27:
482:
857:
245:
401:
341:
886:
118:
626:
261:
424:, and thus the gene may become less- or non-functional or "deactivated". These are the same mechanisms by which non-processed genes become pseudogenes, but the difference in this case is that the gene was not duplicated before pseudogenization. Normally, such a pseudogene would be unlikely to become fixed in a population, but various population effects, such as
375:, since an intact functional copy still exists. According to some evolutionary models, shared duplicated pseudogenes indicate the evolutionary relatedness of humans and the other primates. If pseudogenization is due to gene duplication, it usually occurs in the first few million years after the gene duplication, provided the gene has not been subjected to any
324:. However, because they are derived from an RNA product, processed pseudogenes also lack the upstream promoters of normal genes; thus, they are considered "dead on arrival", becoming non-functional pseudogenes immediately upon the retrotransposition event. However, these insertions occasionally contribute exons to existing genes, usually via
649:
normally. However, when BRAFP1 RNA expression is increased (either experimentally or by natural mutations), less miRNA is available to control the expression of BRAF, and the increased amount of BRAF protein causes cancer. This sort of competition for regulatory elements by RNAs that are endogenous to the genome has given rise to the term
328:
transcripts. A further characteristic of processed pseudogenes is common truncation of the 5' end relative to the parent sequence, which is a result of the relatively non-processive retrotransposition mechanism that creates processed pseudogenes. Processed pseudogenes are continually being created in
648:
that, when mutated, is associated with many cancers. Normally, the amount of BRAF protein is kept under control in cells through the action of miRNA. In normal situations, the amount of RNA from BRAF and the pseudogene BRAFP1 compete for miRNA, but the balance of the 2 RNAs is such that cells grow
690:
due to its similarity to the PTEN gene, and overexpression of the 3' UTR resulted in an increase of PTEN protein level. That is, overexpression of the PTENP1 3' UTR leads to increased regulation and suppression of cancerous tumors. The biology of this system is basically the inverse of the BRAF
79:
Most non-bacterial genomes contain many pseudogenes, often as many as functional genes. This is not surprising, since various biological processes are expected to accidentally create pseudogenes, and there are no specialized mechanisms to remove them from genomes. Eventually pseudogenes may be
160:
sequence that is similar to some functional gene, they are usually unable to produce functional final protein products. Pseudogenes are sometimes difficult to identify and characterize in genomes, because the two requirements of similarity and loss of functionality are usually implied through
383:
and it is not normally advantageous to carry two identical genes. Mutations that disrupt either the structure or the function of either of the two genes are not deleterious and will not be removed through the selection process. As a result, the gene that has been mutated gradually becomes a
603:
appear to be derived from pseudogenes, and thus some pseudogenes play a role in regulating protein-coding transcripts, as reviewed. One of the many examples is psiPPM1K. Processing of RNAs transcribed from psiPPM1K yield siRNAs that can act to suppress the most common type of liver cancer,
103:
elements inherited from the ancestral gene or arising by new mutations. Although most of these transcripts will have no more functional significance than chance transcripts from other parts of the genome, some have given rise to beneficial regulatory RNAs and new proteins.
392:. According to evolutionary context, these pseudogenes will either be deleted or become so distinct from the parental genes so that they will no longer be identifiable. Relatively young pseudogenes can be recognized due to their sequence similarity.
1443:
Pavlícek A, Paces J, Zíka R, Hejnar J (October 2002). "Length distribution of long interspersed nucleotide elements (LINEs) and processed pseudogenes of human endogenous retroviruses: implications for retrotransposition and pseudogene detection".
173:
is computed. A high sequence identity means that it is highly likely that these two sequences diverged from a common ancestral sequence (are homologous), and highly unlikely that these two sequences have evolved independently (see
517:
of such premature stop codons in mammals. As alluded to in the figure above, a small amount of the protein product of such readthrough may still be recognizable and function at some level. If so, the pseudogene can be subject to
580:
of peptides identified at least 19,262 human proteins produced from 16,271 genes or clusters of genes, with 8 new protein-coding genes identified that were previously considered pseudogenes. An earlier analysis found that human
197:
are all required parts of this process. If any of these steps fails, then the sequence may be considered nonfunctional. In high-throughput pseudogene identification, the most commonly identified disablements are premature
214:
genes are usually more difficult to discover as they do not need to be translated and thus do not have "reading frames". A number of rRNA pseudogenes have been identified on the basis of changes in rDNA array ends.
538:
species actually encode proteins with biologically important functions, "suggesting that such 'pseudo-pseudogenes' could represent a widespread phenomenon". For example, the functional protein (a glutamate
460:
While the vast majority of pseudogenes have lost their function, some cases have emerged in which a pseudogene either re-gained its original or a similar function or evolved a new function. In the
233:
programs, often being misidentified as real genes or exons. It has been proposed that the identification of processed pseudogenes can help improve the accuracy of gene prediction methods.
739:. Thus, they do not require many genes that are needed by free-living bacteria, such as gene associated with metabolism and DNA repair. However, there is not an order to which functional
819:
Although genome reduction focuses on what genes are not needed by getting rid of pseudogenes, selective pressures from the host can sway what is kept. In the case of a symbiont from the
670:. The PTEN pseudogene, PTENP1 is a processed pseudogene that is very similar in its genetic sequence to the wild-type gene. However, PTENP1 has a missense mutation which eliminates the
464:, a number of examples have been identified that were originally classified as pseudogenes but later discovered to have a functional, although not necessarily protein-coding, role.
222:
may simultaneously amplify a pseudogene that shares similar sequences. This is known as PCR bias or amplification bias. Similarly, pseudogenes are sometimes annotated as genes in
840:. The domino theory suggests that if one gene of a cellular process becomes inactivated, then selection in other genes involved relaxes, leading to gene loss. When comparing
3737:
2401:
Dierick HA, Mercer JF, Glover TW (October 1997). "A phosphoglycerate mutase brain isoform (PGAM 1) pseudogene is localized within the human Menkes disease gene (ATP7 A)".
588:
A number of pseudo-pseudogenes were also found in prokaryotes, where some stop codon substitutions in essential genes appear to be retained, even positively selected for.
784:
Since most bacteria that carry pseudogenes are either symbionts or obligate intracellular parasites, genome size eventually reduces. An extreme example is the genome of
92:
changes that they are no longer recognizable as former genes. Analysis of these degeneration events helps clarify the effects of non-selective processes in genomes.
444:(vitamin C), but it exists as a disabled gene (GULOP) in humans and other primates. Another more recent example of a disabled gene links the deactivation of the
252:
There are four main types of pseudogenes, all with distinct mechanisms of origin and characteristic features. The classifications of pseudogenes are as follows:
1916:"Cloning and chromosomal mapping of the human nonfunctional gene for L-gulono-gamma-lactone oxidase, the enzyme for L-ascorbic acid biosynthesis missing in man"
3794:
351:
is another common and important process in the evolution of genomes. A copy of a functional gene may arise as a result of a gene duplication event caused by
3819:
3769:
618:
are derived from pseudogenes located in piRNA clusters. Those piRNAs regulate genes via the piRNA pathway in mammalian testes and are crucial for limiting
312:
system that they can create retrotransposed copies of random genes, too. Once these pseudogenes are inserted back into the genome, they usually contain a
3829:
363:
that cause the copy to lose the original gene's function. Duplicated pseudogenes usually have all the same characteristics as genes, including an intact
3951:
3869:
1875:"Guinea pigs possess a highly mutated gene for L-gulono-gamma-lactone oxidase, the key enzyme for L-ascorbic acid biosynthesis missing in this species"
3859:
3774:
56:
transcript. Pseudogenes are usually identified when genome sequence analysis finds gene-like sequences that lack regulatory sequences needed for
3234:
Cole ST, Eiglmeier K, Parkhill J, James KD, Thomson NR, Wheeler PR, et al. (February 2001). "Massive gene decay in the leprosy bacillus".
3779:
608:. This and much other research has led to considerable excitement about the possibility of targeting pseudogenes with/as therapeutic agents
2589:""Pseudo-pseudogenes" in bacterial genomes: Proteogenomics reveals a wide but low protein expression of pseudogenes in Salmonella enterica"
3505:
2479:"A single nucleotide polymorphism within the novel sex-linked testis-specific retrotransposed PGAM4 gene influences human male fertility"
3092:"The "domino theory" of gene death: gradual and mass gene extinction events in three lineages of obligate symbiotic bacterial pathogens"
308:
back into DNA and inserted into chromosomal DNA. Although retrotransposons usually create copies of themselves, it has been shown in an
3874:
280:
is a fairly common event that has had a huge impact on the composition of the genome. For example, somewhere between 30 and 44% of the
3839:
3789:
3784:
2250:
Long M, Langley CH (April 1993). "Natural selection and the origin of jingwei, a chimeric processed functional gene in
Drosophila".
