993:(especially trans regulatory expression) when removed. Singleton and duplicate capacitors both largely represent instances of incomplete functional redundancy; differentially expressed paralogs of duplicate capacitors continue some functionality of the original gene, and the protein-protein interaction complexes within which singleton capacitors reside largely exhibit overlapping functionality. In general the phenotypic capacitors identified by knockouts in yeast are genes expressed in several key regulatory areas which, while non-lethal when removed, do not have enough redundancy to maintain complete functionality. The concept of functional redundancy may also be involved in the high number of synthetic-lethal interactions which capacitor genes participate in. When a gene has its functionality resumed by a paralog or functional analog, its removal is not inherently lethal, however when the gene and its redundancy are removed, the result is lethality.
956:
36:
591:
972:), while duplicate capacitors are more highly connected and tend to interact with multiple large complexes. The gene ontologies of singleton and duplicate capacitors also differ notably. Singleton capacitors are concentrated in the categories of DNA maintenance and organization, response to stimuli, and RNA transcription and localization, whereas duplicate capacitors are concentrated in the categories of
578:
848:, in matched and strains in a variety of stressful environments. Sometimes the strain grows faster, sometimes : this depends on the genetic background of the strain, suggesting that taps into pre-existing cryptic genetic variation. Mathematical models suggest that may have evolved, as an evolutionary capacitor, to promote
1001:
Computational simulations of knockouts in complex gene interaction networks have demonstrated that many, and possibly all expressed genes have the potential to reveal phenotypic variation of some kind when removed, and that previously identified capacitor genes are simply especially strong examples.
967:
elsewhere in the genome; most capacitors identified in yeast are either singleton genes, or have historical paralogs from which they have diverged substantially in terms of expression. Singleton and duplicate capacitors largely exhibit disjoint behavior in the interactome. Singleton capacitors are
992:
at various levels of the genome. Coding regions that are necessary for the synthesis of key proteins which do not have paralogs elsewhere in the genome are lethal when removed. Conversely, coding regions with many paralogs or strongly expressed paralogs have a minimal effect on overall expression
878:
or genomic regions which function as evolutionary capacitors. When a gene is knocked out, and its removal reveals phenotypic variation that was not previously observable, that gene is functioning as a phenotypic capacitor. If any of the variation is adaptive, it is functioning as an evolutionary
669:
to new environmental conditions. Switching rates may be a function of stress, making genetic variation more likely to affect the phenotype at times when it is most likely to be useful for adaptation. In addition, strongly deleterious variation may be purged while in a partially cryptic state, so
919:. Unlike the chromatin regulators, the removal of genes which code for metabolic enzymes does not have a consistent effect on the difference in expression between species, with different enzyme knockouts either increasing, decreasing, or not significantly affecting the expression difference.
746:. It has been proposed that the presence of chaperones may, by providing additional robustness to errors in folding, allow the exploration of a larger set of genotypes. When chaperones are overworked at times of environmental stress, this may "switch on" previously cryptic genetic variation.
772:). This was thought to prove that the new phenotypes depended on pre-existing cryptic genetic variation that had merely been revealed. More recent evidence suggests that these data might be explained by new mutations caused by the reactivation of formally dormant
951:
observed for its expressed protein. However, proteins with the highest amount of interactions have reduced phenotypic capacitance, possibly due to increased duplication of regions coding these proteins in the genome, reducing the effect of a single knockout.
862:
Similar transient increases in error rates can evolve emergently in the absence of a "widget" like . The primary advantage of a -like widget is to facilitate the subsequent evolution of lower error rates once genetic assimilation has occurred.
688:
Switches that turn robustness to phenotypic rather than genetic variation on and off do not fit the capacitance analogy, as their presence does not cause variation to accumulate over time. They have instead been called phenotypic stabilizers.
