210:
949:. 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
421:
1024:
1010:
624:
668:: 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.
545:; 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".
981:
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
592:
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
163:
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
974:
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
803:, 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
1902:
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
42:
597:. 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
713:
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
164:
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
517:
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
834:
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
664:: 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.
691:, 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.
530:, 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.
1193:
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".
762:
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.
155:
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
89:
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
2504:
945:), 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
781:
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
1144:
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).
568:). 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
1404:
1438:
1238:"The Adh-related gene of Drosophila melanogaster is expressed as a functional dicistronic messenger RNA: multigenic transcription in higher organisms"
978:
An alternative method to predict operons is based on finding gene clusters where gene order and orientation is conserved in two or more genomes.
209:
777:, a regulatory repressor protein is normally bound to the operator, which prevents the transcription of the genes on the operon. If an
120:
was shown in the early 1990s, and are considered to be rare. In general, expression of prokaryotic operons leads to the generation of
2452:
1997:
746:
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
3329:
178:
2620:
2143:
1778:
1726:
1540:
1125:
1098:
915:(from above: negative inducible) model. So it is a negative inducible operon induced by presence of lactose or allolactose.
3294:
2375:
994:. The 517 polycistronic operons are listed in a 2009 study describing the global changes in transcription that occur in
1957:
3139:
2867:
2484:
1037:
911:. Lactose binds to the repressor protein and prevents it from repressing gene transcription. This is an example of the
879:
was the first operon to be discovered and provides a typical example of operon function. It consists of three adjacent
846:
is bound by the activator, it is prevented from binding the DNA. This stops activation and transcription of the system.
17:
739:(small molecule) can displace a repressor (protein) from the operator site (DNA), resulting in an uninhibited operon.
2839:
2592:
3096:
2882:
2877:
2746:
2690:
2347:
1412:
2685:
2400:
1462:
1114:
Lodish H, Zipursky L, Matsudaira P, Baltimore D, Darnel J (2000). "Chapter 9: Molecular
Definition of a Gene".
601:, ribosomal protein coding genes occur in clusters that are less conserved in their organization than in other
3273:
2342:
3124:
2773:
2704:
2445:
2395:
2363:
2337:
2509:
3091:
1990:
1015:
941:
was the first repressible operon to be discovered. While the lac operon can be activated by a chemical (
3283:
3149:
2981:
2724:
2023:
2976:
2220:
2156:
2126:
2104:
152:
3348:
2768:
2556:
2469:
2438:
2014:
2006:
1357:"Displacements of prohead protease genes in the late operons of double-stranded-DNA bacteriophages"
684:
457:
353:
732:. The regulatory gene does not need to be in, adjacent to, or even near the operon to control it.
3217:
3154:
3129:
3012:
2932:
2729:
2546:
2531:
2321:
2010:
1047:
70:
822:
protein stimulates transcription by binding to DNA (usually at a site other than the operator).
3202:
3101:
3086:
3055:
2956:
2734:
2659:
2551:
2230:
1983:
1329:
990:
954:
594:
523:
490:
402:
1088:
3313:
3207:
3134:
3033:
2971:
2966:
2897:
2832:
2649:
2634:
2514:
2414:
2409:
2277:
2179:
950:
946:
888:
709:
as a segment between the promoter and the genes of the operon. The main operator (O1) in the
465:
441:
388:
381:
374:
367:
346:
78:
1529:
186:
3222:
3065:
3060:
2942:
2937:
2924:
2756:
2654:
2572:
2311:
2225:
2191:
2085:
1912:
1807:
1572:
1057:
842:, the activator proteins are normally bound to the pertinent DNA segment. However, when an
819:
527:
576:
to bind and initiate transcription. Close to the promoter lies a section of DNA called an
8:
3318:
2582:
2294:
2262:
2148:
2122:
799:, transcription of the operon normally takes place. Repressor proteins are produced by a
461:
453:
445:
437:
304:
276:
269:
262:
254:
246:
239:
232:
225:
66:
1975:
1916:
1811:
1685:
1658:
1576:
3007:
2717:
2600:
2063:
1936:
1262:
1237:
1218:
1170:
1145:
1052:
988:'s laboratory was the first to experimentally identify all operons of a microorganism,
892:
884:
843:
729:
676:
569:
514:
449:
297:
290:
160:
1634:
1609:
1381:
1356:
814:(which itself acts as the corepressor), is a negatively controlled repressible operon.