236:
In 2014, 140 human pseudogenes have been shown to be translated. However, the function, if any, of the protein products is unknown.
3814:
3764:
2211:"The origin of the Jingwei gene and the complex modular structure of its parental gene, yellow emperor, in Drosophila melanogaster"
585:(phosphoglycerate mutase), previously thought to be a pseudogene, is not only functional, but also causes infertility if mutated.
3809:
3714:
2004:
Cheetham SW, Faulkner GJ, Dinger ME (March 2020). "Overcoming challenges and dogmas to understand the functions of pseudogenes".
582:
181:
Nonfunctionality can manifest itself in many ways. Normally, a gene must go through several steps to a fully functional protein:
3472:
3834:
2794:
2655:
371:
structure and regulatory sequences. The loss of a duplicated gene's functionality usually has little effect on an organism's
2438:"Evolution of the phosphoglycerate mutase processed gene in human and chimpanzee revealing the origin of a new primate gene"
1341:
Graur D, Shuali Y, Li WH (April 1989). "Deletions in processed pseudogenes accumulate faster in rodents than in humans".
2681:"Transcribed pseudogene ψPPM1K generates endogenous siRNA to suppress oncogenic cell growth in hepatocellular carcinoma"
3691:
3678:
926:
3701:
1255:
Dewannieux M, Heidmann T (2005). "LINEs, SINEs and processed pseudogenes: parasitic strategies for genome modeling".
1198:
440:(GULO) in primates. In all mammals studied besides primates (except guinea pigs), GULO aids in the biosynthesis of
2818:
Siomi MC, Sato K, Pezic D, Aravin AA (April 2011). "PIWI-interacting small RNAs: the vanguard of genome defence".
1298:
Dewannieux M, Esnault C, Heidmann T (September 2003). "LINE-mediated retrotransposition of marked Alu sequences".
701:
and other mutational events that may give rise to new or newly functional genes. This has led to the concept that
547:. This finding of tissue-specific biologically-functional genes that could have been classified as pseudogenes by
3498:
736:
436:, can lead to fixation. The classic example of a unitary pseudogene is the gene that presumably coded the enzyme
384:
pseudogene and will be either unexpressed or functionless. This kind of evolutionary fate is shown by population
1799:"Molecular fossils in the human genome: identification and analysis of the pseudogenes in chromosomes 21 and 22"
876:
Homologous genes are indicated by identical colors and thin blue vertical bars. Modified after Cole et al. 2001.
165:
Homology is implied by sequence similarity between the DNA sequences of the pseudogene and a known gene. After
329:
primates. Human populations, for example, have distinct sets of processed pseudogenes across its individuals.
3732:
3578:
3565:
825:
phylum, there are seven additional copies of the gene coding the mandelalide pathway. The host, species from
678:
and thus prevents translation of the normal PTEN protein. In spite of that, PTENP1 appears to play a role in
640:
decoys. Perhaps the earliest definitive example of such a pseudogene involved in cancer is the pseudogene of
3483:
572:
As of 2012, it appeared that there are approximately 12,000–14,000 pseudogenes in the human genome. A 2016
3462:
332:
It has been shown that processed pseudogenes accumulate mutations faster than non-processed pseudogenes.
3467:
1797:
Harrison PM, Hegyi H, Balasubramanian S, Luscombe NM, Bertone P, Echols N, et al. (February 2002).
3849:
3491:
816:
because it can survive outside the host; therefore, the genome must contain the genes needed to do so.
356:
2538:"Analysis of Stop Codons within Prokaryotic Protein-Coding Genes Suggests Frequent Readthrough Events"
219:
1581:"Pseudogenes in the ENCODE regions: consensus annotation, analysis of transcription, and evolution"
1528:
Schrider DR, Navarro FC, Galante PA, Parmigiani RB, Camargo AA, Hahn MW, de Souza SJ (2013-01-24).
605:
352:
182:
3463:
Pseudogene interaction database, miRNA-pseudogene and protein-pseudogene interaction maps database
3999:
3543:
3518:
2101:
Prieto-Godino LL, Rytz R, Bargeton B, Abuin L, Arguello JR, Peraro MD, Benton R (November 2016).
1656:
Lynch M, Conery JS (November 2000). "The evolutionary fate and consequences of duplicate genes".
1175:
1021:"Ribosomal RNA Genes Contribute to the Formation of Pseudogenes and Junk DNA in the Human Genome"
514:
486:
437:
417:
57:
3973:
3901:
936:
675:
3369:
Bischof JM, Chiang AP, Scheetz TE, Stone EM, Casavant TL, Sheffield VC, Braun TA (June 2006).
218:
Pseudogenes can complicate molecular genetic studies. For example, amplification of a gene by
2346:"Improving GENCODE reference gene annotation using a high-stringency proteogenomics workflow"
941:
853:
it was found that positive epistasis furthers gene loss while negative epistasis hinders it.
800:
794:. It has been reported to have 1,133 pseudogenes which give rise to approximately 50% of its
667:
562:
506:
429:
421:
305:
190:
61:
49:
3044:
2863:"The BRAF pseudogene functions as a competitive endogenous RNA and induces lymphoma in vivo"
3994:
3946:
3660:
3522:
3298:
3243:
3021:
2964:
2490:
2357:
2344:
Wright JC, Mudge J, Weisser H, Barzine MP, Gonzalez JM, Brazma A, et al. (June 2016).
2259:
2167:
2114:
1665:
1350:
1130:
786:
758:
745:
619:
615:
325:
175:
20:
2777:
Olovnikov I, Le Thomas A, Aravin AA (2014). "A Framework for piRNA Cluster
Manipulation".
2454:
2437:
2227:
2210:
1579:
Zheng D, Frankish A, Baertsch R, Kapranov P, Reymond A, Choo SW, et al. (June 2007).
8:
3724:
3595:
1633:
931:
842:
683:
317:
203:
186:
156:
to a known gene and loss of some functionality. That is, although every pseudogene has a
100:
65:
3302:
3247:
2968:
2910:
Dahia PL, FitzGerald MG, Zhang X, Marsh DJ, Zheng Z, Pietsch T, et al. (May 1998).
2494:
2361:
2263:
2171:
2118:
2065:
1957:"Spread of an inactive form of caspase-12 in humans is due to recent positive selection"
1669:
1354:
1134:
1117:
Kim MS, Pinto SM, Getnet D, Nirujogi RS, Manda SS, Chaerkady R, et al. (May 2014).
899:
Please expand the article to include this information. Further details may exist on the
131:
Please expand the section to include this information. Further details may exist on the
3896:
3804:
3623:
3438:
3413:
3400:
3267:
3211:
3186:
3159:
3132:
2985:
2952:
2887:
2862:
2843:
2754:
2729:
2705:
2680:
2615:
2588:
2564:
2537:
2513:
2478:
2378:
2345:
2321:
2294:
2191:
2135:
2102:
2075:
2048:
2029:
1981:
1956:
1774:
1749:
1725:
1700:
1605:
1580:
1556:
1529:
1505:
1480:
1420:
1393:
1374:
1323:
1280:
1151:
1118:
1094:
1069:
1045:
1020:
1001:
540:
481:
376:
289:
285:
277:
166:
3371:"Genome-wide identification of pseudogenes capable of disease-causing gene conversion"
3352:
3327:
3310:
2861:
Karreth FA, Reschke M, Ruocco A, Ng C, Chapuy B, Léopold V, et al. (April 2015).
2477:
Okuda H, Tsujimura A, Irie S, Yamamoto K, Fukuhara S, Matsuoka Y, et al. (2012).
2414:
2293:
Pei B, Sisu C, Frankish A, Howald C, Habegger L, Mu XJ, et al. (September 2012).
1932:
1915:
1891:
1874:
1859:
1823:
1798:
1457:
988:
971:
798:. The effect of pseudogenes and genome reduction can be further seen when compared to
3443:
3392:
3357:
3314:
3259:
3216:
3187:"Increased Biosynthetic Gene Dosage in a Genome-Reduced Defensive Bacterial Symbiont"
3164:
3113:
3067:
3025:
2990:
2953:"A coding-independent function of gene and pseudogene mRNAs regulates tumour biology"
2933:
2892:
2835:
2800:
2790:
2759:
2710:
2661:
2651:
2638:
Chan WL, Chang JG (2014). "Pseudogene-Derived
Endogenous siRNAs and Their Function".