658:. After that, the rest of variation, most of which is presumably deleterious, can be switched off, leaving the population with a newly evolved advantageous trait, but no long-term handicap. For evolutionary capacitance to increase
859:, mimicking the effects of , than would be expected from mutation bias or than are observed in other taxa that do not form the prion. These observations are compatible with acting as an evolutionary capacitor in the wild.
922:
Broader knockout samples in yeast have identified at least 300 genes which, when absent, increase morphological variation between yeast individuals. These capacitor genes predominantly occupy a few key domains in
654:. An evolutionary capacitor is a molecular switch mechanism that can "toggle" genetic variation between hidden and revealed states. If some subset of newly revealed variation is adaptive, it becomes fixed by
840:
form (). When is present, this depletes the amount of normal Sup35p available. As a result, the rate of errors in which translation continues beyond a stop codon increases from about 0.3% to about 1%.
1555:
Specchia V; Piacentini L; Tritto P; Fanti L; D’Alessandro R; Palumbo G; Pimpinelli S; Bozzetti MP (2010). "Hsp90 prevents phenotypic variation by suppressing the mutagenic activity of transposons".
707:
perform side reactions. Similarly, binding proteins may spend some proportion of their time bound to off-target proteins. These reactions or interactions may be of no consequence to current
764:, a broad range of different phenotypes are seen, where the identity of the phenotype depends on the genetic background. Also, a recent study on the model insect, the red flour beetle
307:
681:
about the extent to which capacitance might contribute to evolution in natural populations. The possibility of evolutionary capacitance is considered to be part of the
723:
resistance. In populations exposed only to ampicillin, such mutations may be present in a minority of members since there is not fitness cost (i.e. are within the
650:
effect. But when the system is disturbed (perhaps by stress), robustness breaks down, and the variation has phenotypic effects and is subject to the full force of
622:
1466:
Matsumura, I; Ellington, AD (Jan 12, 2001). "In vitro evolution of beta-glucuronidase into a beta-galactosidase proceeds through non-specific intermediates".
230:
959:
Singleton capacitors (light blue) are generally part of large complexes, while duplicate capacitors (dark blue) often interact with several major complexes.
541:
911:
regulating genes increases the differences in expression between yeast species. The majority of the variation in protein expression is attributable to
727:). This represents cryptic genetic variation since if the population is newly exposed to cefotaxime, the minority members will exhibit some resistance.
711:
but under altered conditions, may provide the starting point for adaptive evolution. For example, several mutations in the antibiotic resistance gene
2427:
349:
1906:"The Spontaneous Appearance Rate of the Yeast Prion PSI+ and Its Implications for the Evolution of the Evolvability Properties of the PSI+ System"
1767:
Firoozan M, Grant CM, Duarte JA, Tuite MF (1991). "Quantitation of readthrough of termination codons in yeast using a novel gene fusion assay".
55:
776:. However, this finding regarding transposable elements may be dependent on the strong nature of the Hsp90 knockdown used in that experiment.
768:, showed that Hsp90 impairment revealed a new phenotype, reduced-eye phenotype, which was stably inherited without further HSP90 inhibition (
536:
483:
939:, and response to stimuli such as stress. More generally, capacitor genes are likely to express proteins which act as network hubs in the
615:
344:
175:
1390:
Mohamed, MF; Hollfelder, F (Jan 2013). "Efficient, crosswise catalytic promiscuity among enzymes that catalyze phosphoryl transfer".
670:
cryptic variation that remains is more likely to be adaptive than random mutations are. Capacitance can help cross "valleys" in the
77:
932:
682:
334:
302:
1018:
556:
339:
891:. While some of the variation revealed by these knockouts is deleterious, other variation has a relatively minor effect on
2422:
1033:
can act as an evolutionary capacitor by breaking up allele combinations with phenotypic effects that normally cancel out.
955:
608:
595:
948:
526:
1716:
Nobuhiko
Tokuriki; Dan S. Tawfik (2009). "Chaperonin overexpression promotes genetic variation and enzyme evolution".