3268:
3048:
3043:
2952:
2382:
1928:
1884:
1879:
1854:
1835:
1830:
1795:
1774:
1722:
1718:
1712:
1690:
1639:
1590:
1536:
1509:
1454:
1386:
1372:
1311:
1267:
1210:
1206:
1175:
1121:
1115:
1094:
505:
genes often exist in operons that have been found in a range of eukaryotes including
136:
3252:
2801:
1556:
1222:
182:
3353:
3278:
3247:
3242:
3197:
2825:
2304:
2287:
1940:
1920:
1874:
1866:
1825:
1815:
1743:
1680:
1670:
1629:
1621:
1580:
1499:
1439:"[Operon: a group of genes with the expression coordinated by an operator]"
1376:
1368:
1301:
1257:
1249:
1202:
1165:
1157:
967:
875:
606:
598:
173:
135:
have two operons. The first operon codes for various products, including a special
1441:[Operon: a group of genes with the expression coordinated by an operator]
3288:
3192:
3002:
2778:
2615:
2461:
2168:
1969:
985:
880:
759:
725:
717:
589:
554:
510:
139:
which can bind to and transcribe the second operon. The second operon includes a
90:
1527:
Miller JH, Suzuki DT, Griffiths AJ, Lewontin RC, Wessler SR, Gelbart WM (2005).
971:. As a result, predictions can be made based on an organism's genomic sequence.
151:
The term "operon" was first proposed in a short paper in the
Proceedings of the
2817:
2605:
2577:
2378:
2366:
2316:
1800:
Proceedings of the
National Academy of Sciences of the United States of America
1161:
1067:
1029:
870:
830:, activator proteins are normally unable to bind to the pertinent DNA. When an
800:
688:
613:, the more dispersed is the arrangement of the ribosomal protein coding genes.
573:
1253:
3342:
3256:
3170:
3038:
2872:
2849:
2806:
2493:
2080:
1773:(8th ed.). Upper Saddle River, NJ: Pearson Education. pp. 394–402.
1513:
934:
558:
502:
190:
128:
74:
1870:
1585:
1560:
1504:
1487:
3081:
2902:
2892:
2680:
2610:
2479:
1932:
1888:
1839:
1820:
1694:
1675:
1643:
1594:
1458:
1390:
1315:
1306:
1289:
1179:
1042:
965:
The number and organization of operons has been studied most critically in
912:
788:
is a negatively controlled inducible operon, where the inducer molecule is
82:
1271:
1214:
724:
Not always included within the operon, but important in its function is a
3017:
2997:
2783:
2712:
2267:
2235:
2138:
1793:
942:
908:
789:
743:
426:
325:
318:
1924:
1657:
Tirumalai MR, Sivaraman RV, Kutty LA, Song EL, Fox GE (September 2003).
3261:
2961:
2914:
2741:
2419:
2113:
2068:
2058:
2053:
2048:
2043:
953:
sequence which allows for graded regulation. This is an example of the
928:
860:
811:
804:
782:
747:
706:
680:
496:
478:
339:
332:
165:
101:
1794:
Salgado H, Moreno-Hagelsieb G, Smith TF, Collado-Vides J (June 2000).
1625:
3212:
3180:
2947:
2853:
2075:
899:
operon is regulated by several factors including the availability of
818:
Operons can also be positively controlled. With positive control, an
766:
702:
610:
519:
482:
156:
117:
105:
960:
3175:
3144:
2761:
2751:
2675:
2282:
2272:
1447:
Comptes Rendus
Hebdomadaires des Séances de l'Académie des Sciences
1062:
1023:
538:
506:
486:
311:
283:
132:
113:
50:
2430:
2907:
2497:
2242:
2109:
1901:
1113:
904:
900:
831:
778:
736:
602:
583:
565:
542:
534:
409:
193:
for their discoveries concerning the operon and virus synthesis.
121:
109:
86:
31:
420:
1796:"Operons in Escherichia coli: genomic analyses and predictions"
1009:
35:
2519:
2299:
2151:
1526:
623:
140:
124:
mRNAs, while eukaryotic operons lead to monocistronic mRNAs.
1610:"Genomic gene clustering analysis of pathways in eukaryotes"
3185:
2370:
2200:
2131:
1744:"Bacteriology – Chapter Nine Genetic Regulatory Mechanisms"
1656:
1086:
561:
433:
395:
62:
41:
1717:(4th ed.). Oxford: Oxford University Press. pp.
1659:"Ribosomal Protein Cluster Organization in Asgard Archaea"
1535:(8th ed.). San Francisco: W.H. Freeman. p. 740.