2620:
2569:
2518:
2459:
2418:
2383:
2326:
2275:
2232:
2183:
2140:
2080:
2033:
2021:
1986:
1937:
1896:
1828:
1779:
1730:
1681:
1610:
1561:
1510:
1461:
1425:
1366:
1315:
1272:
1237:
1156:
1099:
1050:
993:
821:
763:
577:
519:
449:
433:
413:
372:
153:
3404:
1378:
1327:
1284:
1005:
3958:
3888:
3526:
3433:
3425:
3382:
3347:
3339:
3306:
3271:
3251:
3206:
3198:
3154:
3144:
3103:
3059:
3017:
2980:
2972:
2923:
2882:
2874:
2847:
2827:
2782:
2749:
2741:
2700:
2692:
2643:
2610:
2600:
2559:
2549:
2508:
2498:
2449:
2410:
2373:
2365:
2316:
2306:
2267:
2222:
2195:
2175:
2130:
2122:
2070:
2060:
2013:
1976:
1968:
1927:
1886:
1855:
1818:
1810:
1769:
1761:
1720:
1712:
1673:
1600:
1592:
1551:
1541:
1500:
1492:
1453:
1415:
1405:
1358:
1307:
1264:
1229:
1146:
1138:
1089:
1081:
1040:
1032:
983:
848:
348:
293:
206:, which almost universally prevent the translation of a functional protein product.
45:
3008:
Balakirev ES, Ayala FJ (2003). "Pseudogenes: are they "junk" or functional DNA?".
1765:
1677:
19:
This article is about nonfunctional segments of DNA. For a species of beetle, see
3669:
3641:
3610:
3605:
3149:
2912:"A highly conserved processed PTEN pseudogene is located on chromosome band 9p21"
2647:
2503:
1716:
1546:
1220:
Jurka J (December 2004). "Evolutionary impact of human Alu repetitive elements".
951:
900:
774:
698:
498:
389:
380:
313:
230:
194:
132:
81:
2786:
2679:
Chan WL, Yuo CY, Yang WK, Hung SY, Chang YS, Chiu CC, et al. (April 2013).
731:. Most are found in bacteria that are not free-living; that is, they are either
3941:
3799:
3756:
3584:
3574:
2951:
Poliseno L, Salmena L, Zhang J, Carver B, Haveman WJ, Pandolfi PP (June 2010).
2878:
750:
645:
573:
494:
3202:
3063:
2017:
1955:
Xue Y, Daly A, Yngvadottir B, Liu M, Coop G, Kim Y, et al. (April 2006).
1233:
501:
techniques. Pseudogenes are often identified by the appearance of a premature
497:
technologies has led to the identification of many apparent pseudogenes using
3988:
3906:
3646:
3569:
3560:
3552:
3513:
3478:
3429:
3412:
Syberg-Olsen MJ, Garber AI, Keeling PJ, McCutcheon JP, Husnik F (July 2022).
2311:
1070:"Iterative gene prediction and pseudogene removal improves genome annotation"
795:
441:
425:
385:
297:
3108:
3091:
2271:
1410:
3918:
3447:
3396:
3361:
3318:
3263:
3220:
3168:
3117:
3071:
3029:
2994:
2928:
2911:
2896:
2839:
2804:
2763:
2714:
2665:
2624:
2573:
2536:
Belinky F, Ganguly I, Poliakov E, Yurchenko V, Rogozin IB (February 2021).
2522:
2463:
2387:
2330:
2236:
2179:
2144:
2084:
2025:
1990:
1832:
1783:
1685:
1614:
1565:
1514:
1465:
1429:
1319:
1276:
1241:
1160:
1103:
1054:
997:
856:
754:
461:
281:
44:. Most arise as superfluous copies of functional genes, either directly by
2937:
2605:
2422:
2279:
2187:
1941:
1900:
1734:
1629:
1370:
3913:
3824:
3686:
2696:
2554:
1496:
1036:
679:
663:
641:
505:
in a predicted mRNA sequence, which would, in theory, prevent synthesis (
2976:
2587:
Feng Y, Wang Z, Chien KY, Chen HL, Liang YH, Hua X, Chiu CH (May 2022).
2369:
2158:
Jeffs P, Ashburner M (May 1991). "Processed pseudogenes in
Drosophila".
2126:
1142:
244:
3742:
3387:
3370:
3343:
3133:"Decoding the similarities and differences among mycobacterial species"
2745:
1596:
1392:
Baertsch R, Diekhans M, Kent WJ, Haussler D, Brosius J (October 2008).
1362:
1085:
946:
778:
524:
502:
445:
199:
85:
69:
1814:
1530:"Gene copy-number polymorphism caused by retrotransposition in humans"
1268:
3844:
3535:
3255:
833:
687:
549:
269:
170:
26:
2831:
2049:"Progress, Challenges, and Surprises in Annotating the Human Genome"
1914:
Nishikimi M, Fukuyama R, Minoshima S, Shimizu N, Yagi K (May 1994).
1796:
534:
In 2016 it was reported that four predicted pseudogenes in multiple
3968:
3854:
3618:
3514:
3411:
3289:
Gerstein M, Zheng D (August 2006). "The real life of pseudogenes".
1972:
1311:
805:
769:
732:
728:
637:
360:
89:
73:
1750:"The probability of preservation of a newly arisen gene duplicate"
3864:
2642:. Methods in Molecular Biology. Vol. 1167. pp. 227–39.
1913:
791:
697:. Pseudogenes can, over evolutionary time scales, participate in
544:
528:
510:
470:
400:
340:
260:
2781:. Methods in Molecular Biology. Vol. 1093. pp. 47–58.
2209:
Wang W, Zhang J, Alvarez C, Llopart A, Long M (September 2000).
1527:
885:
117:
3963:
3414:"Pseudofinder: Detection of Pseudogenes in Prokaryotic Genomes"
2535:
970:
Mighell AJ, Smith NR, Robinson PA, Markham AF (February 2000).
368:
273:
223:
3185:
Lopera J, Miller IJ, McPhail KL, Kwan JC (November 21, 2017).
2100:
1394:"Retrocopy contributions to the evolution of the human genome"
1019:
Robicheau BM, Susko E, Harrigan AM, Snyder M (February 2017).
686:
of PTENP1 mRNA functions as a decoy of PTEN mRNA by targeting
513:
product of the original gene. There have been some reports of
3709:
1578:
671:
650:
636:. There are many reports of pseudogene transcripts acting as
625:
600:
409:
359:
sequences on misaligned chromosomes and subsequently acquire
301:
1846:
Zhang J (2003). "Evolution by gene duplication: an update".
1391:
1018:
553:
analysis complicates the analysis of sequence data. Another
2730:"Not so pseudo anymore: pseudogenes as therapeutic targets"
1481:"A Genome-Wide Landscape of Retrocopies in Primate Genomes"
969:
740:
364:
321:
64:, or whose coding sequences are obviously defective due to
53:
41:
3233:
2950:
2476:
296:). In the process of retrotransposition, a portion of the
152:
Pseudogenes are usually characterized by a combination of
3368:
3326:
Torrents D, Suyama M, Zdobnov E, Bork P (December 2003).
3325:
2909:
2343:
1747:
1297:
248:
Mechanism of classical and processed pseudogene formation
211:
157:
96:
37:
2776:
522:. That appears to have happened during the evolution of
3184:
3130:
2208:
1442:
743:
are lost first. For example, the oldest pseudogenes in
2860:
2003:
1116:
868:, showing three pseudogenes (indicated by crosses) in
2292:
2096:
2094:
2046:
1954:
1748:
Lynch M, O'Hely M, Walsh B, Force A (December 2001).
1701:"How often do duplicated genes evolve new functions?"
829:, use mandelalides as part of its defense mechanism.
161:
sequence alignments rather than biologically proven.
2817:
1651:
1649:
1196:
3131:Malhotra S, Vedithi SC, Blundell TL (August 2017).
2400:
1061:
836:and the domino theory of gene loss was observed in
3089:
2091:
1872:
1254:
1190:
790:, an obligate parasite and the causative agent of
455:
2586:
2435:
1646:
3986:
2047:Zerbino DR, Frankish A, Flicek P (August 2020).
2678:
719:ential genes for evolutionary diversification.