724:
582:
754:
The hypothesis that chaperones can act as evolutionary capacitors is closely associated with the heat shock protein
1810:
True HL, Lindquist SL (2000). "A yeast prion provides a mechanism for genetic variation and phenotypic diversity".
947:
interactions. The confidence that a specific gene acts as a phenotypic capacitor is correlated with the number of
82:
638:
is the storage and release of variation, just as electric capacitors store and release charge. Living systems are
448:
423:
403:
383:
60:
2323:"The consequences of rare sexual reproduction by means of selfing in an otherwise clonally reproducing species"
1242:"Cryptic genetic variation can make "irreducible complexity" a common mode of adaptation in sexual populations"
1002:
Capacitance, then, is simply a feature of complex gene networks that arises in conjunction with canalization.
912:
904:
825:
438:
433:
408:
363:
329:
323:
312:
2270:
Bergman A, Siegal ML (July 2003). "Evolutionary capacitance as a general feature of complex gene networks".
1042:
928:
916:
888:
760:
551:
458:
453:
398:
359:
160:
855:
appears more frequently in response to environmental stress. In yeast, more stop codon disappearances are
833:
463:
418:
240:
135:
2143:
Kazuo
Takahashi (2013). "Multiple capacitors for natural genetic variation in Drosophila melanogaster".
988:
The mechanism of phenotypic capacitor genes in yeast appears to be closely related to the modalities of
795:
674:, where a combination of two mutations would be beneficial, even though each is deleterious on its own.
639:
561:
443:
388:
354:
267:
2279:
1819:
1725:
1672:
1564:
1513:
1026:
845:
678:
655:
413:
185:
70:
27:
944:
765:
743:
735:
503:
493:
428:
393:
277:
180:
105:
40:
2432:
2399:
2391:
2303:
1886:
1843:
1792:
1749:
1698:
1588:
1537:
1320:
1164:
1022:
989:
973:
698:
662:
in this way, the switching rate should not be faster than the timescale of genetic assimilation.
546:
478:
262:
190:
155:
1861:
Masel J, Bergman A (2003). "The evolution of the evolvability properties of the yeast prion ".
2352:
2295:
2252:
2201:
2125:
2076:
2035:
1986:
1935:
1878:
1835:
1784:
1741:
1690:
1659:
Mario A. Fares; Mario X. Ruiz-González; Andrés Moya; Santiago F. Elena; Eladio Barrio (2002).
1641:
1580:
1529:
1483:
1448:
1407:
1372:
1312:
1271:
1219:
1156:
1112:
964:
708:
671:
651:
643:
247:
125:
115:
110:
35:
2383:
2342:
2334:
2287:
2242:
2232:
2191:
2183:
2152:
2115:
2107:
2066:
2025:
2017:
1976:
1966:
1925:
1917:
1870:
1827:
1776:
1733:
1680:
1631:
1623:
1572:
1521:
1475:
1438:
1399:
1362:
1354:
1302:
1261:
1253:
1209:
1201:
1146:
1102:
1092:
969:
790:
282:
2055:"Evolutionary Capacitance Emerges Spontaneously during Adaptation to Environmental Changes"
1240:; Weissman, Daniel B.; Peterson, Grant I.; Peck, Kayla M.; Masel, Joanna (December 2014).
1057:
887:
Deficiency in at least 15 different genes reveals cryptic variation in wing morphology in
739:
235:
225:
87:
2371:
2283:
1823:
1729:
1676:
1568:
1517:
2347:
2322:
2247:
2220:
2196:
2171:
2120:
2095:
2030:
2005:
1981:
1954:
1930:
1905:
1874:
1636:
1611:
1504:
Rutherford SL, Lindquist S (1998). "Hsp90 as a capacitor for morphological evolution".
