89:
162:
309:) project reported that detectable biochemical activity was observed in regions covering at least 80% of the human genome, with biochemical activity defined primarily as being transcribed. While these findings were announced as the demise of junk DNA it is important to point out that transcription does not mean a sequence is "functional", analogous to some meaningless text that can be transcribed or copied without having any meaning.
196:"Dear Francis, I am sure that you realize how frightfully angry a lot of people will be if you say that much of the DNA is junk. The geneticists will be angry because they think that DNA is sacred. The Darwinian evolutionists will be outraged because they believe every change in DNA that is accepted in evolution is necessarily an adaptive change. To suggest anything else is an insult to the sacred memory of Darwin."
85:. The paradox was resolved with the discovery of repetitive DNA and the observation that most of the differences in genome size could be attributed to repetitive DNA. Some scientists thought that most of the repetitive DNA was involved in regulating gene expression but many scientists thought that the excess repetitive DNA was nonfunctional.
245:
non-coding DNA. But
Comings never said that. In that paper he discusses non-coding genes for ribosomal RNA and tRNAs and non-coding regulatory DNA and he proposes several possible functions for the bulk of non-coding DNA. In another publication from the same year Comings again discusses the term junk
131:
popularized the term in a 1972 paper titled "So much 'junk' DNA in our genome" where he summarized the current evidence that had accumulated by then. In a second paper that same year, he concluded that 90% of mammalian genomes consisted of nonfunctional DNA. The case for junk DNA was summarized in a
76:
In 1966 Muller reviewed these predictions and concluded that the human genome could only contain about 30,000 genes based on the number of deleterious mutations that the species could tolerate. Similar predictions were made by other leading experts in molecular evolution who concluded that the human
51:
It is difficult to determine whether other regions of the genome are functional or nonfunctional. There is considerable controversy over which criteria should be used to identify function. Many scientists have an evolutionary view of the genome and they prefer criteria based on whether DNA sequences
43:
All protein-coding regions are generally considered to be functional elements in genomes. Additionally, non-protein coding regions such as genes for ribosomal RNA and transfer RNA, regulatory sequences, origins of replication, centromeres, telomeres, and scaffold attachment regions are considered as
374:
Opponents of junk DNA argue that biochemical activity detects functional regions of the genome that are not identified by sequence conservation or purifying selection. According to some scientists, until a region in question has been shown to have additional features, beyond what is expected of the
60:
The idea that only a fraction of the human genome could be functional dates back to the late 1940s. The estimated mutation rate in humans suggested that if a large fraction of those mutations were deleterious then the human species could not survive such a mutation load (genetic load). This led to
111:
was refined to include RNA:DNA hybridization leading to the discovery that considerably less than 10% of the human genome was complementary to mRNA and this DNA was in the unique (non-repetitive) fraction. This confirmed the predictions made from genetic load arguments and was consistent with the
157:
in the 1970s seemed to confirm the views of junk DNA proponents because it meant that genes were very large and even huge genomes could not accommodate large numbers of genes. The proponents of junk DNA tended to dismiss intron sequences as mostly nonfunctional DNA (junk) but junk DNA opponents
187:
The positions of the two sides of the controversy hardened with one side believing that evolution was consistent with large amounts of junk DNA and the other side believing that natural selection should eliminate junk DNA. These differing views of evolution were highlighted in a letter from
80:
The size of genomes in various species was known to vary considerably and there did not seem to be a correlation between genome size and the complexity of the species. Even closely related species could have very different genome sizes. This observation led to what came to be known as the
324:
However, in most animal or plant genomes, a large fraction of DNA is non-functional, given that there is no obvious selective pressure on these sequences. More importantly, there is strong evidence that these sequences are not functional in other ways (using the human genome as example):
253:
It is simply not true that noncoding DNA has long been dismissed as worthless junk and that functional hypotheses have only recently been proposed - despite the frequency with which this cliché is repeated in media reports and in the introduction of far too many scientific
179:
Opponents of junk DNA interpreted these results as evidence that most of the genome is functional and they developed several hypotheses advocating that transposon sequences could benefit the organism or the species. The most important opponent of junk DNA at this time was
232:
When it was first discovered, the nongenic DNA was sometimes called—somewhat derisively by people who did not know better—"junk DNA" because it had no obvious utility, and they foolishly assumed that if it was not carrying coding information it must be useless trash.
115:
The idea that large amounts of eukaryotic genomes could be nonfunctional conflicted with the prevailing view of evolution in 1968 since it seemed likely that nonfunctional DNA would be eliminated by natural selection. The development of the
359:
in the human genome of which 23.4% affected multiple genes (by deleting them or part of them). This study also found 47 deletions of >1 MB, showing that large chunks of the human genome can get deleted without obvious consequences.
95:(Tomoko Harada) developed the nearly neutral theory that led to an understanding of how slightly deleterious junk DNA could be maintained in the genomes of species with small effective population sizes. In 2015 she was awarded the
262:
Revisionist claims that equate noncoding DNA with junk merely reveal that people who are allowed to exhibit their logorrhea in Nature and other glam journals are as ignorant as the worst young-earth creationists.
271:
The main challenge of identifying junk DNA is to distinguish between "functional" and "non-functional " sequences. This is non-trivial, but there is some good evidence for both categories.
124:
provided a way out of this problem since it allowed for the preservation of slightly deleterious nonfunctional DNA in accordance with fundamental principles of population genetics.
2095:
Cavalier-Smith T (December 1978). "Nuclear volume control by nucleoskeletal DNA, selection for cell volume and cell growth rate, and the solution of the DNA C-value paradox".
