199:
938:. It also contains a promoter which binds to RNA polymerase and an operator which blocks transcription when bound to the protein synthesized by the repressor gene (trp R) that binds to the operator. In the lac operon, lactose binds to the repressor protein and prevents it from repressing gene transcription, while in the trp operon, tryptophan binds to the repressor protein and enables it to repress gene transcription. Also unlike the lac operon, the trp operon contains a leader peptide and an
410:
1013:
999:
613:
657:: The gene is turned on. Lactose is inhibiting the repressor, allowing the RNA polymerase to bind with the promoter, and express the genes, which synthesize lactase. Eventually, the lactase will digest all of the lactose, until there is none to bind to the repressor. The repressor will then bind to the operator, stopping the manufacture of lactase.
534:; whereas operons contain a set of genes regulated by the same operator, regulons contain a set of genes under regulation by a single regulatory protein, and stimulons contain a set of genes under regulation by a single cell stimulus. According to its authors, the term "operon" is derived from the verb "to operate".
970:
Operon prediction is even more accurate if the functional class of the molecules is considered. Bacteria have clustered their reading frames into units, sequestered by co-involvement in protein complexes, common pathways, or shared substrates and transporters. Thus, accurate prediction would involve
581:
of an operon are turned ON or OFF together, due to a single promoter and operator upstream to them, but sometimes more control over the gene expression is needed. To achieve this aspect, some bacterial genes are located near together, but there is a specific promoter for each of them; this is called
152:
located before the first gene. Later, it was discovered that genes could be positively regulated and also regulated at steps that follow transcription initiation. Therefore, it is not possible to talk of a general regulatory mechanism, because different operons have different mechanisms. Today, the
963:
One prediction method uses the intergenic distance between reading frames as a primary predictor of the number of operons in the genome. The separation merely changes the frame and guarantees that the read through is efficient. Longer stretches exist where operons start and stop, often up to 40–50
792:, but they are unable to bind to the operator in their normal conformation. However, certain molecules called corepressors are bound by the repressor protein, causing a conformational change to the active site. The activated repressor protein binds to the operator and prevents transcription. The
1891:
Toledo-Arana A, Dussurget O, Nikitas G, Sesto N, Guet-Revillet H, Balestrino D, Loh E, Gripenland J, Tiensuu T, Vaitkevicius K, Barthelemy M, Vergassola M, Nahori MA, Soubigou G, Régnault B, Coppée JY, Lecuit M, Johansson J, Cossart P (June 2009). "The
Listeria transcriptional landscape from
31:
586:. Usually these genes encode proteins which will work together in the same pathway, such as a metabolic pathway. Gene clustering helps a prokaryotic cell to produce metabolic enzymes in a correct order. In one study, it has been posited that in the
702:
operon is located slightly downstream of the promoter; two additional operators, O2 and O3 are located at -82 and +412, respectively. In the case of a repressor, the repressor protein physically obstructs the RNA polymerase from transcribing the
153:
operon is simply defined as a cluster of genes transcribed into a single mRNA molecule. Nevertheless, the development of the concept is considered a landmark event in the history of molecular biology. The first operon to be described was the
506:
and regulated by a common operator. It is defined as a set of adjacent structural genes, plus the adjacent regulatory signals that affect transcription of the structural genes. The regulators of a given operon, including
823:
is bound by the activator protein, it undergoes a change in conformation so that it can bind to the DNA and activate transcription. Examples of positive inducible operons include the MerR family of transcriptional
653:: The gene is essentially turned off. There is no lactose to inhibit the repressor, so the repressor binds to the operator, which obstructs the RNA polymerase from binding to the promoter and making lactase.
680:, which then initiates transcription. In RNA synthesis, promoters indicate which genes should be used for messenger RNA creation – and, by extension, control which proteins the cell produces.
519:, are not necessarily coded for by that operon. The location and condition of the regulators, promoter, operator and structural DNA sequences can determine the effects of common mutations.
1182:
Spieth J, Brooke G, Kuersten S, Lea K, Blumenthal T (May 1993). "Operons in C. elegans: polycistronic mRNA precursors are processed by trans-splicing of SL2 to downstream coding regions".
751:
that enables organisms to regulate the expression of various genes depending on environmental conditions. Operon regulation can be either negative or positive by induction or repression.
144:
in 1960. From this paper, the so-called general theory of the operon was developed. This theory suggested that in all cases, genes within an operon are negatively controlled by a
78:
mRNAs that are translated separately, i.e. several strands of mRNA that each encode a single gene product. The result of this is that the genes contained in the operon are either
2493:
934:), the tryptophan (Trp) operon is inhibited by a chemical (tryptophan). This operon contains five structural genes: trp E, trp D, trp C, trp B, and trp A, which encodes
770:
molecule is present, it binds to the repressor and changes its conformation so that it is unable to bind to the operator. This allows for expression of the operon. The
1133:
Kominek J, Doering DT, Opulente DA, Shen XX, Zhou X, DeVirgilio J, Hulfachor AB, Groenewald M, Mcgee MA, Karlen SD, Kurtzman CP, Rokas A, Hittinger CT (March 2019).
557:). However, the definition of an operon does not require the mRNA to be polycistronic, though in practice, it usually is. Upstream of the structural genes lies a
1393:
1427:
1227:"The Adh-related gene of Drosophila melanogaster is expressed as a functional dicistronic messenger RNA: multigenic transcription in higher organisms"
967:
An alternative method to predict operons is based on finding gene clusters where gene order and orientation is conserved in two or more genomes.
198:
766:, a regulatory repressor protein is normally bound to the operator, which prevents the transcription of the genes on the operon. If an
109:
was shown in the early 1990s, and are considered to be rare. In general, expression of prokaryotic operons leads to the generation of
2441:
1986:
735:
can bind to the repressor to allow its binding to the operator site. A good example of this type of regulation is seen for the
3318:
167:
2609:
2132:
1767:
1715:
1529:
1114:
1087:
904:(from above: negative inducible) model. So it is a negative inducible operon induced by presence of lactose or allolactose.