1367:
1342:
1266:
1241:
1214:
1189:
1107:
1080:
317:
220:
1443:
1426:
2416:
2111:
1753:
1541:
1307:
1151:
1134:
1052:
924:
871:
856:
837:
468:
140:
1890:
1796:
1168:
2403:
1847:
1702:
1592:
1338:
1324:
1237:
1185:
1076:
1030:
892:
849:
799:
659:
488:
473:
257:
252:
170:
2307:
2172:"Chromatin regulators as capacitors of interspecies variations in gene expression"
836:
to stop correctly at the ends of proteins. Sup35p comes in a normal form () and a
2237:
2071:
2054:
1971:
1612:"Drosophila Piwi functions in Hsp90-mediated suppression of phenotypic variation"
1403:
1921:
1205:
1047:
977:
940:
712:
498:
215:
165:
1013:
mutations can be thought of as cryptic when they are present overwhelmingly in
2338:
1607:
1358:
936:
829:
773:
720:
716:
666:
272:
195:
150:
130:
44:
2170:
Itay Tirosh; Sharon
Reikhav; Nadejda Sigal; Yael Assia; Naama Barkai (2010).
2096:"The Evolution of Reversible Switches in the Presence of Irreversible Mimics"
1097:
915:
effects, suggesting that trans-regulatory processes are strongly involved in
2021:
1010:
908:
647:
531:
145:
65:
2356:
2299:
2256:
2205:
2129:
2080:
2039:
1990:
1939:
1882:
1839:
1745:
1694:
1645:
1584:
1487:
1479:
1452:
1411:
1376:
1316:
1275:
1223:
1160:
1116:
1029:
population by creating homozygotes. Facultative sex that takes the form of
1788:
1780:
1606:
Vamsi K Gangaraju; Hang Yin; Molly M Weiner; Jianquan Wang; Xiao A Huang;
1533:
577:
2221:"Network hubs buffer environmental variation in Saccharomyces cerevisiae"
1014:
521:
120:
2291:
2187:
1737:
1576:
2395:
704:
2156:
1257:
1831:
1661:"Endosymbiotic bacteria: GroEL buffers against deleterious mutations"
1427:"Catalytic promiscuity and the evolution of new enzymatic activities"
821:
1685:
1660:
1290:
2387:
1627:
1525:
954:
895:, and could even improve the flight capability of an individual.
785:
755:
742:. The need to fold proteins correctly is a big restriction on the
1190:"Cryptic Genetic Variation Is Enriched for Potential Adaptations"
844:
This can lead to different growth rates, and sometimes different
769:
1953:
Tyedmers J, Madariaga ML, Lindquist S (2008). Weissman J (ed.).
968:
most often part of highly interconnected complexes (such as the
875:
1135:"Rate of adaptive peak shifts with partial genetic robustness"
1392:
Biochimica et
Biophysica Acta (BBA) - Proteins and Proteomics
642:
to mutations. This means that living systems accumulate
927:, including chromosome organization and DNA integrity,
1955:"Prion Switching in Response to Environmental Stress"
2094:
Lancaster, Alex K.; Masel, Joanna (September 2009).
1025:
can act as an evolutionary capacitor in a primarily
1904:Lancaster AK, Bardill JP, True HL, Masel J (2010).
1499:
1497:
1291:"Do We Need an Extended Evolutionary Synthesis?"
2006:"The conversion of 3′ UTRs into coding regions"
758:. When Hsp90 is downregulated in the fruit fly
2219:Levy SF, Siegal ML (2008). Levchenko A (ed.).
616:
8:
2053:Nelson, Paul; Masel, Joanna (October 2018).
1180:
1178:
703:In addition to their native reaction, many
2372:"Phenotypic evolution and parthenogenesis"
2004:Giacomelli M, Hancock AS, Masel J (2007).