1209:
Britten RJ, Kohne DE (August 1968). "Repeated sequences in DNA. Hundreds of thousands of copies of DNA sequences have been incorporated into the genomes of higher organisms".
176:. This prompted a series of papers and letters describing transposons as selfish DNA that acted as a parasite in genomes and produced no fitness advantage for the organism.
352:(2.9% of total genome). Many of these sequences are the descendents of ancient virus infections and are thus "non-functional" in terms of human genome function.
184:
who argued that the extra DNA was required to increase the volume of the nucleus in order to promote more efficient transport across the nuclear membrane.
224:
Close to 99 percent of our genome has been historically classified as noncoding, useless "junk" DNA. Consequently, these sequences were rarely studied.
249:
The idea that all non-coding DNA was thought to be junk has been criticized by numerous authors for distorting the history of junk DNA; for example:
208:"A concept that is repugnant to us is that about half of the DNA of higher organisms is trivial or permanently inert (on an evolutionary timescale)."
237:
The common theme is that the original proponents of junk DNA thought that all non-coding DNA was junk. This claim has been attributed to a paper by
217:
There is considerable confusion in the popular press and in the scientific literature about the distinction between non-coding DNA and junk DNA.
644:
121:
282:
are the most obvious functional sequences in genomes. However, they make up only 1-2% of most vertebrate genomes. However, there are also
2346:
964:
Lin Y, Zhai T, Zhang X (April 2014). "Nanoscale heat transfer in direct nanopatterning into gold films by a nanosecond laser pulse".
69:, that only a small percentage of the human genome contains functional DNA elements (genes) that can be destroyed by mutation. (see
355:(2) Many sequences can be deleted as shown by comparing genomes. For instance, an analysis of 14,623 individuals identified 42,765
168:
and others promoted the idea that transposons were examples of selfish DNA and were responsible for the proliferation of junk DNA.
3026:"8.2% of the Human genome is constrained: variation in rates of turnover across functional element classes in the human lineage"
368:
367:
on these sequences, so they can rather freely mutate. About 11% or less of the human genome is conserved and about 7% is under
2477:
Dunham I, Kundaje A, Aldred SF, Collins PJ, Davis CA, Doyle F, et al. (The ENCODE Project
Consortium) (September 2012).
117:
2605:
2249:"On the immortality of television sets: "function" in the human genome according to the evolution-free gospel of ENCODE"
333:
1115:
Ohta T, Kimura M (September 1971). "Functional organization of genetic material as a product of molecular evolution".
52:
are preserved by natural selection. Other scientists dispute this view or have different interpretations of the data.
3008:
341:
2131:
345:
363:(3) Only a small fraction of the human genome is conserved, indicating that there is no strong (functional)
3134:
Halldorsson BV, Eggertsson HP, Moore KH, Hauswedell H, Eiriksson O, Ulfarsson MO, et al. (July 2022).
349:
328:(1) Repetitive elements, especially mobile elements make up a large fraction of the human genome, such as
143:
current estimates of the number of genes (in 1972) are much less than the number that can be accommodated,
747:
Germain PL, Ratti E, and Boem F (2014). "Junk or functional DNA? ENCODE and the function controversy".
321:
which typically have an extremely high gene density, with only a few percent being not protein-coding.
329:
28:) is a DNA sequence that has no known biological function. Most organisms have some junk DNA in their
1765:
Doolittle WF, Sapienza C (April 1980). "Selfish genes, the phenotype paradigm and genome evolution".
77:
genome could not contain more than 40,000 genes and that less than 10% of the genome was functional.
3254:"A Kuhnian revolution in molecular biology: Most genes in complex organisms express regulatory RNAs"
302:. These sequences are usually conserved in evolution and make up another 3-8% of the human genome.
2844:"Keeping up with the genomes: efficient learning of our increasing knowledge of the tree of life"
2395:
Graur D (2017). "Rubbish DNA: The functionless fraction of the human genome". In Saitou N (ed.).
1328:
Lewin B (1974). "Sequence
Organization of Eukaryotic DNA: Defining the Unit of Gene Expression".
2428:
Siepel A, Bejerano G, Pedersen JS, Hinrichs AS, Hou M, Rosenbloom K, et al. (August 2005).
88:
40:
and viruses—but it is possible that some organisms have substantial amounts of junk DNA.
364:
295:
181:
66:
1440:
Kimura M, Ohta T (February 1971). "Protein polymorphism as a phase of molecular evolution".
831:
Kellis M, Wold B, Snyder MP, Bernstein BE, Kundaje A, Marinov GK, et al. (April 2014).
132:
lengthy paper by David
Comings in 1972 where he listed four reasons for proposing junk DNA:
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1994:
1937:
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1449:
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844:
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445:
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356:
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DNA with the clear understanding that it does not include non-coding regulatory sequences.
172:
By 1980 it was apparent that most of the repetitive DNA in the human genome was related to
158:
advanced a number of hypotheses attributing functions of various sort to intron sequences.
3075:
Christmas MJ, Kaplow IM, Genereux DP, Dong MX, Hughes GM, Li X, et al. (April 2023).
8:
291:
3151:
3111:
3076:
2959:
2751:
2692:
2640:
2494:
2209:
Niu DK, Jiang L (2013). "Can ENCODE tell us how much junk DNA we carry in our genome?".