3283:
2364:
983:. The 517 polycistronic operons are listed in a 2009 study describing the global changes in transcription that occur in
1946:
3128:
2856:
2473:
1026:
900:. Lactose binds to the repressor protein and prevents it from repressing gene transcription. This is an example of the
868:
was the first operon to be discovered and provides a typical example of operon function. It consists of three adjacent
835:
is bound by the activator, it is prevented from binding the DNA. This stops activation and transcription of the system.
728:(small molecule) can displace a repressor (protein) from the operator site (DNA), resulting in an uninhibited operon.
2828:
2581:
3085:
2871:
2866:
2735:
2679:
2336:
1401:
2674:
2389:
1451:
1103:
Lodish H, Zipursky L, Matsudaira P, Baltimore D, Darnel J (2000). "Chapter 9: Molecular
Definition of a Gene".
590:, ribosomal protein coding genes occur in clusters that are less conserved in their organization than in other
3262:
2331:
3113:
2762:
2693:
2434:
2384:
2352:
2326:
2498:
3080:
1979:
1004:
930:
was the first repressible operon to be discovered. While the lac operon can be activated by a chemical (
3272:
3138:
2970:
2713:
2012:
2965:
2209:
2145:
2115:
2093:
141:
3337:
2757:
2545:
2458:
2427:
2003:
1995:
1346:"Displacements of prohead protease genes in the late operons of double-stranded-DNA bacteriophages"
673:
446:
342:
721:. The regulatory gene does not need to be in, adjacent to, or even near the operon to control it.
3206:
3143:
3118:
3001:
2921:
2718:
2535:
2520:
2310:
1999:
1036:
59:
811:
protein stimulates transcription by binding to DNA (usually at a site other than the operator).
3191:
3090:
3075:
3044:
2945:
2723:
2648:
2540:
2219:
1972:
1318:
979:
943:
583:
512:
479:
391:
1077:
3302:
3196:
3123:
3022:
2960:
2955:
2886:
2821:
2638:
2623:
2503:
2403:
2398:
2266:
2168:
939:
935:
877:
698:
as a segment between the promoter and the genes of the operon. The main operator (O1) in the
454:
430:
377:
370:
363:
356:
335:
67:
1518:
175:
3211:
3054:
3049:
2931:
2926:
2913:
2745:
2643:
2561:
2300:
2214:
2180:
2074:
1901:
1796:
1561:
1046:
831:, the activator proteins are normally bound to the pertinent DNA segment. However, when an
808:
516:
565:
to bind and initiate transcription. Close to the promoter lies a section of DNA called an
8:
3307:
2571:
2283:
2251:
2137:
2111:
788:, transcription of the operon normally takes place. Repressor proteins are produced by a
450:
442:
434:
426:
293:
265:
258:
251:
243:
235:
228:
221:
214:
55:
1964:
1905:
1800:
1674:
1647:
1565:
2996:
2706:
2589:
2052:
1925:
1251:
1226:
1207:
1159:
1134:
1041:
977:'s laboratory was the first to experimentally identify all operons of a microorganism,
881:
873:
832:
718:
665:
558:
503:
438:
286:
279:
149:
1623:
1598:
1370:
1345:
803:(which itself acts as the corepressor), is a negatively controlled repressible operon.
3257:
3037:
3032:
2941:
2371:
1917:
1873:
1868:
1843:
1824:
1819:
1784:
1763:
1711:
1707:
1701:
1679:
1628:
1579:
1525:
1498:
1443:
1375:
1361:
1300:
1256:
1199:
1195:
1164:
1110:
1104:
1083:
494:
genes often exist in operons that have been found in a range of eukaryotes including
125:
3241:
2790:
1545:
1211:
171:
3342:
3267:
3236:
3231:
3186:
2814:
2293:
2276:
1929:
1909:
1863:
1855:
1814:
1804:
1732:
1669:
1659:
1618:
1610:
1569:
1488:
1428:"[Operon: a group of genes with the expression coordinated by an operator]"
1365:
1357:
1290:
1246:
1238:
1191:
1154:
1146:
956:
864:
595:
587:
162:
124:
have two operons. The first operon codes for various products, including a special
1430:[Operon: a group of genes with the expression coordinated by an operator]
3277:
3181:
2991:
2767:
2604:
2450:
2157:
1958:
974:
869:
748:
714:
706:
578:
543:
499:
128:
which can bind to and transcribe the second operon. The second operon includes a
79:
1516:
Miller JH, Suzuki DT, Griffiths AJ, Lewontin RC, Wessler SR, Gelbart WM (2005).
960:. As a result, predictions can be made based on an organism's genomic sequence.
140:
The term "operon" was first proposed in a short paper in the
Proceedings of the
2806:
2594:
2566:
2367:
2355:
2305:
1789:
Proceedings of the
National Academy of Sciences of the United States of America
1150:
1056:
1018:
859:
819:, activator proteins are normally unable to bind to the pertinent DNA. When an
789:
677:
602:, the more dispersed is the arrangement of the ribosomal protein coding genes.
562:
1242:
3331:
3245:
3159:
3027:
2861:
2838:
2795:
2482:
2069:
1762:(8th ed.). Upper Saddle River, NJ: Pearson Education. pp. 394–402.
1502:
923:
547:
491:
179:
117:
63:
1859:
1574:
1549:
1493:
1476:
3070:
2891:
2881:
2669:
2599:
2468:
1921:
1877:
1828:
1809:
1683:
1664:
1632:
1583:
1447:
1379:
1304:
1295:
1278:
1168:
1031:
954:
The number and organization of operons has been studied most critically in
901:
777:
is a negatively controlled inducible operon, where the inducer molecule is
71:
1260:
1203:
713:
Not always included within the operon, but important in its function is a
3006:
2986:
2772:
2701:
2256:
2224:
2127:
1782:
931:
897:
778:
732:
415:
314:
307:
1913:
1646:
Tirumalai MR, Sivaraman RV, Kutty LA, Song EL, Fox GE (September 2003).