1128:
1126:
623:
609:
18:
2346:
2246:
2236:
2195:
2119:
2070:
2029:
1980:
1970:
1929:
1684:
1635:
1442:
1366:
1343:"Robustness: mechanisms and consequences"
1306:
1265:
1213:
1150:
1106:
1096:
963:Capacitor genes are less likely to have
1068:
26:
1425:O'Brien, PJ; Herschlag, D (Apr 1999).
665:This mechanism would allow for rapid
7:
1081:"Q&A: Evolutionary capacitance"
1875:10.1111/j.0014-3820.2003.tb00358.x
14:
943:of a cell, and in the network of
2112:10.1111/j.1558-5646.2009.00729.x
1308:10.1111/j.1558-5646.2007.00246.x
1152:10.1111/j.1558-5646.2007.00166.x
828:protein involved in recognising
590:
589:
576:
34:
2428:Extended evolutionary synthesis
2010:Molecular Biology and Evolution
683:extended evolutionary synthesis
646:without the variation having a
583:Evolutionary biology portal
16:Evolutionary biology hypothesis
2327:Theoretical Population Biology
874:can be used to identify novel
770:https://doi.org/10.1101/690727
744:evolution of protein sequences
542:Creation–evolution controversy
296:History of evolutionary theory
1:
1444:10.1016/s1074-5521(99)80033-7
719:resistance but do not affect
2238:10.1371/journal.pbio.0060264
2072:10.1016/j.celrep.2018.09.008
1972:10.1371/journal.pbio.0060294
1468:Journal of Molecular Biology
1404:10.1016/j.bbapap.2012.07.015
949:protein-protein interactions
527:Evolution as fact and theory
2370:Lynch M, Gabriel W (1983).
2321:Masel J, Lyttle DN (2011).
1922:10.1534/genetics.109.110213
1289:Pigliucci, Massimo (2007).
1206:10.1534/genetics.105.051649
2449:
907:, the knockout of certain
696:
562:Nature-nurture controversy
2339:10.1016/j.tpb.2011.08.004
2176:Molecular Systems Biology
1359:10.1016/j.tig.2009.07.005
1017:rather than homozygotes.
449:Evolutionary neuroscience
424:Evolutionary epistemology
404:Evolutionary anthropology
384:Applications of evolution
2378:(Submitted manuscript).
1098:10.1186/1741-7007-11-103
636:Evolutionary capacitance
439:Evolutionary linguistics
434:Evolutionary game theory
409:Evolutionary computation
1431:Chemistry & Biology
1043:Canalization (genetics)
1021:that takes the form of
889:Drosophila melanogaster
761:Drosophila melanogaster
552:Objections to evolution
459:Evolutionary psychology
454:Evolutionary physiology
399:Evolutionary aesthetics
378:Fields and applications
360:History of paleontology
1480:10.1006/jmbi.2000.4259
960:
784:The overproduction of
677:There is currently no
484:Speciation experiments
464:Experimental evolution
419:Evolutionary economics
241:Recent human evolution
99:Processes and outcomes
2022:10.1093/molbev/msl172
1781:10.1002/yea.320070211
990:functional redundancy
958:
796:mutational robustness
774:transposable elements
444:Evolutionary medicine
389:Biosocial criminology
355:History of speciation
268:Evolutionary taxonomy
231:Timeline of evolution
2423:Evolutionary biology
1341:; Siegal ML (2009).
1238:Trotter, Meredith V.
933:protein modification
798:. This can increase
656:genetic assimilation
414:Evolutionary ecology
28:Evolutionary biology
2376:American Naturalist
2292:10.1038/nature01765
2284:2003Natur.424..549B
2188:10.1038/msb.2010.84
1824:2000Natur.407..477T
1738:10.1038/nature08009
1730:2009Natur.459..668T
1677:2002Natur.417..398F
1577:10.1038/nature08739
1569:2010Natur.463..662S
1518:1998Natur.396..336R
766:Tribolium castaneum
516:Social implications
504:Universal Darwinism
494:Island biogeography
429:Evolutionary ethics
394:Ecological genetics
340:Molecular evolution
278:Transitional fossil
106:Population genetics
22:Part of a series on
2151:(435): 1356–1365.