1143:
2489:
2005:
698:
360:
58:
1437:
Jacob F, Perrin D, Sanchez C, Monod J (February 1960).
1436:
1283:
1281:
1192:
1852:
27:
Group of open reading frames under the same regulation
1965:
1087:
Sadava DE, Hillis DM, Heller HC, Berenbaum M (2009).
564:(a single mRNA molecule that codes for more than one
1278:
1005:
200:
100:
Originally, operons were thought to exist solely in
1449:(Facsimile version reprinted in 2005) (in French).
1528:
1853:Ermolaeva MD, White O, Salzberg SL (March 2001).
1750:. University of South Carolina School of Medicine
1294:Briefings in Functional Genomics & Proteomics
961:Predicting the number and organization of operons
440:controls when expression occurs for the multiple
3340:
2847:
720:– the genes that are co-regulated by the operon.
672:An operon is made up of 3 basic DNA components:
1235:
1479:
1146:"Eukaryotic Acquisition of a Bacterial Operon"
728:, a constantly expressed gene which codes for
584:Operons versus clustering of prokaryotic genes
93:together or not at all. Several genes must be
2833:
2446:
1991:
1607:
548:
1855:"Prediction of operons in microbial genomes"
1768:
1432:
1430:
1354:
982:all of these data, a difficult task indeed.
143:gene meant to cause the host cell to burst.
1488:"Eukaryotic and prokaryotic gene structure"
765:Negative control involves the binding of a
2840:
2826:
2601:Precursor mRNA (pre-mRNA / hnRNA)
2453:
2439:
1998:
1984:
1485:
1287:
769:to the operator to prevent transcription.
127:Operons are also found in viruses such as
1878:
1829:
1819:
1706:
1704:
1684:
1674:
1633:
1584:
1503:
1427:
1380:
1305:
1261:
1169:
1139:
1137:
705:binds. It is classically defined in the
557:which are generally transcribed into one
1093:(9th ed.). Macmillan. p. 349.
622:
40:
1348:
14:
3341:
3330:Index of evolutionary biology articles
1701:
1134:
389:Protein coding region
386:
382:Protein coding region
379:
372:
365:
361:DNA
344:
337:
330:
323:
316:
179:Nobel Prize in Physiology and Medicine
81:together in the cytoplasm, or undergo
2821:
2621:Histone acetylation and deacetylation
2434:
2144:Histone acetylation and deacetylation
1979:
1710:
1555:
407:
400:
393:
358:
351:
309:
302:
295:
288:
281:
274:
267:
251:
244:
230:
223:
2686:Ribosome-nascent chain complex (RNC)
1486:Shafee, Thomas; Lowe, Rohan (2017).
1236:Brogna S, Ashburner M (April 1997).
616:
259:
237:
216:
2460:
1411:. Dartmouth College. Archived from
683:sequence that enables a gene to be
572:sequence which provides a site for
460:of the gene into an mRNA. The mRNA
24:
3140:Evolutionary developmental biology
1748:Microbiology and Immunology Online
1608:Lee JM, Sonnhammer EL (May 2003).
1038:Evolutionary developmental biology
937:and colleagues, the trp operon in
758:Control of an operon is a type of
509:. An operon is made up of several
410:Protein
396:mRNA
25:
3365:
1951:
1741:
918:
850:
3097:Evolution of sexual reproduction
1961:H37Rv Operon Correlation Browser
1531:Introduction to genetic analysis
1373:10.1128/JB.186.13.4369-4375.2004
1355:Liu J, Mushegian A (July 2004).
1022:
1008:
687:. The promoter is recognized by
468:into the final protein products.
419:
208:
2691:Post-translational modification
2348:Archaeal transcription factor B
1895:
1846:
1787:
1762:
1735:
1650:
1601:
1549:
1520:
810:, involved in the synthesis of
553:An operon contains one or more
2868:Genotype–phenotype distinction
1397:
1322:
1288:Blumenthal T (November 2004).
1229:
1186:
1107:
1080:
456:regions (yellow) regulate the
65:under the control of a single
13:
1:
3125:Regulation of gene expression
1972:(a bit awkward to use though)
1769:Cummings MS, Klug WS (2006).
1073:
753:
3295:Endless Forms Most Beautiful
3075:Evolution of genetic systems
2883:Gene–environment correlation
2878:Gene–environment interaction
1903:saprophytism to virulence".