3250:
2950:
2903:
2730:
2408:
2102:
2057:
2047:
2042:
2037:
2032:
942:
sequence which allows for graded regulation. This is an example of the
917:
849:
800:
793:
771:
736:
695:
669:
485:
467:
328:
321:
154:
90:
1783:
Salgado H, Moreno-Hagelsieb G, Smith TF, Collado-Vides J (June 2000).
1614:
3201:
3169:
2936:
2842:
2064:
888:
operon is regulated by several factors including the availability of
807:
Operons can also be positively controlled. With positive control, an
755:
691:
599:
508:
471:
145:
106:
94:
949:
3164:
3133:
2750:
2740:
2664:
2271:
2261:
1436:
Comptes Rendus
Hebdomadaires des Séances de l'Académie des Sciences
1051:
1012:
527:
495:
475:
300:
272:
121:
102:
39:
2419:
2896:
2486:
2231:
2098:
1890:
1102:
893:
889:
820:
767:
725:
591:
572:
554:
531:
523:
398:
182:
for their discoveries concerning the operon and virus synthesis.
110:
98:
75:
20:
409:
1785:"Operons in Escherichia coli: genomic analyses and predictions"
998:
24:
2508:
2288:
2140:
1515:
612:
129:
113:
mRNAs, while eukaryotic operons lead to monocistronic mRNAs.
1599:"Genomic gene clustering analysis of pathways in eukaryotes"
3174:
2359:
2189:
2120:
1733:"Bacteriology – Chapter Nine Genetic Regulatory Mechanisms"
1645:
1075:
550:
422:
384:
51:
30:
1706:(4th ed.). Oxford: Oxford University Press. pp.
1648:"Ribosomal Protein Cluster Organization in Asgard Archaea"
1524:(8th ed.). San Francisco: W.H. Freeman. p. 740.
1132:
2478:
1994:
687:
349:
47:
1426:
Jacob F, Perrin D, Sanchez C, Monod J (February 1960).
1425:
1272:
1270:
1181:
1841:
16:
Group of open reading frames under the same regulation
1954:
1076:
Sadava DE, Hillis DM, Heller HC, Berenbaum M (2009).
553:(a single mRNA molecule that codes for more than one
1267:
994:
189:
89:
Originally, operons were thought to exist solely in
1438:(Facsimile version reprinted in 2005) (in French).
1517:
1842:Ermolaeva MD, White O, Salzberg SL (March 2001).
1739:. University of South Carolina School of Medicine
1283:Briefings in Functional Genomics & Proteomics
950:Predicting the number and organization of operons
429:controls when expression occurs for the multiple
3329:
2836:
709:– the genes that are co-regulated by the operon.
661:An operon is made up of 3 basic DNA components:
1224:
1468:
1135:"Eukaryotic Acquisition of a Bacterial Operon"
717:, a constantly expressed gene which codes for
573:Operons versus clustering of prokaryotic genes
82:together or not at all. Several genes must be
2822:
2435:
1980:
1596:
537:
1844:"Prediction of operons in microbial genomes"
1757:
1421:
1419:
1343:
971:all of these data, a difficult task indeed.
132:gene meant to cause the host cell to burst.
1477:"Eukaryotic and prokaryotic gene structure"
754:Negative control involves the binding of a
2829:
2815:
2590:Precursor mRNA (pre-mRNA / hnRNA)
2442:
2428:
1987:
1973:
1474:
1276:
758:to the operator to prevent transcription.
116:Operons are also found in viruses such as
1867:
1818:
1808:
1695:
1693:
1673:
1663:
1622:
1573:
1492:
1416:
1369:
1294:
1250:
1158:
1128:
1126:
694:binds. It is classically defined in the
546:which are generally transcribed into one
1082:(9th ed.). Macmillan. p. 349.
611:
29:
1337:
3330:
3319:Index of evolutionary biology articles
1690:
1123:
378:Protein coding region
375:
371:Protein coding region
368:
361:
354:
350:DNA
333:
326:
319:
312:
305:
168:Nobel Prize in Physiology and Medicine
70:together in the cytoplasm, or undergo
2810:
2610:Histone acetylation and deacetylation
2423:
2133:Histone acetylation and deacetylation
1968:
1699:
1544:
396:
389:
382:
347:
340:
298:
291:
284:
277:
270:
263:
256:
240:
233:
219:
212:
2675:Ribosome-nascent chain complex (RNC)
1475:Shafee, Thomas; Lowe, Rohan (2017).
1225:Brogna S, Ashburner M (April 1997).
605:
248:
226:
205:
2449:
1400:. Dartmouth College. Archived from
672:sequence that enables a gene to be
561:sequence which provides a site for
449:of the gene into an mRNA. The mRNA
13:
3129:Evolutionary developmental biology
1737:Microbiology and Immunology Online
1597:Lee JM, Sonnhammer EL (May 2003).
1027:Evolutionary developmental biology
926:and colleagues, the trp operon in
747:Control of an operon is a type of
498:. An operon is made up of several
399:Protein
385:mRNA
14:
3354:
1940:
1730:
907:
839:
3086:Evolution of sexual reproduction
1950:H37Rv Operon Correlation Browser
1520:Introduction to genetic analysis
1362:10.1128/JB.186.13.4369-4375.2004
1344:Liu J, Mushegian A (July 2004).
1011:
997:
676:. The promoter is recognized by
457:into the final protein products.
408:
197:
2680:Post-translational modification
2337:Archaeal transcription factor B
1884:
1835:
1776:
1751:
1724:
1639:
1590:
1538:
1509:
799:, involved in the synthesis of
542:An operon contains one or more
2857:Genotype–phenotype distinction
1386:
1311:
1277:Blumenthal T (November 2004).
1218:
1175:
1096:
1069:
445:regions (yellow) regulate the
54:under the control of a single
1:
3114:Regulation of gene expression
1961:(a bit awkward to use though)
1758:Cummings MS, Klug WS (2006).