2055:
1998:
1941:
1884:
1829:
1778:
1733:
1686:
1643:
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1128:
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848:
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449:
3321:
3294:
3229:
3202:
3178:
3135:
3052:
3025:
2976:
2943:
2919:
2895:"Repetitive DNA and next-generation sequencing: computational challenges and solutions"
2894:
2870:
2843:
2819:
2794:
2770:
2735:
2711:
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2511:
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2429:
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2273:
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2077:
2020:
1963:
1906:
1849:
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1747:
1612:
1473:
1148:
1051:
997:
981:
949:
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867:
832:
810:
764:
724:
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675:
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550:
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3116:
3057:
3004:
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2924:
2875:
2824:
2775:
2716:
2672:
2654:
2586:
2551:
2516:
2459:
2369:
2347:"Conceptual and empirical challenges of ascribing functions to transposable elements"
2327:
2278:
2226:
2112:
2069:
2012:
1955:
1898:
1841:
1790:
1698:
1655:
1604:
1520:
1465:
1422:
1418:
1387:
1341:
1277:
1234:
1188:
1140:
1097:
1043:
989:
913:
872:
814:
802:
768:
729:
667:
626:
582:
531:
463:
2942:
Abel HJ, Larson DE, Regier AA, Chiang C, Das I, Kanchi KL, et al. (July 2020).
2381:
2081:
2024:
1152:
1001:
679:
475:
3316:
3306:
3265:
3224:
3214:
3173:
3163:
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3096:
3088:
3047:
3037:
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2963:
2914:
2906:
2865:
2855:
2814:
2806:
2765:
2755:
2706:
2696:
2644:
2578:
2543:
2534:
Pennisi E (September 2012). "Genomics. ENCODE project writes eulogy for junk DNA".
2506:
2498:
2449:
2441:
2361:
2317:
2309:
2268:
2260:
2218:
2104:
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2002:
1967:
1945:
1910:
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1833:
1802:
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1751:
1737:
1690:
1647:
1616:
1594:
1477:
1457:
1414:
1379:
1337:
1269:
1252:
Britten RJ, Davidson EH (July 1969). "Gene regulation for higher cells: a theory".
1226:
1180:
1132:
1089:
1055:
1035:
973:
944:
903:
862:
852:
794:
756:
719:
709:
659:
622:
572:
562:
521:
511:
453:
318:
201:
2547:
2430:"Evolutionarily conserved elements in vertebrate, insect, worm, and yeast genomes"
605:
Ohno S (1972). "An argument for the genetic simplicity of man and other mammals".
3042:
2582:
1273:
1230:
1093:
516:
189:
82:
3203:"A meta-analysis of the genomic and transcriptomic composition of complex life"
3159:
2860:
2740:
Proceedings of the
National Academy of Sciences of the United States of America
2681:
Proceedings of the
National Academy of Sciences of the United States of America
2222:
1370:
O'Brien SJ (March 1973). "On estimating functional gene number in eukaryotes".
837:
Proceedings of the
National Academy of Sciences of the United States of America
404:
399:
96:
62:
45:
2967:
2944:"Mapping and characterization of structural variation in 17,795 human genomes"
2649:
2624:
798:
760:
714:
458:
433:
3344:
3311:
3168:
3077:"Evolutionary constraint and innovation across hundreds of placental mammals"
1492:
279:
238:
165:
61:
predictions in the late 1940s by one of the founders of population genetics,
3092:
2760:
2701:
1651:
857:
3350:
3330:
3279:
3270:
3253:
3238:
3187:
3120:
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2985:
2928:
2879:
2828:
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2720:
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2590:
2555:
2520:
2463:
2373:
2331:
2282:
2230:
1694:
1383:
1192:
1144:
993:
917:
876:
806:
733:
671:
586:
535:
467:
389:
337:
104:
70:
3219:
2073:
2016:
1959:
1902:
1845:
1794:
1659:
1524:
1469:
1426:
1391:
1281:
1238:
1101:
1047:
663:
2810:
2313:
2264:
2116:
2108:
1702:
1608:
567:
173:
128:
92:
37:
2502:
375:
null hypothesis, it should provisionally be labelled as non-functional.
146:
the mutation load would be too large if all the DNA were functional, and
3101:
2445:
1816:
Orgel LE, Crick FH (April 1980). "Selfish DNA: the ultimate parasite".
985:
299:
33:
2149:
Mortola E, Long M (2021). "Turning Junk into Us: How Genes Are Born".
1494:
Requiem for a Gene: The
Problem of Junk DNA for the Molecular Paradigm
1018:
Kimura M (February 1968). "Evolutionary rate at the molecular level".
136:
some organisms have a lot more DNA than they seem to require (C-value
2064:
2039:
2007:
1982:
1950:
1925:
1893:
1868:
1837:
1786:
1742:
1717:
1599:
1574:
1461:
1136:
1039:
429:
161:
103:
At about the same time (late 1960s) the newly developed technique of
3133:
2910:
2365:
2247:
Graur D, Zheng Y, Price N, Azevedo RB, Zufall RA, Elhaik E (2013).
977:
2795:"Distinguishing between "function" and "effect" in genome biology"
1355:
Lewin B (1974). "Chapter 5: Transcription and
Processing of RNA".
241:
in 1972 where he is reported to have said that junk DNA refers to
192:, a proponent of junk DNA, to Francis Crick on December 20, 1979:
2606:"Scientists attacked over claim that 'junk DNA' is vital to life"
220:
According to an article published in 2021 in
American Scientist:
137:
2479:"An integrated encyclopedia of DNA elements in the human genome"
1560:
The Black Box of Biology: A History of the Molecular Revolution
384:
306:
154:
29:
2792:
2427:
1543:
Comings DE (1972). "The structure and function of chromatin".
1171:
Thomas CA (1971). "The genetic organization of chromosomes".