229:Processed pseudogenes often pose a problem for
169:the two sequences, the percentage of identical
3090:Dagan T, Blekhman R, Graur D (February 2006).
3007:
2157:
1907:
1866:
1478:
1291:
1248:
1067:
3499:
3288:
3042:
2727:
1997:
1873:Nishikimi M, Kawai T, Yagi K (October 1992).
1790:
1436:
1385:
1340:
1222:Current Opinion in Genetics & Development
335:
95:Pseudogene sequences may be transcribed into
2436:Betrán E, Wang W, Jin L, Long M (May 2002).
2053:Annual Review of Genomics and Human Genetics
2040:
1741:
3328:"A genome-wide survey of human pseudogenes"
2542:International Journal of Molecular Sciences
2429:
2337:
2249:
2243:
1692:
1655:
1630:"Plagiarized Errors and Molecular Genetics"
1176:"Plagiarized Errors and Molecular Genetics"
963:
3506:
3492:
1839:
1334:
3437:
3386:
3351:
3210:
3158:
3148:
3107:
2984:
2927:
2886:
2753:
2704:
2637:
2614:
2604:
2563:
2553:
2512:
2502:
2453:
2377:
2320:
2310:
2226:
2134:
2074:
2064:
1980:
1931:
1890:
1822:
1773:
1724:
1604:
1555:
1545:
1504:
1419:
1409:
1213:
1150:
1093:
1044:
987:
416:) can prevent a gene from being normally
2286:
1948:
855:
722:
624:
480:
399:
379:. Gene duplication generates functional
339:
284:consists of repetitive elements such as
259:
243:
80:deleted from their genomes by chance of
25:
3479:RCPedia - Processed Pseudogene database
3085:
3083:
3081:
2728:Roberts TC, Morris KV (December 2013).
2103:"Olfactory receptor pseudo-pseudogenes"
1167:
872:that still have functional homologs in
88:errors, or they may accumulate so many
3987:
3180:
3178:
3045:"Taking the pseudo out of pseudogenes"
3043:Goodhead I, Darby AC (February 2015).
3022:10.1146/annurev.genet.37.040103.103949
2820:Nature Reviews. Molecular Cell Biology
1627:
1199:"Origins of new genes and pseudogenes"
395:
320:; these are both hallmark features of
304:transcript of a gene is spontaneously
129:about faster accumulation of mutation.
3487:
2455:10.1093/oxfordjournals.molbev.a004124
2228:10.1093/oxfordjournals.molbev.a026413
1845:
1698:
1572:
1219:
591:
316:, and usually have had their introns
3078:
1479:Navarro FC, Galante PA (July 2015).
879:
111:
3468:Yale University pseudogene database
3175:
2066:10.1146/annurev-genom-121119-083418
1920:The Journal of Biological Chemistry
1879:The Journal of Biological Chemistry
1621:
1197:Chandrasekaran C, Betrán E (2008).
1173:
1119:"A draft map of the human proteome"
1068:van Baren MJ, Brent MR (May 2006).
543:) from gene Ir75a is found only in
452:) to positive selection in humans.
404:2 ways a pseudogene may be produced
239:
13:
3575:Short tandem repeat/Microsatellite
3475:(homologous processed pseudogenes)
3281:
1961:American Journal of Human Genetics
927:List of disabled human pseudogenes
14:
4011:
3456:
3311:10.1038/scientificamerican0806-48
2295:"The GENCODE pseudogene resource"
3137:PLOS Neglected Tropical Diseases
2160:Proceedings. Biological Sciences
884:
812:has a larger genome compared to
737:obligate intracellular parasites
467:Examples include the following:
116:
3418:Molecular Biology and Evolution
3227:
3124:
3096:Molecular Biology and Evolution
3052:Current Opinion in Microbiology
3036:
3001:
2944:
2903:
2854:
2811:
2770:
2721:
2672:
2631:
2580:
2529:
2470:
2442:Molecular Biology and Evolution
2394:
2215:Molecular Biology and Evolution
2202:
2151:
1848:Trends in Ecology and Evolution
1521:
1472:
1257:Cytogenetic and Genome Research
629:BRAF pseudogene acts as a ceRNA
456:Examples of pseudogene function
264:Processed pseudogene production
16:Functionless relative of a gene
3579:Trinucleotide repeat disorders
1343:Journal of Molecular Evolution
1110:
1012:
344:One way a pseudogene may arise
36:are nonfunctional segments of
1:
3566:Variable number tandem repeat
2415:10.1016/s0378-1119(97)00289-8
1933:10.1016/S0021-9258(17)36884-9
1892:10.1016/S0021-9258(19)36707-9
1860:10.1016/S0169-5347(03)00033-8
1678:10.1126/science.290.5494.1151
1458:10.1016/S0378-1119(02)01047-8
989:10.1016/S0014-5793(00)01199-6
957:
561:, which encodes a functional
107:
3150:10.1371/journal.pntd.0005883
2648:10.1007/978-1-4939-0835-6_15
2504:10.1371/journal.pone.0035195
1547:10.1371/journal.pgen.1003242
1485:Genome Biology and Evolution
1025:Genome Biology and Evolution
355:at, for example, repetitive
255:
72:. Pseudogenes are a type of
7:
2787:10.1007/978-1-62703-694-8_5
1766:10.1093/genetics/159.4.1789
920:
493:The rapid proliferation of
408:Various mutations (such as
10:
4016:
2879:10.1016/j.cell.2015.02.043
1717:10.1093/genetics/139.1.421
1180:Creation Evolution Journal
438:L-gulono-γ-lactone oxidase
336:Non-processed (duplicated)
18:
3934:
3887:
3755:
3723:
3700:
3677:
3668:
3659:
3634:
3594:
3551:
3542:
3533:
3203:10.1128/msystems.00096-17
3064:10.1016/j.mib.2014.11.012
3010:Annual Review of Genetics
2018:10.1038/s41576-019-0196-1
1699:Walsh JB (January 1995).
1234:10.1016/j.gde.2004.08.008
832:The relationship between
761:while the oldest ones in
727:Pseudogenes are found in
707:genes could be viewed as
515:translational readthrough
40:that resemble functional
2312:10.1186/gb-2012-13-9-r51
2006:Nature Reviews. Genetics
972:"Vertebrate pseudogenes"
691:system described above.
606:hepatocellular carcinoma
353:homologous recombination
2272:10.1126/science.7682012
1411:10.1186/1471-2164-9-466
487:Drosophila melanogaster
3974:Protein tandem repeats
3902:Tandemly arrayed genes
3430:10.1093/molbev/msac153
2929:10.1038/sj.onc.1201762
2685:Nucleic Acids Research
2593:Nucleic Acids Research
2180:10.1098/rspb.1991.0064
937:Molecular paleontology
895:is missing information
877:
808:from the same family.
630:
490:
405:
345:
265:
249:
127:is missing information
99:at low levels, due to
30:
3109:10.1093/molbev/msj036
2779:PIWI-Interacting RNAs
2350:Nature Communications
1628:Max EE (2003-05-05).
942:Pseudogene (database)
859:
810:Mycobacteirum marinum
801:Mycobacterium marinum
777:, recombination, and
759:secondary metabolites
723:Bacterial pseudogenes
676:initiating methionine
668:tumor suppressor gene
644:. The BRAF gene is a
628:
622:damage to the genome.
563:alcohol dehydrogenase
557:pseudo-pseudogene is
484:
432:, or, in some cases,
430:population bottleneck
403:
343:
326:alternatively spliced
263:
247:
50:reverse transcription
29:
3947:Pathogenicity island
2555:10.3390/ijms22041876
862:Mycobacterium leprae
814:Mycobacterium leprae
787:Mycobacterium leprae
746:Mycobacterium leprae
620:transposable element
176:Convergent evolution
21:Pseudogenes (beetle)
3303:2006SciAm.295b..48G
3291:Scientific American
3248:2001Natur.409.1007C
3242:(6823): 1007–1011.
2977:10.1038/nature09144
2969:2010Natur.465.1033P
2963:(7301): 1033–1038.
2606:10.1093/nar/gkac302
2495:2012PLoSO...735195O
2370:10.1038/ncomms11778
2362:2016NatCo...711778W
2264:1993Sci...260...91L
2172:1991RSPSB.244..151J
2127:10.1038/nature19824
2119:2016Natur.539...93P
1926:(18): 13685–13688.
1885:(30): 21967–21972.
1670:2000Sci...290.1151L
1664:(5494): 1151–1155.