1062:
742:
3284:Endless Forms Most Beautiful
3064:Evolution of genetic systems
2872:Gene–environment correlation
2867:Gene–environment interaction
1892:saprophytism to virulence".
1196:10.1016/0092-8674(93)90139-H
1079:Life: The Science of Biology
987:under different conditions.
829:positive repressible operons
786:negative repressible operons
301:3'UTR
273:5'UTR
222:Regulatory sequence
215:Regulatory sequence
7:
3263:Christiane NĂĽsslein-Volhard
1394:"Bacteriophage Use Operons"
1005:Evolutionary biology portal
990:
466:Operons occur primarily in
185:
10:
3359:
3139:Hedgehog signaling pathway
3016:Developmental architecture
2013:Transcriptional regulation
1948:Mycobacterium tuberculosis
1151:10.1016/j.cell.2019.01.034
915:
847:
817:positive inducible operons
764:negative inducible operons
538:As a unit of transcription
135:
105:), but their discovery in
18:
3316:
3295:
3224:
3152:
3106:
3099:
3063:
3015:
2979:
2966:Transgressive segregation
2912:
2849:
2783:
2692:
2657:
2631:
2622:
2580:
2554:
2528:
2519:
2457:
2380:
2345:
2319:
2244:
2210:Transcription coregulator
2202:
2179:
2156:
2146:Histone acetyltransferase
2116:Histone methyltransferase
2094:Histone-modifying enzymes
2092:
2085:
2020:
2011:
1550:"The birth of the operon"
896:. It can be activated by
142:French Academy of Science
46:is a functioning unit of
2741:sequestration (P-bodies)
1398:Prokaryotic Gene Control
502:arranged under a common
470:but also rarely in some
50:containing a cluster of
19:Not to be confused with
3144:Notch signaling pathway
3119:Gene regulatory network
3002:Dual inheritance theory
2719:Gene regulatory network
2311:Internal control region
1575:10.1126/science.1207943
1481:WikiJournal of Medicine
1350:Journal of Bacteriology
1279:"Operons in eukaryotes"
1243:10.1093/emboj/16.8.2023
1037:Gene regulatory network
522:Operons are related to
487:Drosophila melanogaster
3192:cis-regulatory element
3100:Control of development
2980:Non-genetic influences
2946:evolutionary landscape
2724:cis-regulatory element
1848:Nucleic Acids Research
1810:10.1073/pnas.110147297
1319:"Definition of Operon"
1106:Molecular Cell Biology
980:Listeria monocytogenes
922:Discovered in 1953 by
658:
431:protein coding regions
101:that are derived from
35:
3303:Nature versus nurture
3207:Cell surface receptor
3124:Evo-devo gene toolkit
3023:Developmental biology
2961:Polygenic inheritance
2887:Quantitative genetics
2404:Intrinsic termination
2169:DNA methyltransferase
1860:10.1093/nar/29.5.1216
1494:10.15347/wjm/2017.002
936:tryptophan synthetase
615:
86:to define an operon.
33:
3212:Transcription factor
2927:Genetic assimilation
2914:Genetic architecture
2746:alternative splicing
2736:Post-transcriptional
2562:Transcription factor
2181:Chromatin remodeling
1760:Concepts of genetics
1665:10.1155/2023/5512414
1296:10.1093/bfgp/3.3.199
1145:(6): 1356–1366.e10.
1047:Protein biosynthesis
451:untranslated regions
208:Polycistronic operon
3308:Morphogenetic field
3225:Influential figures
2670:Transfer RNA (tRNA)
2138:Histone deacetylase
2128:Histone demethylase
2112:Histone methylation
1914:10.1038/nature08080
1906:2009Natur.459..950T
1801:2000PNAS...97.6652S
1566:2011Sci...332..767J
484:and the fruit fly,
427:Regulatory sequence
414:The structure of a
148:acting at a single
2997:Genomic imprinting
2784:Influential people
2763:Post-translational
2582:Post-transcription
1404:on 28 January 2013
1323:Medical Dictionary
1042:L-arabinose operon
719:repressor proteins
659:
620:: RNA Polymerase,
421:of protein-coding
176:André Michel Lwoff
66:strand and either
36:
3325:
3324:
3258:Eric F. Wieschaus
3220:
3219:
3038:Pattern formation
2942:Fitness landscape
2804:
2803:
2688:
2687:
2618:
2617:
2494:Special transfers
2417:
2416:
2372:RNA polymerase II
2240:
2239:
2198:
2197:
1769:978-0-13-191833-7
1717:978-0-19-854267-4
1615:10.1101/gr.737703
1531:978-0-7167-4939-4
1325:. MedicineNet.com
1116:978-0-7167-3136-8
1109:. W. H. Freeman.
1089:978-1-4292-1962-4
731:Alternatively, a
606:General structure
464:
463:
126:T7 RNA polymerase
62:together into an
3350:
3268:William McGinnis
3237:Richard Lewontin
3232:C. H. Waddington
3104:
3103:
3081:Neutral networks
2831:
2824:
2817:
2808:
2807:
2629:
2628:
2526:
2525:
2444:
2437:
2430:
2421:
2420:
2294:Response element
2277:Response element
2090:
2089:
2018:
2017:
1989:
1982:
1975:
1966:
1965:
1934:
1933:
1888:
1882:
1881:
1871:
1839:
1833:
1832:
1822:
1812:
1780:
1774:
1773:
1755:
1749:
1748:
1746:
1744:
1728:
1722:
1721:
1700:Lewin B (1990).
