: 2971: 2969: 2967: 2965: 2963: 2961: 2959: 2957: 2955: 2953: 2951: 2942: 2938: 2933: 2928: 2923: 2918: 2914: 2910: 2909:PLOS Genetics 2906: 2899: 2890: 2889: 2884: 2877: 2863: 2859: 2853: 2845: 2841: 2836: 2831: 2826: 2821: 2817: 2813: 2810:(3): 707–12. 2809: 2805: 2801: 2794: 2786: 2782: 2778: 2774: 2771:(4): 279–85. 2770: 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Index

Amplification (molecular biology)
molecular evolution
DNA
gene
DNA replication
repair
ectopic recombination
retrotransposition
aneuploidy
polyploidy
replication slippage
unequal crossing-over
transposable

Replication slippage
DNA polymerase
Retrotransposons
L1
Aneuploidy
Down syndrome
Polyploidy
nondisjunction
2R event
Hox genes

copy number variation
C. elegans
Neofunctionalization
selective pressure
mutations

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