1634:TalkOrigins Archive
1355:1989JMolE..28..279G
1143:10.1038/nature13302
1135:2014Natur.509..575K
932:Molecular evolution
843:Buchnera aphidicola
838:Buchnera aphidicola
396:Unitary pseudogenes
306:reverse transcribed
187:pre-mRNA processing
3897:Gene amplification
3473:Hoppsigen database
3388:10.1002/humu.20335
3344:10.1101/gr.1455503
2746:10.2217/pgs.13.172
2697:10.1093/nar/gkt047
1597:10.1101/gr.5586307
1497:10.1093/gbe/evv142
1363:10.1007/BF02103423
1086:10.1101/gr.4766206
1037:10.1093/gbe/evw307
878:
631:
599:. Some endogenous
592:Non-protein-coding
541:olfactory receptor
491:
474:pseudo-pseudogenes
414:nonsense mutations
406:
377:selection pressure
346:
278:retrotransposition
266:
250:
31:
3982:
3981:
3883:
3882:
3751:
3750:
3655:
3654:
3544:Repeated sequence
3519:repeated sequence
3338:(12): 2559–2567.
2922:(18): 2403–2406.
2796:978-1-62703-693-1
2740:(16): 2023–2034.
2657:978-1-4939-0834-9
2166:(1310): 151–159.
1815:10.1101/gr.207102
1269:10.1159/000084936
1129:(7502): 575–581.
918:
917:
860:The proS loci in
822:Verrucomicrobiota
764:Shigella flexneri
578:mass spectrometry
520:natural selection
471:Protein-coding: "
450:nonsense mutation
434:natural selection
150:
149:
48:or indirectly by
4007:
3959:Low copy repeats
3952:Symbiosis island
3889:Gene duplication
3675:
3674:
3666:
3665:
3549:
3548:
3527:gene duplication
3508:
3501:
3494:
3485:
3484:
3451:
3441:
3408:
3390:
3365:
3355:
3322:
3276:
3275:
3256:10.1038/35059006
3231:
3225:
3224:
3214:
3182:
3173:
3172:
3162:
3152:
3128:
3122:
3121:
3111:
3087:
3076:
3075:
3049:
3040:
3034:
3033:
3005:
2999:
2998:
2988:
2948:
2942:
2941:
2931:
2907:
2901:
2900:
2890:
2858:
2852:
2851:
2815:
2809:
2808:
2774:
2768:
2767:
2757:
2734:Pharmacogenomics
2725:
2719:
2718:
2708:
2691:(6): 3734–3747.
2676:
2670:
2669:
2635:
2629:
2628:
2618:
2608:
2599:(9): 5158–5170.
2584:
2578:
2577:
2567:
2557:
2533:
2527:
2526:
2516:
2506:
2474:
2468:
2467:
2457:
2433:
2427:
2426:
2398:
2392:
2391:
2381:
2341:
2335:
2334:
2324:
2314:
2290:
2284:
2283:
2247:
2241:
2240:
2230:
2221:(9): 1294–1301.
2206:
2200:
2199:
2155:
2149:
2148:
2138:
2098:
2089:
2088:
2078:
2068:
2044:
2038:
2037:
2001:
1995:
1994:
1984:
1952:
1946:
1945:
1935:
1911:
1905:
1904:
1894:
1870:
1864:
1863:
1843:
1837:
1836:
1826:
1794:
1788:
1787:
1777:
1760:(4): 1789–1804.
1745:
1739:
1738:
1728:
1696:
1690:
1689:
1653:
1644:
1643:
1641:
1640:
1625:
1619:
1618:
1608:
1576:
1570:
1569:
1559:
1549:
1525:
1519:
1518:
1508:
1491:(8): 2265–2275.
1476:
1470:
1469:
1452:(1–2): 189–194.
1440:
1434:
1433:
1423:
1413:
1389:
1383:
1382:
1338:
1332:
1331:
1295:
1289:
1288:
1252:
1246:
1245:
1217:
1211:
1210:
1203:Nature Education
1194:
1188:
1187:
1171:
1165:
1164:
1154:
1114:
1108:
1107:
1097:
1065:
1059:
1058:
1048:
1016:
1010:
1009:
991:
982:(2–3): 109–114.
967:
913:
910:
904:
888:
880:
874:M. tuberculosis.
849:Escherichia coli
666:gene is a known
509:) of the normal
476:
475:
448:gene (through a
386:genetic modeling
349:Gene duplication
294:retrotransposons
240:Types and origin
210:Pseudogenes for
145:
142:
136:
120:
112:
46:gene duplication
4015:
4014:
4010:
4009:
4008:
4006:
4005:
4004:
3985:
3984:
3983:
3978:
3930:
3879:
3747:
3719:
3696:
3670:Retrotransposon
3651:
3642:Inverted repeat
3630:
3615:DNA transposon
3611:Retrotransposon
3606:Gene conversion
3597:
3590:
3587:
3538:
3529:
3512:
3459:
3454:
3332:Genome Research
3284:
3282:Further reading
3279:
3232:
3228:
3183:
3176:
3143:(8): e0005883.
3129:
3125:
3088:
3079:
3047:
3041:
3037:
3006:
3002:
2949:
2945:
2908:
2904:
2859:
2855:
2832:10.1038/nrm3089
2816:
2812:
2797:
2775:
2771:
2726:
2722:
2677:
2673:
2658:
2636:
2632:
2585:
2581:
2534:
2530:
2475:
2471:
2434:
2430:
2399:
2395:
2342:
2338:
2291:
2287:
2258:(5104): 91–95.
2248:
2244:
2207:
2203:
2156:
2152:
2113:(7627): 93–97.
2099:
2092:
2045:
2041:
2002:
1998:
1953:
1949:
1912:
1908:
1871:
1867:
1844:
1840:
1803:Genome Research
1795:
1791:
1746:
1742:
1697:
1693:
1654:
1647:
1638:
1636:
1626:
1622:
1585:Genome Research
1577:
1573:
1540:(1): e1003242.
1526:
1522:
1477:
1473:
1441:
1437:
1390:
1386:
1339:
1335:
1300:Nature Genetics
1296:
1292:
1253:
1249:
1218:
1214:
1195:
1191:
1174:Max EE (1986).
1172:
1168:
1115:
1111:
1074:Genome Research
1066:
1062:
1017:
1013:
968:
964:
960:
952:Retrotransposon
923:
914:
908:
905:
898:
889:
866:M. tuberculosis
775:DNA replication
751:RNA polymerases
725:
699:gene conversion
594:
576:analysis using
499:gene prediction
479:
473:
472:
458:
398:
390:genome analysis
338:
272:, particularly
258:
242:
231:gene prediction
195:protein folding
146:
140:
137:
130:
121:
110:
82:DNA replication
24:
17:
12:
11:
5:
4013:
4003:
4002:
4000:Non-coding DNA
3997:
3980:
3979:
3977:
3976:
3971:
3966:
3961:
3956:
3955:
3954:
3949:
3942:Genomic island
3938:
3936:
3932:
3931:
3929:
3928:
3923:
3922:
3921:
3911:
3910:
3909:
3899:
3893:
3891:
3885:
3884:
3881:
3880:
3878:
3877:
3872:
3867:
3862:
3857:
3852:
3847:
3842:
3837:
3832:
3827:
3822:
3817:
3812:
3807:
3802:
3797:
3792:
3787:
3782:
3777:
3772:
3767:
3761:
3759:
3757:DNA transposon
3753:
3752:
3749:
3748:
3746:
3745:
3740:
3735:
3729:
3727:
3721:
3720:
3718:
3717:
3712:
3706:
3704:
3698:
3697:
3695:
3694:
3689:
3683:
3681:
3672:
3663:
3657:
3656:
3653:
3652:
3650:
3649:
3644:
3638:
3636:
3632:
3631:
3629:
3628:
3627:
3626:
3621:
3613:
3608:
3602:
3600:
3592:
3591:
3589:
3588:
3585:Macrosatellite
3582:
3572:
3563:
3557:
3555:
3553:Tandem repeats
3546:
3540:
3539:
3534:
3531:
3530:
3511:
3510:
3503:
3496:
3488:
3482:
3481:
3476:
3470:
3465:
3458:
3457:External links
3455:
3453:
3452:
3409:
3381:(6): 545–552.
3375:Human Mutation
3366:
3323:
3285:
3283:
3280:
3278:
3277:
3226:
3174:
3123:
3102:(2): 310–316.
3077:
3035:
3000:
2943:
2902:
2873:(2): 319–332.
2853:
2826:(4): 246–258.