1697:
1688:
1687:
1677:
1667:
1643:
1637:
1636:
1626:
1594:
1588:
1587:
1577:
1542:
1536:
1535:
1523:
1513:
1507:
1506:
1496:
1472:
1466:
1465:
1463:
1462:
1456:
1450:. Archived from
1433:
1423:
1414:
1413:
1411:
1409:
1390:
1384:
1383:
1373:
1341:
1335:
1334:
1332:
1330:
1315:
1309:
1308:
1298:
1274:
1265:
1264:
1254:
1231:The EMBO Journal
1222:
1216:
1215:
1179:
1173:
1172:
1162:
1130:
1121:
1120:
1100:
1094:
1093:
1073:
1021:
1016:
1015:
1007:
1002:
1001:
985:L. monocytogenes
870:structural genes
865:Escherichia coli
707:Structural genes
596:Asgard (archaea)
594:; the closer an
588:Asgard (archaea)
579:structural genes
544:structural genes
500:structural genes
453:(blue) regulate
412:
411:
401:
394:
387:
380:
373:
366:
359:
352:
345:
338:
331:
324:
317:
310:
303:
296:
289:
282:
275:
268:
261:
254:
246:
238:
231:
224:
217:
210:
201:
190:
93:(which includes
58:. The genes are
34:A typical operon
3358:
3357:
3353:
3352:
3351:
3349:
3348:
3347:
3338:Gene expression
3328:
3327:
3326:
3321:
3312:
3291:
3278:Sean B. Carroll
3216:
3148:
3095:
3059:
3011:
2992:Maternal effect
2975:
2908:
2845:
2835:
2805:
2800:
2779:
2714:Transcriptional
2684:
2653:
2614:
2605:Polyadenylation
2576:
2550:
2515:
2509:Protein→Protein
2460:
2453:
2451:Gene expression
2448:
2418:
2413:
2388:
2382:
2376:
2341:
2315:
2236:
2194:
2175:
2158:DNA methylation
2152:
2096:
2081:
2007:
1993:
1959:Operon database
1943:
1938:
1937:
1900:(7249): 950–6.
1889:
1885:
1840:
1836:
1781:
1777:
1770:
1756:
1752:
1742:
1740:
1729:
1725:
1718:
1698:
1691:
1644:
1640:
1603:Genome Research
1595:
1591:
1543:
1539:
1532:
1514:
1510:
1473:
1469:
1460:
1458:
1454:
1431:
1424:
1417:
1407:
1405:
1392:
1391:
1387:
1356:(13): 4369–75.
1342:
1338:
1328:
1326:
1317:
1316:
1312:
1275:
1268:
1223:
1219:
1180:
1176:
1131:
1124:
1117:
1101:
1097:
1090:
1074:
1070:
1065:
1017:
1010:
1003:
996:
993:
975:Pascale Cossart
952:
920:
914:
852:
846:
749:gene regulation
745:
715:regulatory gene
686:– a segment of
608:
584:gene clustering
575:
540:
460:
459:
458:
413:
405:
404:
403:
402:
397:
395:
390:
388:
383:
381:
376:
374:
369:
367:
362:
360:
355:
353:
348:
346:
341:
339:
334:
332:
327:
325:
320:
318:
313:
311:
306:
304:
299:
297:
292:
290:
285:
283:
278:
276:
271:
269:
264:
262:
257:
255:
249:
247:
241:
239:
234:
232:
227:
225:
220:
218:
213:
211:
206:
202:
188:
170:was awarded to
138:
120:. For example,
28:
17:
12:
11:
5:
3356:
3346:
3345:
3340:
3323:
3322:
3317:
3314:
3313:
3311:
3310:
3305:
3299:
3297:
3293:
3292:
3290:
3289:
3288:
3287:
3275:
3270:
3265:
3260:
3255:
3254:
3253:
3242:François Jacob
3239:
3234:
3228:
3226:
3222:
3221:
3218:
3217:
3215:
3214:
3209:
3204:
3199:
3194:
3189:
3184:
3179:
3178:
3177:
3167:
3162:
3156:
3154:
3150:
3149:
3147:
3146:
3141:
3136:
3131:
3126:
3121:
3116:
3110:
3108:
3101:
3097:
3096:
3094:
3093:
3088:
3083:
3078:
3073:
3067:
3065:
3061:
3060:
3058:
3057:
3052:
3047:
3042:
3041:
3040:
3035:
3025:
3019:
3017:
3013:
3012:
3010:
3009:
3004:
2999:
2994:
2989:
2983:
2981:
2977:
2976:
2974:
2973:
2971:Sequence space
2968:
2963:
2958:
2953:
2948:
2939:
2934:
2929:
2924:
2918:
2916:
2910:
2909:
2907:
2906:
2901:
2900:
2899:
2889:
2884:
2879:
2874:
2869:
2864:
2859:
2853:
2851:
2847:
2846:
2834:
2833:
2826:
2819:
2811:
2802:
2801:
2799:
2798:
2793:
2791:François Jacob
2787:
2785:
2781:
2780:
2778:
2777:
2776:
2775:
2770:
2760:
2755:
2754:
2753:
2748:
2743:
2733:
2728:
2727:
2726:
2721:
2711:
2710:
2709:
2698:
2696:
2690:
2689:
2686:
2685:
2683:
2682:
2677:
2672:
2667:
2661:
2659:
2655:
2654:
2652:
2651:
2646:
2641:
2635:
2633:
2626:
2620:
2619:
2616:
2615:
2613:
2612:
2607:
2602:
2597:
2592:
2586:
2584:
2578:
2577:
2575:
2574:
2569:
2567:RNA polymerase
2564:
2558:
2556:
2552:
2551:
2549:
2548:
2543:
2538:
2532:
2530:
2523:
2517:
2516:
2514:
2513:
2512:
2511:
2506:
2501:
2491:
2490:
2489:
2471:
2465:
2463:
2455:
2454:
2447:
2446:
2439:
2432:
2424:
2415:
2414:
2412:
2411:
2406:
2401:
2395:
2393:
2378:
2377:
2375:
2374:
2368:RNA polymerase
2362:
2356:RNA polymerase
2349:
2347:
2343:
2342:
2340:
2339:
2334:
2329:
2323:
2321:
2317:
2316:
2314:
2313:
2308:
2303:
2298:
2297:
2296:
2291:
2281:
2280:
2279:
2274:
2269:
2264:
2259:
2248:
2246:
2242:
2241:
2238:
2237:
2235:
2234:
2229:
2228:
2227:
2222:
2217:
2206:
2204:
2200:
2199:
2196:
2195:
2193:
2192:
2186:
2184:
2177:
2176:
2174:
2173:
2172:
2171:
2163:
2161:
2154:
2153:
2151:
2150:
2149:
2148:
2143:
2130:
2125:
2124:
2123:
2108:
2106:
2087:
2083:
2082:
2080:
2079:
2078:
2077:
2072:
2062:
2061:
2060:
2055:
2050:
2045:
2040:
2035:
2024:
2022:
2015:
2009:
2008:
1992:
1991:
1984:
1977:
1969:
1963:
1962:
1952:
1942:
1941:External links
1939:
1936:
1935:
1883:
1854:(5): 1216–21.