2793:
Doolittle WF, Brunet TD, Linquist S, Gregory TR (May 2014).
3074:
127:
The term "junk DNA" began to be used in the late 1950s but
2476:
2344:
1758:
1558:
Morange M (2020). "Chapter 17: Split Genes and Splicing".
1313:
Lewin B (1974). "Chapter 4: Sequences of Eukaryotic DNA".
2727:
1072:
King JL, Jukes TH (May 1969). "Non-Darwinian evolution".
830:
3023:
1245:
1204:
1202:
691:
689:
1917:
1298:
Gregory TR (2005). "Genome Size Evolution in Animals".
645:"Genome as a multipurpose structure built by evolution"
112:
idea that much of the repetitive DNA is nonfunctional.
3295:"Non-coding RNA: what is functional and what is junk?"
3194:
3024:
Rands CM, Meader S, Ponting CP, Lunter G (July 2014).
2941:
2625:"The ENCODE project: missteps overshadowing a success"
2246:
1673:
Crick F (April 1979). "Split genes and RNA splicing".
833:"Defining functional DNA elements in the human genome"
1199:
686:
99:
by the Royal Swedish Academy (with Richard Lewontin).
3136:"The sequences of 150,119 genomes in the UK Biobank"
3068:
2786:
2298:"An evolutionary classification of genomic function"
2204:
2202:
746:
493:
491:
489:
487:
485:
204:
and Kohne in their seminal paper on repetitive DNA.
2568:
2211:
Biochemical and Biophysical Research Communications
2172:. New York, New York, USA: Oxford University Press.
1809:
1630:Gilbert W (May 1985). "Genes-in-pieces revisited".
1511:Ohno S (1972). "So much "junk" DNA in our genome".
1433:
1108:
3127:
2088:
2031:
2841:
2562:
2295:
2199:
1764:
740:
482:
3342:
3003:. Sunderland MA (USA): Sinauer Associates, Inc.
2892:
2242:
2240:
1405:Bishop JO (June 1974). "The gene numbers game".
1067:
1065:
2733:
2414:. New York, New York, USA: W. A. Benjamin, Inc.
2345:Elliott TA, Linquist S, and Gregory TR (2014).
1538:
1536:
1534:
1251:
266:
3286:
3200:
2842:Zhao Z, Cristian A, Rosen G (September 2020).
2338:
2094:
2037:
1923:
1293:
1291:
497:
212:
3245:
2665:
2237:
1709:
1166:
1164:
1162:
1062:
1013:
1011:
780:
778:
695:
638:
636:
2527:
2470:
2176:
2170:DNA Demystified: Unraveling the Double Helix
1551:
1531:
1506:
1504:
1208:
963:
2148:
1718:"Genes in pieces: were they ever together?"
1623:
1566:
1369:
1288:
924:
600:
598:
596:
3292:
2893:Treangen TJ, Salzberg SL (November 2011).
2569:Casane D, Fumey J, Laurenti P (2015). "".
2403:
1815:
1439:
1398:
1363:
1348:
1321:
1159:
1114:
1008:
957:
883:
785:Mattick JS (2023). "RNA out of the mist".
775:
633:
548:
434:"The C-value paradox, junk DNA and ENCODE"
424:
422:
420:
3320:
3310:
3269:
3228:
3218:
3177:
3167:
3110:
3100:
3051:
3041:
2975:
2918:
2869:
2859:
2818:
2769:
2759:
2710:
2700:
2671:
2648:
2616:
2510:
2453:
2388:
2321:
2272:
2063:
2006:
1949:
1892:
1860:
1741:
1715:
1666:
1598:
1501:
1357:Gene Expression-2: Eukaryotic Chromosomes
1315:Gene Expression-2: Eukaryotic Chromosomes
1306:
1071:
948:
907:
866:
856:
723:
713:
576:
566:
525:
515:
457:
258:Some of the criticisms have been strong:
200:The other point of view was expressed by
3201:Liu G, Mattick JS, Taft RJ (July 2013).
2677:"Is junk DNA bunk? A critique of ENCODE"
2597:
2208:
2167:
1974:
826:
824:
593:
160:
87:
16:Those parts of a genome with no function
3251:
2734:Brunet TD, Doolittle WF (August 2014).
2533:
2182:
1924:Dover G, Doolittle WF (December 1980).
1629:
1572:
1557:
1542:
1297:
930:
784:
642:
417:
3343:
2409:
2129:
1404:
1170:
1017:
889:
3017:
2998:
2603:
2423:
2421:
2394:
2296:Graur D, Zheng Y, Azevedo RB (2015).
1866:
1672:
1490:
1354:
1327:
1312:
933:"The rate of mutation of human genes"
821:
551:"Factors behind junk DNA in bacteria"
2622:
1980:
1510:
652:Perspectives in Biology and Medicine
604:
428:
344:(20.4% of total genome), SVAs (SINE-
2136:National Institutes of Health (USA)
1185:10.1146/annurev.ge.05.120171.001321
498:Palazzo AF, Gregory TR (May 2014).
13:
2418:
2189:American Journal of Human Genetics
950:10.1111/j.1601-5223.1949.tb03339.x
896:American Journal of Human Genetics
336:(13.1% of total genome) including
305:The Encyclopedia of DNA Elements (
14:
3362:
698:"Non-Darwinian Molecular Biology"
549:Gil R, Latorre A (October 2012).
312:
228:A book published in 2020 states:
2992:
2935:
2886:
2835:
2397:Evolution of the Human Genome I
2289:
2161:
2142:
2123:
1484:
3001:Molecular and Genome Evolution
2183:Comings DE (1972). "Review of
2130:Thomas J (December 29, 1979).