2810:
2795:
2769:
2720:
2671:
2656:
2630:
2579:
2528:
2469:
2448:(5): 654–663.
2428:
2409:(1–2): 37–41.
2393:
2336:
2299:Genome Biology
2285:
2242:
2201:
2150:
2090:
2039:
2012:(3): 191–201.
1996:
1973:10.1086/503116
1967:(4): 659–670.
1947:
1906:
1865:
1854:(6): 292–298.
1838:
1809:(2): 272–280.
1789:
1740:
1711:(1): 421–428.
1691:
1645:
1620:
1591:(6): 839–851.
1571:
1520:
1471:
1435:
1384:
1349:(4): 279–285.
1333:
1312:10.1038/ng1223
1290:
1263:(1–4): 35–48.
1247:
1228:(6): 603–608.
1212:
1189:
1166:
1109:
1080:(5): 678–685.
1060:
1031:(2): 380–397.
1011:
961:
959:
956:
955:
954:
949:
944:
939:
934:
929:
922:
919:
916:
915:
897:about archaea.
892:
890:
883:
770:Shigella typhi
724:
721:
646:proto-oncogene
593:
590:
574:proteogenomics
495:DNA sequencing
478:
469:
457:
454:
397:
394:
337:
334:
257:
254:
241:
238:
208:
207:
179:
148:
147:
124:
122:
115:
109:
106:
15:
9:
6:
4:
3:
2:
4012:
4001:
3998:
3996:
3993:
3992:
3990:
3975:
3972:
3970:
3967:
3965:
3962:
3960:
3957:
3953:
3950:
3948:
3945:
3944:
3943:
3940:
3939:
3937:
3933:
3927:
3924:
3920:
3917:
3916:
3915:
3912:
3908:
3907:Ribosomal DNA
3905:
3904:
3903:
3900:
3898:
3895:
3894:
3892:
3890:
3886:
3876:
3873:
3871:
3868:
3866:
3863:
3861:
3858:
3856:
3853:
3851:
3848:
3846:
3843:
3841:
3838:
3836:
3833:
3831:
3828:
3826:
3823:
3821:
3818:
3816:
3813:
3811:
3808:
3806:
3803:
3801:
3798:
3796:
3793:
3791:
3788:
3786:
3783:
3781:
3778:
3776:
3773:
3771:
3768:
3766:
3763:
3762:
3760:
3758:
3754:
3744:
3741:
3739:
3736:
3734:
3731:
3730:
3728:
3726:
3722:
3716:
3713:
3711:
3708:
3707:
3705:
3703:
3699:
3693:
3690:
3688:
3685:
3684:
3682:
3680:
3676:
3673:
3671:
3667:
3664:
3662:
3658:
3648:
3647:Direct repeat
3645:
3643:
3640:
3639:
3637:
3633:
3625:
3622:
3620:
3617:
3616:
3614:
3612:
3609:
3607:
3604:
3603:
3601:
3599:
3593:
3586:
3583:
3580:
3576:
3573:
3571:
3570:Minisatellite
3567:
3564:
3562:
3561:Satellite DNA
3559:
3558:
3556:
3554:
3550:
3547:
3545:
3541:
3537:
3532:
3528:
3524:
3520:
3516:
3509:
3504:
3502:
3497:
3495:
3490:
3489:
3486:
3480:
3477:
3474:
3471:
3469:
3466:
3464:
3461:
3460:
3449:
3445:
3440:
3435:
3431:
3427:
3423:
3419:
3415:
3410:
3406:
3402:
3398:
3394:
3389:
3384:
3380:
3376:
3372:
3367:
3363:
3359:
3354:
3349:
3345:
3341:
3337:
3333:
3329:
3324:
3320:
3316:
3312:
3308:
3304:
3300:
3296:
3292:
3287:
3286:
3273:
3269:
3265:
3261:
3257:
3253:
3249:
3245:
3241:
3237:
3230:
3222:
3218:
3213:
3208:
3204:
3200:
3196:
3192:
3188:
3181:
3179:
3170:
3166:
3161:
3156:
3151:
3146:
3142:
3138:
3134:
3127:
3119:
3115:
3110:
3105:
3101:
3097:
3093:
3086:
3084:
3082:
3073:
3069:
3065:
3061:
3057:
3053:
3046:
3039:
3031:
3027:
3023:
3019:
3015:
3011:
3004:
2996:
2992:
2987:
2982:
2978:
2974:
2970:
2966:
2962:
2958:
2954:
2947:
2939:
2935:
2930:
2925:
2921:
2917:
2913:
2906:
2898:
2894:
2889:
2884:
2880:
2876:
2872:
2868:
2864:
2857:
2849:
2845:
2841:
2837:
2833:
2829:
2825:
2821:
2814:
2806:
2802:
2798:
2792:
2788:
2784:
2780:
2773:
2765:
2761:
2756:
2751:
2747:
2743:
2739:
2735:
2731:
2724:
2716:
2712:
2707:
2702:
2698:
2694:
2690:
2686:
2682:
2675:
2667:
2663:
2659:
2653:
2649:
2645:
2641:
2634:
2626:
2622:
2617:
2612:
2607:
2602:
2598:
2594:
2590:
2583:
2575:
2571:
2566:
2561:
2556:
2551:
2547:
2543:
2539:
2532:
2524:
2520:
2515:
2510:
2505:
2500:
2496:
2492:
2489:(5): e35195.
2488:
2484:
2480:
2473:
2465:
2461:
2456:
2451:
2447:
2443:
2439:
2432:
2424:
2420:
2416:
2412:
2408:
2404:
2397:
2389:
2385:
2380:
2375:
2371:
2367:
2363:
2359:
2355:
2351:
2347:
2340:
2332:
2328:
2323:
2318:
2313:
2308:
2304:
2300:
2296:
2289:
2281:
2277:
2273:
2269:
2265:
2261:
2257:
2253:
2246:
2238:
2234:
2229:
2224:
2220:
2216:
2212:
2205:
2197:
2193:
2189:
2185:
2181:
2177:
2173:
2169:
2165:
2161:
2154:
2146:
2142:
2137:
2132:
2128:
2124:
2120:
2116:
2112:
2108:
2104:
2097:
2095:
2086:
2082:
2077:
2072:
2067:
2062:
2058:
2054:
2050:
2043:
2035:
2031:
2027:
2023:
2019:
2015:
2011:
2007:
2000:
1992:
1988:
1983:
1978:
1974:
1970:
1966:
1962:
1958:
1951:
1943:
1939:
1934:
1929:
1925:
1921:
1917:
1910:
1902:
1898:
1893:
1888:
1884:
1880:
1876:
1869:
1861:
1857:
1853:
1849:
1842:
1834:
1830:
1825:
1820:
1816:
1812:
1808:
1804:
1800:
1793:
1785:
1781:
1776:
1771:
1767:
1763:
1759:
1755:
1751:
1744:
1736:
1732:
1727:
1722:
1718:
1714:
1710:
1706:
1702:
1695:
1687:
1683:
1679:
1675:
1671:
1667:
1663:
1659:
1652:
1650:
1635:
1631:
1624:
1616:
1612:
1607:
1602:
1598:
1594:
1590:
1586:
1582:
1575:
1567:
1563:
1558:
1553:
1548:
1543:
1539:
1535:
1534:PLOS Genetics
1531:
1524:
1516:
1512:
1507:
1502:
1498:
1494:
1490:
1486:
1482:
1475:
1467:
1463:
1459:
1455:
1451:
1447:
1439:
1431:
1427:
1422:
1417:
1412:
1407:
1403:
1399:
1395:
1388:
1380:
1376:
1372:
1368:
1364:
1360:
1356:
1352:
1348:
1344:
1337:
1329:
1325:
1321:
1317:
1313:
1309:
1305:
1301:
1294:
1286:
1282:
1278:
1274:
1270:
1266:
1262:
1258:
1251:
1243:
1239:
1235:
1231:
1227:
1223:
1216:
1208:
1204:
1200:
1193:
1185:
1181:
1177:
1170:
1162:
1158:
1153:
1148:
1144:
1140:
1136:
1132:
1128:
1124:
1120:
1113:
1105:
1101:
1096:
1091:
1087:
1083:
1079:
1075:
1071:
1064:
1056:
1052:
1047:
1042:
1038:
1034:
1030:
1026:
1022:
1015:
1007:
1003:
999:
995:
990:
985:
981:
977:
973:
966:
962:
953:
950:
948:
945:
943:
940:
938:
935:
933:
930:
928:
925:
924:
912:
909:December 2023
902:
896:
893:This article
891:
887:
882:
881:
875:
871:
867:
863:
858:
854:
852:
850:
845:
844:
839:
835:
830:
828:
824:
823:
817:
815:
811:
807:
803:
802:
797:
796:transcriptome
793:
789:
788:
782:
780:
776:
772:
771:
766:
765:
760:
756:
752:
748:
747:
742:
738:
734:
730:
720:
718:
717:
712:
711:
706:
705:
700:
696:
692:
689:
685:
681:
677:
673:
669:
665:
661:
657:
655:
653:
647:
643:
639:
635:
627:
623:
621:
617:
613:
609:
607:
602:
598:
589:
586:
584:
579:
575:
570:
568:
564:
560:
556:
552:
551:
546:
542:
537:
532:
530:
527:
526:
521:
516:
512:
508:
504:
500:
496:
489:
488:
483:
468:
465:
463:
453:
451:
447:
443:
442:ascorbic acid
439:
435:
431:
427:
426:genetic drift
423:
419:
415:
411:
402:
393:
391:
387:
382:
378:
374:
370:
366:
362:
358:
354:
350:
342:
333:
330:
327:
323:
319:
315:
311:
307:
303:
299:
295:
291:
287:
283:
279:
275:
271:
262:
253:
246:
237:
234:
232:
227:
225:
221:
216:
213:
205:
201:
196:
192:
188:
184:
183:Transcription
180:
177:
172:
168:
164:
163:
162:
159:
155:
144:
141:December 2023
134:
128:
125:This section
123:
119:
114:
113:
105:
102:
98:
93:
91:
87:
83:
77:
75:
71:
68:or premature
67:
63:
59:
58:transcription
55:
51:
47:
43:
39:
35:
28:
22:
3925:
3919:Gene cluster
3687:Alu sequence
3596:Interspersed
3421:
3417:
3378:
3374:
3335:
3331:
3297:(2): 48–55.