1834:
1795:(12): 6652–7.
1775:
1768:
1750:
1723:
1716:
1689:
1638:
1589:
1537:
1530:
1508:
1467:
1415:
1385:
1336:
1310:
1289:(3): 199–211.
1266:
1237:(8): 2023–31.
1217:
1174:
1122:
1115:
1095:
1088:
1067:
1066:
1064:
1061:
1060:
1059:
1057:Umu Chromotest
1054:
1049:
1044:
1039:
1034:
1029:
1023:
1022:
1019:Biology portal
1008:
992:
989:
951:
948:
916:Main article:
913:
906:
858:operon of the
848:Main article:
845:
838:
837:
836:
825:
805:
804:
790:regulator gene
782:
744:
741:
711:
710:
704:
681:
678:RNA polymerase
607:
604:
574:
571:
563:RNA polymerase
539:
536:
462:
461:
407:
406:
204:
203:
196:
195:
194:
193:
187:
184:
172:François Jacob
137:
134:
118:bacteriophages
84:co-transcribed
15:
9:
6:
4:
3:
2:
3355:
3344:
3341:
3339:
3336:
3335:
3333:
3320:
3315:
3309:
3306:
3304:
3301:
3300:
3298:
3294:
3286:
3285:
3281:
3280:
3279:
3276:
3274:
3271:
3269:
3266:
3264:
3261:
3259:
3256:
3252:
3249:
3248:
3247:
3246:Jacques Monod
3243:
3240:
3238:
3235:
3233:
3230:
3229:
3227:
3223:
3213:
3210:
3208:
3205:
3203:
3200:
3198:
3195:
3193:
3190:
3188:
3185:
3183:
3180:
3176:
3173:
3172:
3171:
3168:
3166:
3163:
3161:
3160:Homeotic gene
3158:
3157:
3155:
3151:
3145:
3142:
3140:
3137:
3135:
3132:
3130:
3127:
3125:
3122:
3120:
3117:
3115:
3112:
3111:
3109:
3105:
3102:
3098:
3092:
3089:
3087:
3084:
3082:
3079:
3077:
3074:
3072:
3069:
3068:
3066:
3062:
3056:
3053:
3051:
3048:
3046:
3043:
3039:
3036:
3034:
3031:
3030:
3029:
3028:Morphogenesis
3026:
3024:
3021:
3020:
3018:
3014:
3008:
3005:
3003:
3000:
2998:
2995:
2993:
2990:
2988:
2985:
2984:
2982:
2978:
2972:
2969:
2967:
2964:
2962:
2959:
2957:
2954:
2952:
2949:
2947:
2943:
2940:
2938:
2935:
2933:
2930:
2928:
2925:
2923:
2920:
2919:
2917:
2915:
2911:
2905:
2902:
2898:
2895:
2894:
2893:
2890:
2888:
2885:
2883:
2880:
2878:
2875:
2873:
2870:
2868:
2865:
2863:
2862:Reaction norm
2860:
2858:
2855:
2854:
2852:
2848:
2844:
2840:
2832:
2827:
2825:
2820:
2818:
2813:
2812:
2809:
2797:
2796:Jacques Monod
2794:
2792:
2789:
2788:
2786:
2782:
2774:
2771:
2769:
2766:
2765:
2764:
2761:
2759:
2758:Translational
2756:
2752:
2749:
2747:
2744:
2742:
2739:
2738:
2737:
2734:
2732:
2729:
2725:
2722:
2720:
2717:
2716:
2715:
2712:
2708:
2705:
2704:
2703:
2700:
2699:
2697:
2695:
2691:
2681:
2678:
2676:
2673:
2671:
2668:
2666:
2663:
2662:
2660:
2656:
2650:
2647:
2645:
2642:
2640:
2637:
2636:
2634:
2630:
2627:
2625:
2621:
2611:
2608:
2606:
2603:
2601:
2598:
2596:
2593:
2591:
2588:
2587:
2585:
2583:
2579:
2573:
2570:
2568:
2565:
2563:
2560:
2559:
2557:
2553:
2547:
2544:
2542:
2539:
2537:
2534:
2533:
2531:
2527:
2524:
2522:
2521:Transcription
2518:
2510:
2507:
2505:
2502:
2500:
2497:
2496:
2495:
2492:
2488:
2484:
2480:
2477:
2476:
2475:
2474:Central dogma
2472:
2470:
2467:
2466:
2464:
2462:
2456:
2452:
2445:
2440:
2438:
2433:
2431:
2426:
2425:
2422:
2410:
2407:
2405:
2402:
2400:
2397:
2396:
2394:
2391:
2386:
2379:
2373:
2369:
2366:
2363:
2361:
2357:
2354:
2351:
2350:
2348:
2344:
2338:
2335:
2333:
2330:
2328:
2325:
2324:
2322:
2318:
2312:
2309:
2307:
2304:
2302:
2299:
2295:
2292:
2290:
2287:
2286:
2285:
2282:
2278:
2275:
2273:
2270:
2268:
2265:
2263:
2260:
2258:
2255:
2254:
2253:
2250:
2249:
2247:
2243:
2233:
2230:
2226:
2223:
2221:
2218:
2216:
2213:
2212:
2211:
2208:
2207:
2205:
2201:
2191:
2188:
2187:
2185:
2182:
2178:
2170:
2167:
2166:
2165:
2164:
2162:
2159:
2155:
2147:
2144:
2142:
2139:
2136:
2135:
2134:
2131:
2129:
2126:
2122:
2119:
2118:
2117:
2113:
2110:
2109:
2107:
2104:
2100:
2095:
2091:
2088:
2084:
2076:
2075:trp repressor
2073:
2071:
2070:lac repressor
2068:
2067:
2066:
2063:
2059:
2056:
2054:
2051:
2049:
2046:
2044:
2041:
2039:
2036:
2034:
2031:
2030:
2029:
2026:
2025:
2023:
2019:
2016:
2014:
2010:
2005:
2001:
1997:
1996:Transcription
1990:
1985:
1983:
1978:
1976:
1971:
1970:
1967:
1960:
1956:
1953:
1951:
1949:
1945:
1944:
1931:
1927:
1923:
1919:
1915:
1911:
1907:
1903:
1899:
1895:
1887:
1879:
1875:
1870:
1865:
1861:
1857:
1853:
1849:
1845:
1838:
1830:
1826:
1821:
1816:
1811:
1806:
1802:
1798:
1794:
1790:
1786:
1779:
1771:
1765:
1761:
1754:
1738:
1734:
1727:
1719:
1713:
1709:
1705:
1704:
1696:
1694:
1685:
1681:
1676:
1671:
1666:
1661:
1657:
1653:
1649:
1642:
1634:
1630:
1625:
1620:
1616:
1612:
1609:(5): 875–82.