1347:Trends in Genetics
974:protein metabolism
961:
699:enzyme promiscuity
693:Enzyme promiscuity
547:Theistic evolution
479:Selective breeding
191:Parallel evolution
156:Adaptive radiation
2278:(6948): 549–552.
2157:10.1111/mec.12091
2145:Molecular Ecology
1818:(6803): 477–483.
1724:(7247): 668–673.
1512:(6709): 336–342.
1301:(12): 2743–2749.
1258:10.1111/evo.12517
1252:(12): 3357–3367.
672:fitness landscape
652:natural selection
644:genetic variation
633:
632:
324:Origin of Species
126:Natural selection
2440:
2408:
2407:
2367:
2361:
2360:
2350:
2318:
2312:
2311:
2267:
2261:
2260:
2250:
2240:
2216:
2210:
2209:
2199:
2167:
2161:
2160:
2140:
2134:
2133:
2123:
2106:(9): 2350–2362.
2091:
2085:
2084:
2074:
2050:
2044:
2043:
2033:
2001:
1995:
1994:
1984:
1974:
1950:
1944:
1943:
1933:
1901:
1895:
1894:
1869:(7): 1498–1512.
1858:
1852:
1851:
1832:10.1038/35035005
1807:
1801:
1800:
1764:
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1757:
1713:
1707:
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1335:
1329:
1328:
1310:
1286:
1280:
1279:
1269:
1234:
1228:
1227:
1217:
1200:(3): 1985–1991.
1182:
1173:
1172:
1154:
1145:(8): 1847–1856.
1130:
1121:
1120:
1110:
1100:
1079:(Sep 30, 2013).
1073:
970:mediator complex
945:synthetic-lethal
818:
817:
813:
791:Escherichia coli
625:
618:
611:
598:
593:
592:
585:
581:
580:
557:Level of support
350:Current research
335:Modern synthesis
330:Before synthesis
283:Extinction event
41:Darwin's finches
38:
19:
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2003:
2002:
1998:
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1766:
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1715:
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1686:10.1038/417398a
1658:
1657:
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1616:Nature Genetics
1605:
1604:
1600:
1554:
1553:
1549:
1503:
1502:
1495:
1465:
1464:
1460:
1437:(4): R91–R105.
1424:
1423:
1419:
1389:
1388:
1384:
1337:
1336:
1332:
1288:
1287:
1283:
1236:
1235:
1231:
1184:
1183:
1176:
1132:
1131:
1124:
1075:
1074:
1070:
1066:
1058:Susan Lindquist
1039:
1019:Facultative sex
1008:
1006:Facultative sex
999:
986:
901:
885:
869:
819:
815:
811:
809:
808:
782:
752:
740:protein folding
733:
725:neutral network
701:
695:
629:
588:
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574:
567:
566:
517:
509:
508:
379:
371:
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369:
297:
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236:Human evolution
226:History of life
210:
209:Natural history
202:
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100:
92:
47:
17:
12:
11:
5:
2446:
2444:
2436:
2435:
2430:
2425:
2415:
2414:
2410:
2409:
2388:10.1086/284169
2382:(6): 745–764.
2362:
2333:(4): 317–322.
2313:
2262:
2211:
2162:
2135:
2086:
2065:(1): 249–258.
2045:
2016:(2): 457–464.
1996:
1945:
1916:(2): 393–400.
1896:
1853:
1802:
1775:(2): 173–183.
1759:
1708:
1651:
1628:10.1038/ng.743
1622:(2): 153–158.
1598:
1563:(1): 662–665.
1547:
1493:
1458:
1417:
1382:
1353:(9): 395–403.
1330:
1281:
1229:
1188:(March 2006).
1174:
1133:Kim Y (2007).