2038:Cavalier-Smith T (June 1980).
1513:Brookhaven Symposia in Biology
696:Palazzo AF, Kejiou NS (2022).
542:
317:Non-functional DNA is rare in
1:
3252:Mattick JS (September 2023).
2604:McKie R (February 24, 2013).
2548:10.1126/science.337.6099.1159
2412:Molecular Biology of the Gene
2185:Evolution of Genetics Systems
1547:. Springer. pp. 237–431.
410:
274:
149:some junk DNA clearly exists.
3043:10.1371/journal.pgen.1004525
2799:Genome Biology and Evolution
2302:Genome Biology and Evolution
2253:Genome Biology and Evolution
1497:(MA). University of Chicago.
1419:10.1016/0092-8674(74)90095-6
1342:10.1016/0092-8674(74)90125-1
1274:10.1126/science.165.3891.349
1231:10.1126/science.161.3841.529
1094:10.1126/science.164.3881.788
627:10.1016/0047-2484(72)90011-5
517:10.1371/journal.pgen.1004351
267:Functional vs non-functional
7:
3293:Palazzo AF, Lee ES (2015).
2399:. Springer. pp. 19–60.
1926:"Modes of genome evolution"
1573:Gilbert W (February 1978).
1562:. Harvard University Press.
1300:The Evolution of the Genome
378:
213:Junk DNA and non-coding DNA
10:
3367:
3160:10.1038/s41586-022-04965-x
2861:10.1186/s12859-020-03744-7
2736:"Getting "function" right"
2583:10.1051/medsci/20153106023
2223:10.1016/j.bbrc.2012.12.074
1545:Advances in human genetics
1302:. Elsevier. pp. 3–87.
607:Journal of Human Evolution
55:
44:functional elements. (See
2968:10.1038/s41586-020-2371-0
2650:10.1016/j.cub.2013.03.023
2132:"letter to Francis Crick"
1981:Jain HK (December 1980).
1173:Annual Review of Genetics
799:10.1016/j.tig.2022.11.001
761:10.1007/s10539-014-9441-3
715:10.3389/fgene.2022.831068
459:10.1016/j.cub.2012.10.002
3312:10.3389/fgene.2015.00002
2899:Nature Reviews. Genetics
1359:. John Wiley & Sons.
1317:. John Wiley & Sons.
749:Biology & Philosophy
350:Class II DNA transposons
332:(8.3% of total genome),
280:Protein-coding sequences
65:, and by Nobel laureate
3093:10.1126/science.abn3943
2761:10.1073/pnas.1409762111
2702:10.1073/pnas.1221376110
2354:The American Naturalist
2097:Journal of Cell Science
1652:10.1126/science.4001923
892:"Our load of mutations"
890:Muller HJ (June 1950).
858:10.1073/pnas.1318948111
500:"The case for junk DNA"
48:for more information.)
3271:10.1002/bies.202300080
2623:Eddy SR (April 2013).
1695:10.1126/science.373120
1575:"Why genes in pieces?"
1384:10.1038/newbio242052a0
296:origins of replication
290:DNA sequences such as
264:
256:
235:
226:
210:
198:
169:
100:
73:for more information)
3299:Frontiers in Genetics
3220:10.1186/1877-6566-7-2
2040:"How selfish is DNA?"
1867:Dover G (June 1980).
702:Frontiers in Genetics
664:10.1353/pbm.2014.0008
260:
251:
230:
222:
206:
194:
182:Thomas Cavalier-Smith
164:
122:nearly neutral theory
91:
2542:(6099): 1159, 1161.
2109:10.1242/jcs.34.1.247
1716:Doolittle W (1978).
568:10.3390/genes3040634
395:Comparative genomics
330:LTR retrotransposons
292:regulatory sequences
3152:2022Natur.607..732H
2960:2020Natur.583...83A
2752:2014PNAS..111E3365P
2693:2013PNAS..110.5294D
2641:2013CBio...23.R259E
2503:10.1038/nature11247
2495:2012Natur.489...57T
2168:McHughen A (2020).
2056:1980Natur.285..617C
1999:1980Natur.288..647J
1942:1980Natur.288..646D
1885:1980Natur.285..618D
1830:1980Natur.284..604O
1779:1980Natur.284..601D
1734:1978Natur.272..581D
1687:1979Sci...204..264C
1644:1985Sci...228..823G
1591:1978Natur.271..501G
1454:1971Natur.229..467K
1266:1969Sci...165..349B
1223:1968Sci...161..529B
1129:1971Natur.233..118O
1086:1969Sci...164..788L
1032:1968Natur.217..624K
849:2014PNAS..111.6131K
619:1972JHumE...1..651O
450:2012CBio...22.R898E
369:purifying selection
357:structural variants
3087:(6643): eabn3943.
2848:BMC Bioinformatics
2811:10.1093/gbe/evu098
2446:10.1101/gr.3715005
2314:10.1093/gbe/evv021
2265:10.1093/gbe/evt028
2151:American Scientist
931:Haldane J (1949).
787:Trends in Genetics
643:Morange M (2014).
365:selection pressure
170:
101:
26:non-functional DNA
3213:(13): 2061–2072.
3169:20.500.11815/3726
3146:(7920): 732–740.
2687:(14): 5294–5300.
2571:Medecine Sciences
2410:Watson J (1965).
2050:(5767): 617–618.
1993:(5792): 647–648.
1936:(5792): 646–647.
1879:(5767): 618–620.
1824:(5757): 604–607.
1773:(5757): 601–603.
1728:(5654): 581–582.