1608:
1604:
1600:
1593:
1585:
1581:
1576:
1571:
1567:
1563:
1560:(6031): 767.
1559:
1555:
1551:
1547:
1541:
1533:
1527:
1522:
1521:
1512:
1504:
1500:
1495:
1490:
1486:
1482:
1478:
1471:
1457:on 2016-03-04
1453:
1449:
1445:
1442:(6): 1727–9.
1441:
1437:
1429:
1422:
1420:
1403:
1399:
1395:
1389:
1381:
1377:
1372:
1367:
1363:
1359:
1355:
1351:
1347:
1340:
1324:
1320:
1314:
1306:
1302:
1297:
1292:
1288:
1284:
1280:
1273:
1271:
1262:
1258:
1253:
1248:
1244:
1240:
1236:
1232:
1228:
1221:
1213:
1209:
1205:
1201:
1197:
1193:
1190:(3): 521–32.
1189:
1185:
1178:
1170:
1166:
1161:
1156:
1152:
1148:
1144:
1140:
1136:
1129:
1127:
1118:
1112:
1108:
1107:
1099:
1091:
1085:
1081:
1080:
1072:
1068:
1058:
1055:
1053:
1050:
1048:
1045:
1043:
1040:
1038:
1035:
1033:
1030:
1028:
1025:
1024:
1020:
1014:
1009:
1006:
1000:
995:
988:
986:
982:
981:
976:
972:
968:
965:
961:
959:
958:
947:
945:
944:corepressible
941:
937:
933:
929:
925:
924:Jacques Monod
919:
911:
905:
903:
902:derepressible
899:
895:
891:
887:
883:
879:
875:
871:
867:
866:
861:
857:
851:
843:
834:
830:
826:
822:
818:
814:
813:
812:
810:
802:
798:
796:
791:
787:
783:
780:
776:
774:
769:
765:
761:
760:
759:
757:
752:
750:
740:
738:
734:
729:
727:
722:
720:
716:
708:
705:
701:
697:
693:
689:
685:
682:
679:
675:
671:
667:
664:
663:
662:
656:
652:
649:
647:
643:
639:
635:
631:
627:
624:: Repressor,
623:
619:
614:
610:
603:
601:
597:
593:
589:
585:
580:
570:
568:
564:
560:
556:
552:
549:
548:polycistronic
545:
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448:
447:transcription
444:
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393:
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379:
372:
365:
364:RBS
358:
357:RBS
351:
344:
343:Transcription
337:
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180:Jacques Monod
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111:polycistronic
108:
104:
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96:
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87:
85:
81:
77:
76:monocistronic
73:
69:
65:
61:
57:
53:
49:
45:
41:
32:
26:
22:
3282:
3175:eyeless gene
3071:Evolvability
3045:Segmentation
2922:Canalisation
2892:Heterochrony
2882:Heritability
2876:
2850:Key concepts
2773:irreversible
2658:Key elements
2555:Key elements
2469:Genetic code
2459:Introduction
2027:
1947:
1897:
1893:
1886:
1851:
1847:
1837:
1792:
1788:
1778:
1759:
1753:
1741:. Retrieved
1736:
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1548:(May 2011).
1540:
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1484:
1480:
1470:
1459:. Retrieved
1452:the original
1439:
1435:
1406:. Retrieved
1402:the original
1397:
1388:
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1349:
1339:
1327:. Retrieved
1322:
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1234:
1230:
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1187:
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1177:
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1105:
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1071:
1032:Genetic code
984:
978:
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863:
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772:
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753:
746:
730:
723:
712:
699:
683:
660:
654:
650:
645:
641:
637:
633:
632:: Operator,
629:
628:: Promoter,
625:
621:
617:
616:
609:
576:
566:
541:
521:
513:corepressors
486:
480:
474:, including
465:
418:
207:
166:. The 1965
161:
155:
139:
115:
88:
83:
43:
37:
3273:Mike Levine
3182:Distal-less
3007:Polyphenism
2987:Epigenetics
2839:development
2624:Translation
2461:to genetics
2381:Termination
2257:Pribnow box
2225:Corepressor
2220:Coactivator
2021:prokaryotic
1743:30 December
1408:30 December
1329:30 December
932:allolactose
898:allolactose
824:activators.