3294:
3290:
3239:
3235:
3229:
3194:
3190:
3140:
3136:
3126:
3099:
3095:
3055:
3051:
3038:
3013:
3009:
3003:
2960:
2956:
2946:
2919:
2915:
2905:
2870:
2866:
2856:
2823:
2819:
2813:
2778:
2772:
2737:
2733:
2723:
2688:
2684:
2674:
2639:
2633:
2596:
2592:
2582:
2545:
2541:
2531:
2486:
2482:
2472:
2445:
2441:
2431:
2406:
2402:
2396:
2353:
2349:
2339:
2302:
2298:
2288:
2255:
2251:
2245:
2218:
2214:
2204:
2163:
2159:
2153:
2110:
2106:
2059:(1): 55–79.
2056:
2052:
2042:
2009:
2005:
1999:
1964:
1960:
1950:
1923:
1919:
1909:
1882:
1878:
1868:
1851:
1847:
1841:
1806:
1802:
1792:
1757:
1753:
1743:
1708:
1704:
1694:
1661:
1657:
1637:. Retrieved
1623:
1588:
1584:
1574:
1537:
1533:
1523:
1488:
1484:
1474:
1449:
1445:
1438:
1401:
1398:BMC Genomics
1397:
1387:
1346:
1342:
1336:
1306:(1): 41–48.
1303:
1299:
1293:
1260:
1256:
1250:
1225:
1221:
1215:
1206:
1202:
1192:
1183:
1179:
1169:
1126:
1122:
1112:
1077:
1073:
1063:
1028:
1024:
1014:
979:
976:FEBS Letters
975:
965:
906:
894:
873:
869:
865:
861:
847:
841:
837:
831:
826:
820:
818:
813:
809:
799:
785:
783:
768:
762:
755:biosynthesis
744:
726:
715:
714:
709:
708:
703:
702:
694:
693:
659:
658:
651:
633:
632:
611:
610:
596:
595:
587:
571:
566:
558:
554:
548:
535:
533:
523:
492:
485:
466:
462:human genome
459:
407:
388:and also by
347:
331:
309:
282:human genome
267:
251:
235:
228:
217:
209:
151:
138:
126:
94:
78:
33:
32:
3995:Pseudogenes
3914:Gene family
3825:Tc1/mariner
3780:EnSpm/CACTA
3197:(6): 1–18.
3058:: 102–109.
3016:: 123–151.
2640:Pseudogenes
2548:(4): 1876.
1186:(3): 34–46.
827:Lissoclinum
680:oncogenesis
555:Drosophilia
507:translation
418:transcribed
318:spliced out
314:poly-A tail
226:sequences.
204:frameshifts
200:stop codons
191:translation
70:stop codons
66:frameshifts
62:translation
34:Pseudogenes
3989:Categories
3926:Pseudogene
3743:retroposon
3661:Transposon
3523:transposon
2305:(9): R51.
1639:2008-07-22
958:References
947:Retroposon
688:micro RNAs
536:Drosophila
525:Drosophila
503:stop codon
446:caspase 12
422:translated
381:redundancy
270:eukaryotes
268:In higher
171:base pairs
154:similarity
108:Properties
90:mutational
86:DNA repair
3845:P element
3795:Harbinger
3536:Repeatome
2356:: 11778.
2034:209393216
1209:(1): 181.
901:talk page
870:M. leprae
834:epistasis
733:symbionts
695:Potogenes
682:. The 3'
634:microRNAs
550:in silico
361:mutations
256:Processed
133:talk page
3969:Telomere
3935:See also
3875:Zisupton
3855:Polinton
3850:PiggyBac
3805:Helitron
3624:Helitron
3619:Polinton
3515:Genetics
3448:35801562
3405:20219423
3397:16671097
3362:14656963
3319:16866288
3264:11234002
3221:29181447
3191:mSystems
3169:28854187
3118:16237210
3072:25461580
3030:14616058
2995:20577206
2916:Oncogene
2897:25843629
2840:21427766
2805:24178556
2764:24279857
2715:23376929
2666:24823781
2625:35489061
2574:33672790
2523:22590500
2483:PLOS ONE
2464:11961099
2388:27250503
2331:22951037
2237:10958846
2145:27776356
2085:32421357
2026:31848477
1991:16532395
1833:11827946
1784:11779815
1754:Genetics
1705:Genetics
1686:11073452
1615:17568002
1566:23359205
1515:26224704
1466:12468100
1430:18842134
1379:22437436
1328:32151696
1320:12897783
1285:25083962
1277:16093656
1242:15531153
1161:24870542
1104:16651666
1055:28204512
1006:42204036
998:10692568
921:See also
806:pathogen
753:and the
729:bacteria
713:ogenes:
674:for the
638:microRNA
310:in vitro
167:aligning
101:promoter
74:junk DNA
3865:Transib
3840:Novosib
3820:Kolobok
3790:Ginger2
3785:Ginger1
3770:Crypton
3439:9336565
3299:Bibcode
3272:4307207
3244:Bibcode
3212:5698493
3160:5595346
2986:3206313
2965:Bibcode
2938:9620558
2888:6922011
2848:5710813
2755:4068744
2706:3616710
2616:9122581
2565:7918605
2514:3348931
2491:Bibcode
2423:9370262
2379:4895710
2358:Bibcode
2322:3491395
2280:7682012
2260:Bibcode
2252:Science
2196:1665885
2188:1679549
2168:Bibcode
2136:5164928
2115:Bibcode
2076:7116059
1982:1424700
1942:8175804
1901:1400507
1775:1461922
1735:7705642
1726:1206338
1666:Bibcode
1658:Science
1606:1891343
1557:3554589
1506:4558860
1421:2584115
1404:: 466.
1371:2499684
1351:Bibcode
1152:4403737
1131:Bibcode
1095:1457044
1046:5381670
792:leprosy
773:are in
749:are in
614:. Some
567:in vivo
565:enzyme
559:jingwei
545:neurons
529:species
511:protein
373:fitness
274:mammals
3964:CRISPR
3830:Merlin
3815:ISL2EU
3765:Academ
3598:repeat
3446:
3436:
3403:
3395:
3360:
3353:403797
3350:
3317:
3270:
3262:
3236:Nature
3219:
3209:
3167:
3157:
3116:
3070:
3028:
2993:
2983:
2957:Nature
2936:
2895:
2885:
2846:
2838:
2803:
2793:
2762:
2752:
2713:
2703:
2664:
2654:
2623:
2613:
2572:
2562:
2521:
2511:
2462:
2421:
2386:
2376:
2329:
2319:
2278:
2235:
2194:
2186:
2143:
2133:
2107:Nature
2083:
2073:
2032:
2024:
1989:
1979:
1940:
1899:
1831:
1824:155275
1821:
1782:
1772:
1733:
1723:
1684:
1613:
1603:
1564:
1554:
1513:
1503:
1464:
1428:
1418:
1377:
1369:
1326:
1318:
1283:
1275:
1240:
1159:
1149:
1123:Nature
1102:
1092:
1053:
1043:
1004:
996:
779:repair
704:pseudo
662:. The
616:piRNAs
612:piRNAs
601:siRNAs
597:siRNAs
410:indels
369:intron
224:genome
193:, and
52:of an
3870:Zator
3810:IS3EU
3715:LINE2
3710:LINE1
3702:LINEs
3679:SINEs
3635:Other
3424:(7).