1122:
1067:
1065:
1062:
1061:
1060:
1055:
1050:
1045:
1038:
1035:
1007:
1004:
998:
995:
985:
982:
929:RNA elongation
900:
897:
884:
881:
872:Gene knockouts
868:
867:Gene knockouts
865:
807:
804:
781:
778:
751:
748:
732:
729:
697:Main article:
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537:Social effects
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263:Classification
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221:Common descent
218:
216:Origin of life
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2027:
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2019:
2015:
2011:
2007:
2000:
1997:
1992:
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1983:
1978:
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1968:
1964:
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1927:
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1911:
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1712:
1709:
1704:
1700:
1696:
1692:
1687:
1682:
1678:
1674:
1671:(6887): 398.
1670:
1666:
1662:
1655:
1652:
1647:
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1629:
1625:
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1617:
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1401:
1398:(1): 417–24.
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1378:
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1186:Masel, Joanna
1181:
1179:
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1053:Preadaptation
1051:
1049:
1046:
1044:
1041:
1040:
1036:
1034:
1032:
1028:
1024:
1020:
1016:
1015:heterozygotes
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1005:
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994:
991:
983:
981:
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925:gene ontology
920:
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797:
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469:Phylogenetics
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348:
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328:
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321:
319:
316:
314:
313:Before Darwin
311:
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141:Genetic drift
139:
137:
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122:
119:
117:
114:
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109:
107:
104:
103:
96:
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50:
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42:
37:
33:
32:
29:
25:
21:
20:
2379:
2375:
2365:
2330:
2326:
2316:
2275:
2271:
2265:
2228:
2225:PLOS Biology
2224:
2214:
2182:(435): 435.
2179:
2175:
2165:
2148:
2144:
2138:
2103:
2099:
2089:
2062:
2059:Cell Reports
2058:
2048:
2013:
2009:
1999:
1965:(11): e294.
1962:
1959:PLOS Biology
1958:
1948:
1913:
1909:
1899:
1866:
1862:
1856:
1815:
1811:
1805:
1772:
1768:
1762:
1721:
1717:
1711:
1668:
1664:
1654:
1619:
1615:
1601:
1560:
1556:
1550:
1509:
1505:
1474:(2): 331–9.
1471:
1467:
1461:
1434:
1430:
1420:
1395:
1391:
1385:
1350:
1346:
1333:
1298:
1294:
1284:
1249:
1245:
1232:
1197:
1193:
1142:
1138:
1088:
1084:
1071:
1009:
1000:
987:
962:
921:
917:canalization
902:
893:aerodynamics
886:
879:capacitor.
870:
861:
854:
850:evolvability
846:morphologies
843:
832:and causing
820:
806:Yeast prion
800:evolvability
789:
783:
759:
753:
734:
702:
687:
676:
664:
660:evolvability
635:
634:
489:Sociobiology
474:Paleontology
322:
258:Biogeography
253:Biodiversity
171:Coextinction
161:Co-operation
136:Polymorphism
61:Introduction
2231:(1): e264.
1085:BMC Biology
1048:Epigenetics
1031:outcrossing
997:Simulations
978:endocytosis
941:interactome
883:Fruit Flies
834:translation
830:stop codons
713:B-lactamase
499:Systematics
308:Renaissance
186:Convergence
176:Contingency
166:Coevolution
2417:Categories
1608:Haifan Lin
1064:References
984:Redundancy
937:cell cycle
794:increases
738:assist in
736:Chaperones
731:Chaperones
721:ampicillin
717:cefotaxime
715:introduce
667:adaptation
648:phenotypic
273:Cladistics
196:Extinction
181:Divergence
151:Speciation
131:Adaptation
45:John Gould
2433:Selection
2100:Evolution
1863:Evolution
1754:205216739
1542:204996106
1295:Evolution
1246:Evolution
1139:Evolution
1011:Recessive
909:chromatin
679:consensus
532:Dysgenics
248:Phylogeny
146:Gene flow
116:Diversity
111:Variation
2357:21888925
2300:12891357
2257:18986213
2206:21119629
2130:19486147
2081:30282033
2040:17099057
1991:19067491
1940:19917766
1910:Genetics
1891:30954684
1883:12940355
1840:11028992
1797:42869007
1746:19494908
1695:12024205
1646:21186352
1610:(2011).