779:allolactose
733:corepressor
690:to which a
674:transcribed
636:: Lactose,
468:prokaryotes
455:translation
416:prokaryotic
392:Translation
91:prokaryotes
60:transcribed
3332:Categories
3251:Lac operon
3076:Robustness
3055:Modularity
3050:Metamerism
2956:Plasticity
2951:Pleiotropy
2904:Heterotopy
2768:reversible
2731:lac operon
2707:imprinting
2702:Epigenetic
2694:Regulation
2649:Eukaryotic
2595:5' capping
2546:Eukaryotic
2409:Rho factor
2399:Terminator
2390:eukaryotic
2365:eukaryotic
2346:Elongation
2332:Eukaryotic
2320:Initiation
2103:nucleosome
2086:eukaryotic
2058:gal operon
2053:ara operon
2048:Gua Operon
2043:gab operon
2038:trp operon
2033:lac operon
2004:Eukaryotic
1461:2015-08-27
1063:References
940:attenuator
918:trp operon
878:terminator
862:bacterium
850:lac operon
801:tryptophan
743:Regulation
737:trp operon
696:lac operon
670:nucleotide
600:eukaryotes
598:is to the
517:activators
509:repressors
481:C. elegans
472:eukaryotes
336:Terminator
329:Stop
322:Stop
315:Start
308:Start
107:eukaryotes
95:organelles
74:to create
68:translated
3202:Morphogen
3187:Engrailed
3170:Pax genes
3091:Tinkering
2937:Epistasis
2932:Dominance
2843:phenotype
2639:Bacterial
2536:Bacterial
2385:bacterial
2353:bacterial
2327:Bacterial
2301:Insulator
2245:Promotion
2215:Activator
2065:Repressor
2000:Bacterial
1731:Mayer G.
1503:2002-4436
880:, and an
833:inhibitor
809:activator
756:repressor
692:repressor
528:stimulons
496:chordates
476:nematodes
146:repressor
122:T7 phages
80:expressed
3165:Hox gene
3153:Elements
3134:Homeobox
2751:microRNA
2665:Ribosome
2644:Archaeal
2600:Splicing
2572:Promoter
2541:Archaeal
2485: →
2481: →
2306:Silencer
2284:Enhancer
2272:CAAT box
2262:TATA box
2252:Promoter
1922:19448609
1878:11222772
1829:10823905
1703:Genes IV
1684:38314098
1675:10833476
1658:: 1–16.
1633:12695325
1584:21566161
1448:14406329
1380:15205439
1305:15642184
1212:26918553
1169:30799038
1052:TATA box
991:See also
882:operator
874:promoter
684:Operator
666:Promoter
648:: lacA.
644:: lacY,
640:: lacZ,
577:All the
567:operator
559:promoter
532:modulons
524:regulons
504:promoter
478:such as
443:enhancer
439:operator
435:Promoter
294:UTR
287:ORF
280:ORF
266:Promoter
259:Operator
252:silencer
244:silencer
236:Enhancer
229:Enhancer
186:Overview
150:operator
103:bacteria
99:plastids
72:splicing
56:promoter
40:genetics
3343:Operons
3296:Debates
3107:Systems
3033:Eyespot
2897:Neoteny
2504:RNA→DNA
2499:RNA→RNA
2487:Protein
2232:Inducer
2099:histone
1930:4341657
1902:Bibcode
1797:Bibcode
1652:Archaea
1562:Bibcode
1554:Science
1546:Jacob F
1261:9155028
1252:1169805
1204:8098272
1160:7295392
964:bases.
957:E. coli
946:model.
928:E. coli
894:lactose
890:glucose
821:inducer
768:inducer
726:inducer
592:Archaea
555:protein
433:(red).
163:E. coli
136:History
21:Opteron
3197:Ligand
2877:Operon
2028:Operon
1928:
1920:
1894:Nature
1876:
1866:
1827:
1817:
1766:
1714:
1708:243–58
1682:
1672:
1631:
1624:430880
1621:
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1528:
1501:
1446:
1378:
1371:421614
1368:
1303:
1259:
1249:
1210:
1202:
1167:
1157:
1113:
1086:
912:operon
884:. The
844:operon
797:operon
775:operon
703:genes.
655:Bottom
515:, and
419:operon
158:operon
44:operon
25:Oberon
2632:Types
2529:Types
2289:E-box
2141:HDAC1
1926:S2CID
1869:29727
1820:18690
1487:(1).
1455:(PDF)
1432:(PDF)
1208:S2CID
860:model
423:genes
130:lysis
97:like
52:genes
42:, an
2837:The
2360:rpoB
2203:both
2190:CHD7
2121:EZH2
1918:PMID
1874:PMID
1825:PMID
1764:ISBN
1745:2012
1712:ISBN
1680:PMID
1656:2023
1629:PMID
1580:PMID
1526:ISBN
1499:ISSN
1444:PMID
1410:2012
1376:PMID
1331:2012
1301:PMID
1257:PMID
1200:PMID
1184:Cell
1165:PMID
1139:Cell
1111:ISBN
1084:ISBN
908:The
892:and
876:, a
872:, a
854:The
840:The
668:– a
551:mRNA
530:and
492:rRNA
441:and
178:and
64:mRNA
2841:of
2483:RNA
2479:DNA
2267:BRE
1955:OBD
1910:doi
1898:459
1864:PMC
1856:doi
1815:PMC
1805:doi
1670:PMC
1660:doi
1619:PMC
1611:doi
1570:doi
1558:332
1489:doi
1440:250
1366:PMC
1358:doi
1354:186
1291:doi
1247:PMC
1239:doi
1192:doi
1155:PMC
1147:doi
1143:176
910:trp
886:lac
856:lac
842:lac
827:In
815:In
795:trp
784:In
773:lac
762:In
724:An
700:lac
688:DNA
651:Top
160:in
156:lac
48:DNA
38:In
23:or
3334::
3244:+
2370::
2358::
2105:):
2002:,
1957:-
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1850:.
1846:.
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1803:.
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1787:.
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1188:73
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1137:.
1125:^
739:.
569:.
526:,
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2944:/
2830:e
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1988:e
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618:1
250:/
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27:.
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