1681:(4390): 264–271.
1638:(4701): 823–824.
1448:(5285): 467–469.
1260:(3891): 349–357.
1217:(3841): 529–540.
1123:(5315): 118–119.
1080:(3881): 788–798.
1026:(5129): 624–626.
843:(17): 6131–6138.
444:(21): R898–R899.
432:(November 2012).
319:bacterial genomes
153:The discovery of
36:and fragments of
3358:
3335:
3334:
3324:
3314:
3290:
3284:
3283:
3273:
3249:
3243:
3242:
3232:
3222:
3198:
3192:
3191:
3181:
3171:
3131:
3125:
3124:
3114:
3104:
3072:
3066:
3065:
3055:
3045:
3021:
3015:
3014:
2999:Graur D (2016).
2996:
2990:
2989:
2979:
2939:
2933:
2932:
2922:
2890:
2884:
2883:
2873:
2863:
2839:
2833:
2832:
2822:
2805:(5): 1234–1237.
2790:
2784:
2783:
2773:
2763:
2731:
2725:
2724:
2714:
2704:
2669:
2663:
2662:
2652:
2635:(7): R259–R261.
2620:
2614:
2613:
2601:
2595:
2594:
2577:(6–7): 680–686.
2566:
2560:
2559:
2531:
2525:
2524:
2514:
2474:
2468:
2467:
2457:
2440:(8): 1034–1050.
2425:
2416:
2415:
2407:
2401:
2400:
2392:
2386:
2385:
2351:
2342:
2336:
2335:
2325:
2293:
2287:
2286:
2276:
2244:
2235:
2234:
2217:(4): 1340–1343.
2206:
2197:
2196:
2180:
2174:
2173:
2165:
2159:
2158:
2146:
2140:
2139:
2127:
2121:
2120:
2092:
2086:
2085:
2067:
2065:10.1038/285617a0
2035:
2029:
2028:
2010:
2008:10.1038/288647a0
1983:"Incidental DNA"
1978:
1972:
1971:
1953:
1951:10.1038/288646a0
1921:
1915:
1914:
1896:
1894:10.1038/285618a0
1864:
1858:
1857:
1838:10.1038/284604a0
1813:
1807:
1806:
1787:10.1038/284601a0
1762:
1756:
1755:
1745:
1743:10.1038/272581a0
1713:
1707:
1706:
1670:
1664:
1663:
1627:
1621:
1620:
1602:
1600:10.1038/271501a0
1570:
1564:
1563:
1555:
1549:
1548:
1540:
1529:
1528:
1508:
1499:
1498:
1491:Sweet A (2022).
1488:
1482:
1481:
1462:10.1038/229467a0
1437:
1431:
1430:
1402:
1396:
1395:
1367:
1361:
1360:
1352:
1346:
1345:
1325:
1319:
1318:
1310:
1304:
1303:
1295:
1286:
1285:
1249:
1243:
1242:
1206:
1197:
1196:
1168:
1157:
1156:
1137:10.1038/233118a0
1112:
1106:
1105:
1069:
1060:
1059:
1040:10.1038/217624a0
1015:
1006:
1005:
972:(7): 8396–8404.
961:
955:
954:
952:
928:
922:
921:
911:
887:
881:
880:
870:
860:
828:
819:
818:
782:
773:
772:
744:
738:
737:
727:
717:
693:
684:
683:
649:
640:
631:
630:
602:
591:
590:
580:
570:
546:
540:
539:
529:
519:
495:
480:
479:
461:
426:
202:Roy John Britten
3366:
3365:
3361:
3360:
3359:
3357:
3356:
3355:
3341:
3340:
3339:
3338:
3291:
3287:
3264:(9): e2300080.
3250:
3246:
3199:
3195:
3132:
3128:
3073:
3069:
3036:(7): e1004525.
3022:
3018:
3011:
2997:
2993:
2954:(7814): 83–89.
2940:
2936:
2911:10.1038/nrg3117
2891:
2887:
2840:
2836:
2791:
2787:
2732:
2728:
2670:
2666:
2629:Current Biology
2621:
2617:
2602:
2598:
2567:
2563:
2532:
2528:
2489:(7414): 57–74.
2475:
2471:
2434:Genome Research
2426:
2419:
2408:
2404:
2393:
2389:
2349:
2343:
2339:
2294:
2290:
2245:
2238:
2207:
2200:
2181:
2177:
2166:
2162:
2147:
2143:
2128:
2124:
2093:
2089:
2036:
2032:
1979:
1975:
1922:
1918:
1869:"Ignorant DNA?"
1865:
1861:
1814:
1810:
1763:
1759:
1714:
1710:
1671:
1667:
1628:
1624:
1571:
1567:
1556:
1552:
1541:
1532:
1509:
1502:
1489:
1485:
1438:
1434:
1403:
1399:
1368:
1364:
1353:
1349:
1326:
1322:
1311:
1307:
1296:
1289:
1250:
1246:
1207:
1200:
1169:
1160:
1113:
1109:
1070:
1063:
1016:
1009:
962:
958:
929:
925:
888:
884:
829:
822:
783:
776:
745:
741:
694:
687:
647:
641:
634:
603:
594:
547:
543:
510:(5): e1004351.
496:
483:
438:Current Biology
427:
418:
413:
381:
315:
277:
269:
215:
108:
83:C-value paradox
58:
32:—mostly,
17:
12:
11:
5:
3364:
3354:
3353:
3337:
3336:
3285:
3244:
3193:
3126:
3067:
3016:
3009:
2991:
2934:
2885:
2834:
2785:
2726:
2675:(April 2013).