3401:S2CID
3268:S2CID
3048:(PDF)
2844:S2CID
2192:S2CID
2030:S2CID
1375:S2CID
1324:S2CID
1281:S2CID
1002:S2CID
741:genes
672:codon
583:PGAM4
322:cDNAs
302:hnRNA
292:(see
290:LINEs
286:SINEs
42:genes
3860:Sola
3835:MuDR
3775:Dada
3738:MER4
3733:HERV
3725:LTRs
3444:PMID
3393:PMID
3358:PMID
3315:PMID
3260:PMID
3217:PMID
3165:PMID
3114:PMID
3068:PMID
3026:PMID
2991:PMID
2934:PMID
2893:PMID
2867:Cell
2836:PMID
2801:PMID
2791:ISBN
2760:PMID
2711:PMID
2662:PMID
2652:ISBN
2621:PMID
2570:PMID
2519:PMID
2460:PMID
2419:PMID
2403:Gene
2384:PMID
2327:PMID
2276:PMID
2233:PMID
2184:PMID
2141:PMID
2081:PMID
2022:PMID
1987:PMID
1938:PMID
1897:PMID
1829:PMID
1780:PMID
1731:PMID
1682:PMID
1611:PMID
1562:PMID
1511:PMID
1462:PMID
1446:Gene
1426:PMID
1367:PMID
1316:PMID
1273:PMID
1238:PMID
1157:PMID
1100:PMID
1051:PMID
994:PMID
864:and
846:and
804:, a
767:and
664:PTEN
660:PTEN
642:BRAF
428:, a
412:and
365:exon
357:SINE
298:mRNA
288:and
202:and
54:mRNA
3800:hAT
3692:MIR
3434:PMC
3426:doi
3383:doi
3348:PMC
3340:doi
3307:doi
3295:295
3252:doi
3240:409
3207:PMC
3199:doi
3155:PMC
3145:doi
3104:doi
3060:doi
3018:doi
2981:PMC
2973:doi
2961:465
2924:doi
2883:PMC
2875:doi
2871:161
2828:doi
2783:doi
2750:PMC
2742:doi
2701:PMC
2693:doi
2644:doi
2611:PMC
2601:doi
2560:PMC
2550:doi
2509:PMC
2499:doi
2450:doi
2411:doi
2407:198
2374:PMC
2366:doi
2317:PMC
2307:doi
2268:doi
2256:260
2223:doi
2176:doi
2164:244
2131:PMC
2123:doi
2111:539
2071:PMC
2061:doi
2014:doi
1977:PMC
1969:doi
1928:doi
1924:269
1887:doi
1883:267
1856:doi
1819:PMC
1811:doi
1770:PMC
1762:doi
1758:159
1721:PMC
1713:doi
1709:139
1674:doi
1662:290
1601:PMC
1593:doi
1552:PMC
1542:doi
1501:PMC
1493:doi
1454:doi
1450:300
1416:PMC
1406:doi
1359:doi
1308:doi
1265:doi
1261:110
1230:doi
1147:PMC
1139:doi
1127:509
1090:PMC
1082:doi
1041:PMC
1033:doi
984:doi
980:468
757:of
735:or
716:pot
710:pot
684:UTR
654:RNA
420:or
300:or
220:PCR
212:RNA
158:DNA
97:RNA
84:or
60:or
38:DNA
3991::
3525:,
3521:,
3517::
3442:.
3432:.
3422:39
3420:.
3416:.
3399:.
3391:.
3379:27
3377:.
3373:.
3356:.
3346:.
3336:13
3334:.
3330:.
3313:.
3305:.
3293:.
3266:.
3258:.
3250:.
3238:.
3215:.
3205:.
3193:.
3189:.
3177:^
3163:.
3153:.
3141:11
3139:.
3135:.
3112:.
3100:23
3098:.
3094:.
3080:^
3066:.
3056:23
3054:.
3050:.
3024:.
3014:37
3012:.
2989:.
2979:.
2971:.
2959:.
2955:.
2932:.
2920:16
2918:.
2914:.
2891:.
2881:.
2869:.
2865:.
2842:.
2834:.
2824:12
2822:.
2799:.
2789:.
2758:.
2748:.
2738:14
2736:.
2732:.
2709:.
2699:.
2689:41
2687:.
2683:.
2660:.
2650:.
2619:.
2609:.
2597:50
2595:.
2591:.
2568:.
2558:.
2546:22
2544:.
2540:.
2517:.
2507:.
2497:.
2485:.
2481:.
2458:.
2446:19
2444:.
2440:.
2417:.
2405:.
2382:.
2372:.
2364:.
2352:.
2348:.
2325:.
2315:.
2303:13
2301:.
2297:.
2274:.
2266:.
2254:.
2231:.
2219:17
2217:.
2213:.
2190:.
2182:.
2174:.
2162:.
2139:.
2129:.
2121:.
2109:.
2105:.
2093:^
2079:.
2069:.
2057:21
2055:.
2051:.
2028:.
2020:.
2010:21
2008:.
1985:.
1975:.
1965:78
1963:.
1959:.
1936:.
1922:.
1918:.
1895:.
1881:.
1877:.
1852:18
1850:.
1827:.
1817:.
1807:12
1805:.
1801:.
1778:.
1768:.
1756:.
1752:.
1729:.
1719:.
1707:.
1703:.
1680:.
1672:.
1660:.
1648:^
1632:.
1609:.
1599:.
1589:17
1587:.
1583:.
1560:.
1550:.
1536:.
1532:.
1509:.
1499:.
1487:.
1483:.
1460:.
1448:.
1424:.
1414:.
1400:.
1396:.
1373:.
1365:.
1357:.
1347:28
1345:.
1322:.
1314:.
1304:35
1302:.
1279:.
1271:.
1259:.
1236:.
1226:14
1224:.
1205:.
1201:.
1182:.
1178:.
1155:.
1145:.
1137:.
1125:.
1121:.
1098:.
1088:.
1078:16
1076:.
1072:.
1049:.
1039:.
1027:.
1023:.
1000:.
992:.
978:.
974:.
781:.
656:.
652:ce
569:.
531:.
276:,
189:,
185:,
178:).
76:.
3581:)
3577:(
3568:/
3507:e
3500:t
3493:v
3450:.
3428::
3407:.
3385::
3364:.
3342::
3321:.
3309::
3301::
3274:.
3254::
3246::
3223:.
3201::
3195:2
3171:.
3147::
3120:.
3106::
3074:.
3062::
3032:.
3020::
2997:.
2975::
2967::
2940:.
2926::
2899:.
2877::
2850:.
2830::
2807:.
2785::
2766:.
2744::
2717:.
2695::
2668:.
2646::
2627:.
2603::
2576:.
2552::
2525:.
2501::
2493::
2487:7
2466:.
2452::
2425:.
2413::
2390:.
2368::
2360::
2354:7
2333:.
2309::
2282:.
2270::
2262::
2239:.
2225::
2198:.
2178::
2170::
2147:.
2125::
2117::
2087:.
2063::
2036:.
2016::
1993:.
1971::
1944:.
1930::
1903:.
1889::
1862:.
1858::
1835:.
1813::
1786:.
1764::
1737:.
1715::
1688:.
1676::
1668::
1642:.
1617:.
1595::
1568:.
1544::
1538:9
1517:.
1495::
1489:7
1468:.
1456::
1432:.
1408::
1402:9
1381:.
1361::
1353::
1330:.
1310::
1287:.
1267::
1244:.
1232::
1207:1
1184:6
1163:.
1141::
1133::
1106:.
1084::
1057:.
1035::
1029:9
1008:.
986::
911:)
907:(
903:.
851:,
477:"
367:-
143:)
139:(
135:.
23:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