1585:20062045
1488:11124909
1453:10099128
1412:22885024
1377:19717203
1317:17924956
1276:25178652
1224:16387877
1194:Genetics
1169:13150906
1161:17683428
1117:24228631
1077:Masel, J
1037:See also
965:paralogs
857:in-frame
596:Category
522:Eugenics
364:timeline
345:Evo-devo
303:Overview
121:Mutation
83:Evidence
78:Glossary
2404:5505336
2396:2460915
2348:3218209
2280:Bibcode
2248:2577700
2197:3010112
2121:2770902
2031:1808353
1982:2586387
1931:2828720
1848:4411231
1820:Bibcode
1789:1905859
1726:Bibcode
1703:4368351
1673:Bibcode
1637:3443399
1593:4429205
1565:Bibcode
1534:9845070
1514:Bibcode
1368:2770586
1339:Masel J
1325:2703146
1267:4258170
1215:1456269
1108:3849687
1091:: 103.
1027:asexual
1023:selfing
709:fitness
705:enzymes
88:History
71:Outline
2402:
2394:
2355:
2345:
2308:775036
2306:
2298:
2272:Nature
2255:
2245:
2204:
2194:
2128:
2118:
2079:
2038:
2028:
1989:
1979:
1938:
1928:
1889:
1881:
1846:
1838:
1812:Nature
1795:
1787:
1752:
1744:
1718:Nature
1701:
1693:
1665:Nature
1644:
1634:
1591:
1583:
1557:Nature
1540:
1532:
1506:Nature
1486:
1451:
1410:
1375:
1365:
1323:
1315:
1274:
1264:
1222:
1212:
1167:
1159:
1115:
1105:
822:Sup35p
810:": -->
640:robust
594:
318:Darwin
2400:S2CID
2392:JSTOR
2304:S2CID
1887:S2CID
1844:S2CID
1793:S2CID
1769:Yeast
1750:S2CID
1699:S2CID
1589:S2CID
1538:S2CID
1321:S2CID
1165:S2CID
913:trans
905:yeast
899:Yeast
876:genes
838:prion
826:yeast
824:is a
786:GroEL
780:GroEL
756:Hsp90
750:Hsp90
56:Index
2353:PMID
2296:PMID
2253:PMID
2202:PMID
2126:PMID
2077:PMID
2036:PMID
1987:PMID
1936:PMID
1879:PMID
1836:PMID
1785:PMID
1742:PMID
1691:PMID
1642:PMID
1581:PMID
1530:PMID
1484:PMID
1449:PMID
1408:PMID
1396:1834
1373:PMID
1313:PMID
1272:PMID
1220:PMID
1157:PMID
1113:PMID
976:and
812:edit
66:Main
2384:doi
2380:122
2343:PMC
2335:doi
2288:doi
2276:424
2243:PMC
2233:doi
2192:PMC
2184:doi
2153:doi
2116:PMC
2108:doi
2067:doi
2026:PMC
2018:doi
1977:PMC
1967:doi
1926:PMC
1918:doi
1914:184
1871:doi
1828:doi
1816:407
1777:doi
1734:doi
1722:459
1681:doi
1669:417
1632:PMC
1624:doi
1573:doi
1561:463
1522:doi
1510:396
1476:doi
1472:305
1439:doi
1400:doi
1363:PMC
1355:doi
1303:doi
1262:PMC
1254:doi
1210:PMC
1202:doi
1198:172
1147:doi
1103:PMC
1093:doi
980:.
903:In
788:in
43:by
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