2664:
2615:
2596:
2561:
2526:
2469:
2417:
2402:
2387:
2366:10.1086/676588
2337:
2308:(3): 642–645.
2288:
2259:(3): 578–590.
2236:
2198:
2175:
2160:
2141:
2122:
2087:
2030:
1973:
1916:
1859:
1808:
1757:
1708:
1665:
1622:
1565:
1550:
1530:
1500:
1483:
1432:
1397:
1378:(115): 52–54.
1362:
1347:
1336:(3): 107–111.
1320:
1305:
1287:
1244:
1198:
1158:
1107:
1061:
1007:
978:10.1086/282445
966:Optics Express
956:
923:
902:(2): 111–176.
882:
820:
793:(3): 187–207.
774:
755:(6): 807–821.
739:
685:
658:(1): 162–171.
632:
613:(6): 651–662.
592:
561:(4): 634–650.
541:
481:
415:
414:
412:
409:
408:
407:
405:Non-coding RNA
402:
400:Non-coding DNA
397:
392:
387:
385:ENCODE Project
380:
377:
314:
313:Non-functional
311:
276:
273:
268:
265:
214:
211:
151:
150:
147:
144:
141:
118:neutral theory
106:
97:Crafoord Prize
67:Hermann Muller
63:J.B.S. Haldane
57:
54:
46:Non-coding DNA
15:
9:
6:
4:
3:
2:
3363:
3352:
3349:
3348:
3346:
3332:
3328:
3323:
3318:
3313:
3308:
3304:
3300:
3296:
3289:
3281:
3277:
3272:
3267:
3263:
3259:
3255:
3248:
3240:
3236:
3231:
3226:
3221:
3216:
3212:
3208:
3204:
3197:
3189:
3185:
3180:
3175:
3170:
3165:
3161:
3157:
3153:
3149:
3145:
3141:
3137:
3130:
3122:
3118:
3113:
3108:
3103:
3098:
3094:
3090:
3086:
3082:
3078:
3071:
3063:
3059:
3054:
3049:
3044:
3039:
3035:
3031:
3030:PLOS Genetics
3027:
3020:
3012:
3010:9781605354699
3006:
3002:
2995:
2987:
2983:
2978:
2973:
2969:
2965:
2961:
2957:
2953:
2949:
2945:
2938:
2930:
2926:
2921:
2916:
2912:
2908:
2904:
2900:
2896:
2889:
2881:
2877:
2872:
2867:
2862:
2857:
2853:
2849:
2845:
2838:
2830:
2826:
2821:
2816:
2812:
2808:
2804:
2800:
2796:
2789:
2781:
2777:
2772:
2767:
2762:
2757:
2753:
2749:
2746:(33): E3365.
2745:
2741:
2737:
2730:
2722:
2718:
2713:
2708:
2703:
2698:
2694:
2690:
2686:
2682:
2678:
2674:
2668:
2660:
2656:
2651:
2646:
2642:
2638:
2634:
2630:
2626:
2619:
2611:
2607:
2600:
2592:
2588:
2584:
2580:
2576:
2572:
2565:
2557:
2553:
2549:
2545:
2541:
2537:
2530:
2522:
2518:
2513:
2508:
2504:
2500:
2496:
2492:
2488:
2484:
2480:
2473:
2465:
2461:
2456:
2451:
2447:
2443:
2439:
2435:
2431:
2424:
2422:
2413:
2406:
2398:
2391:
2383:
2379:
2375:
2371:
2367:
2363:
2359:
2355:
2348:
2341:
2333:
2329:
2324:
2319:
2315:
2311:
2307:
2303:
2299:
2292:
2284:
2280:
2275:
2270:
2266:
2262:
2258:
2254:
2250:
2243:
2241:
2232:
2228:
2224:
2220:
2216:
2212:
2205:
2203:
2194:
2190:
2186:
2179:
2171:
2164:
2156:
2152:
2145:
2137:
2133:
2126:
2118:
2114:
2110:
2106:
2102:
2098:
2091:
2083:
2079:
2075:
2071:
2066:
2061:
2057:
2053:
2049:
2045:
2041:
2034:
2026:
2022:
2018:
2014:
2009:
2004:
2000:
1996:
1992:
1988:
1984:
1977:
1969:
1965:
1961:
1957:
1952:
1947:
1943:
1939:
1935:
1931:
1927:
1920:
1912:
1908:
1904:
1900:
1895:
1890:
1886:
1882:
1878:
1874:
1870:
1863:
1855:
1851:
1847:
1843:
1839:
1835:
1831:
1827:
1823:
1819:
1812:
1804:
1800:
1796:
1792:
1788:
1784:
1780:
1776:
1772:
1768:
1761:
1753:
1749:
1744:
1739:
1735:
1731:
1727:
1723:
1719:
1712:
1704:
1700:
1696:
1692:
1688:
1684:
1680:
1676:
1669:
1661:
1657:
1653:
1649:
1645:
1641:
1637:
1633:
1626:
1618:
1614:
1610:
1606:
1601:
1596:
1592:
1588:
1585:(5645): 501.
1584:
1580:
1576:
1569:
1561:
1554:
1546:
1539:
1537:
1535:
1526:
1522:
1518:
1514:
1507:
1505:
1496:
1495:
1487:
1479:
1475:
1471:
1467:
1463:
1459:
1455:
1451:
1447:
1443:
1436:
1428:
1424:
1420:
1416:
1412:
1408:
1401:
1393:
1389:
1385:
1381:
1377:
1373:
1366:
1358:
1351:
1343:
1339:
1335:
1331:
1324:
1316:
1309:
1301:
1294:
1292:
1283:
1279:
1275:
1271:
1267:
1263:
1259:
1255:
1248:
1240:
1236:
1232:
1228:
1224:
1220:
1216:
1212:
1205:
1203:
1194:
1190:
1186:
1182:
1178:
1174:
1167:
1165:
1163:
1154:
1150:
1146:
1142:
1138:
1134:
1130:
1126:
1122:
1118:
1111:
1103:
1099:
1095:
1091:
1087:
1083:
1079:
1075:
1068:
1066:
1057:
1053:
1049:
1045:
1041:
1037:
1033:
1029:
1025:
1021:
1014:
1012:
1003:
999:
995:
991:
987:
983:
979:
975:
971:
967:
960:
951:
946:
942:
938:
934:
927:
919:
915:
910:
905:
901:
897:
893:
886:
878:
874:
869:
864:
859:
854:
850:
846:
842:
838:
834:
827:
825:
816:
812:
808:
804:
800:
796:
792:
788:
781:
779:
770:
766:
762:
758:
754:
750:
743:
735:
731:
726:
721:
716:
711:
707:
703:
699:
692:
690:
681:
677:
673:
669:
665:
661:
657:
653:
646:
639:
637:
628:
624:
620:
616:
612:
608:
601:
599:
597:
588:
584:
579:
574:
569:
564:
560:
556:
552:
545:
537:
533:
528:
523:
518:
513:
509:
505:
504:PLOS Genetics
501:
494:
492:
490:
488:
486:
477:
473:
469:
465:
460:
455:
451:
447:
443:
439:
435:
431:
425:
423:
421:
416:
406:
403:
401:
398:
396:
393:
391:
388:
386:
383:
382:
376:
372:
370:
366:
361:
358:
353:
351:
347:
343:
339:
335:
331:
326:
322:
320:
310:
308:
303:
301:
297:
293:
289:
285:
281:
272:
263:
259:
255:
250:
247:
244:
240:
239:David Comings
234:
229:
225:
221:
218:
209:
205:
203:
197:
193:
191:
185:
183:
177:
175:
167:
166:Francis Crick
163:
159:
156:
148:
145:
142:
139:
135:
134:
133:
130:
125:
123:
119:
113:
110:
98:
94:
90:
86:
84:
78:
74:
72:
68:
64:
53:
49:
47:
41:
39:
35:
31:
27:
23:
19:
3302:
3298:
3288:
3261:
3257:
3247:
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3000:
2994:
2951:
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2905:(1): 36–46.
2902:
2898:
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2851:
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2802:
2798:
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2729:
2684:
2680:
2673:Doolittle WF
2667:
2632:
2628:
2618:
2610:The Observer
2609:
2599:
2574:
2570:
2564:
2539:
2535:
2529:
2486:
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2472:
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2433:
2411:
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2396:
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2357:
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2305:
2301:
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2256:
2252:
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2178:
2169:
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2154:
2150:
2144:
2135:
2125:
2100:
2096:
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2033:
1990:
1986:
1976:
1933:
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1811:
1770:
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1760:
1725:
1721:
1711:
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1375:
1371:
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610:
606:
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544:
507:
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437:
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373:
362:
354:
338:Alu elements
327:
323:
316:
304:
287:
283:
278:
270:
261:
257:
252:
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231:
227:
223:
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216:
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186:
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2103:: 247–278.
1519:: 366–370.
1179:: 237–256.
943:: 267–273.
300:centromeres
174:transposons
129:Susumu Ohno
93:Tomoko Ohta
38:transposons
34:pseudogenes
3207:Cell Cycle
2854:(1): 412.
2195:: 340–342.
2157:: 174–182.
708:: 831068.
411:References
348:-Alu) and
288:non-coding
284:functional
275:Functional
109:t analysis
3258:BioEssays
937:Hereditas
815:254768457
769:254277794
3345:Category
3331:25674102
3280:37318305
3239:23759593
3188:35859178
3121:37104599
3112:10250106
3062:25057982
2986:32460305
2929:22124482
2880:32957925
2829:24814287
2780:25107292
2721:23479647
2659:23578867
2591:26152174
2556:22955811
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2382:14549993
2374:24921597
2332:25635041
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2082:27111068
2025:31899622
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1153:13344748
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1002:84202145
994:24718213
918:14771033
877:24753594
807:36528415
734:35251134
680:27613442
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587:24705080
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379:See also
254:studies.
120:and the
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3322:4306305
3230:4685169
3179:9329122
3148:Bibcode
3081:Science
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2977:7547914
2956:Bibcode
2920:3324860
2871:7507296
2820:4041003
2771:4143013
2748:Bibcode
2712:3619371
2689:Bibcode
2637:Bibcode
2536:Science
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2491:Bibcode
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2323:5322545
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2074:7393317
2052:Bibcode
2017:7453799
1995:Bibcode
1968:8938434
1960:6256636
1938:Bibcode
1911:4261755
1903:7393318
1881:Bibcode
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1846:7366731
1826:Bibcode
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1795:6245369
1775:Bibcode
1752:4162765
1730:Bibcode
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1640:Bibcode
1632:Science
1617:4216649
1587:Bibcode
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1450:Bibcode
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1282:5789433
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1254:Science
1239:4874239
1219:Bibcode
1211:Science
1125:Bibcode
1102:5767777
1082:Bibcode
1074:Science
1056:4161261
1048:5637732
1028:Bibcode
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909:1716299
868:4035993
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725:8888898
615:Bibcode
578:3899985
527:4014423
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138:paradox
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1966:
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