1207:10.1016/0092-8674(93)90139-H
1090:Life: The Science of Biology
998:under different conditions.
840:positive repressible operons
797:negative repressible operons
312:3'UTR
284:5'UTR
233:Regulatory sequence
226:Regulatory sequence
7:
3274:Christiane NĂĽsslein-Volhard
1405:"Bacteriophage Use Operons"
1016:Evolutionary biology portal
1001:
477:Operons occur primarily in
196:
10:
3370:
3150:Hedgehog signaling pathway
3027:Developmental architecture
2024:Transcriptional regulation
1959:Mycobacterium tuberculosis
1162:10.1016/j.cell.2019.01.034
926:
858:
828:positive inducible operons
775:negative inducible operons
549:As a unit of transcription
146:
116:), but their discovery in
29:
3327:
3306:
3235:
3163:
3117:
3110:
3074:
3026:
2990:
2977:Transgressive segregation
2923:
2860:
2794:
2703:
2668:
2642:
2633:
2591:
2565:
2539:
2530:
2468:
2391:
2356:
2330:
2255:
2221:Transcription coregulator
2213:
2190:
2167:
2157:Histone acetyltransferase
2127:Histone methyltransferase
2105:Histone-modifying enzymes
2103:
2096:
2031:
2022:
1561:"The birth of the operon"
907:. It can be activated by
153:French Academy of Science
57:is a functioning unit of
2752:sequestration (P-bodies)
1409:Prokaryotic Gene Control
513:arranged under a common
481:but also rarely in some
61:containing a cluster of
30:Not to be confused with
3155:Notch signaling pathway
3130:Gene regulatory network
3013:Dual inheritance theory
2730:Gene regulatory network
2322:Internal control region
1586:10.1126/science.1207943
1492:WikiJournal of Medicine
1361:Journal of Bacteriology
1290:"Operons in eukaryotes"
1254:10.1093/emboj/16.8.2023
1048:Gene regulatory network
533:Operons are related to
498:Drosophila melanogaster
3203:cis-regulatory element
3111:Control of development
2991:Non-genetic influences
2957:evolutionary landscape
2735:cis-regulatory element
1859:Nucleic Acids Research
1821:10.1073/pnas.110147297
1330:"Definition of Operon"
1117:Molecular Cell Biology
991:Listeria monocytogenes
933:Discovered in 1953 by
669:
442:protein coding regions
112:that are derived from
46:
3314:Nature versus nurture
3218:Cell surface receptor
3135:Evo-devo gene toolkit
3034:Developmental biology
2972:Polygenic inheritance
2898:Quantitative genetics
2415:Intrinsic termination
2180:DNA methyltransferase
1871:10.1093/nar/29.5.1216
1505:10.15347/wjm/2017.002
947:tryptophan synthetase
626:
97:to define an operon.
44:
3223:Transcription factor
2938:Genetic assimilation
2925:Genetic architecture
2757:alternative splicing
2747:Post-transcriptional
2573:Transcription factor
2192:Chromatin remodeling
1771:Concepts of genetics
1676:10.1155/2023/5512414
1307:10.1093/bfgp/3.3.199
1156:(6): 1356–1366.e10.
1058:Protein biosynthesis
462:untranslated regions
219:Polycistronic operon
3319:Morphogenetic field
3236:Influential figures
2681:Transfer RNA (tRNA)
2149:Histone deacetylase
2139:Histone demethylase
2123:Histone methylation
1925:10.1038/nature08080
1917:2009Natur.459..950T
1812:2000PNAS...97.6652S
1577:2011Sci...332..767J
495:and the fruit fly,
438:Regulatory sequence
425:The structure of a
159:acting at a single
3008:Genomic imprinting
2795:Influential people
2774:Post-translational
2593:Post-transcription
1415:on 28 January 2013
1334:Medical Dictionary
1053:L-arabinose operon
730:repressor proteins
670:
631:: RNA Polymerase,
432:of protein-coding
187:André Michel Lwoff
77:strand and either
47:
18:Operator (biology)
3336:
3335:
3269:Eric F. Wieschaus
3231:
3230:
3049:Pattern formation
2953:Fitness landscape
2815:
2814:
2699:
2698:
2629:
2628:
2505:Special transfers
2428:
2427:
2383:RNA polymerase II
2251:
2250:
2209:
2208:
1780:978-0-13-191833-7
1728:978-0-19-854267-4
1626:10.1101/gr.737703
1542:978-0-7167-4939-4
1336:. MedicineNet.com
1127:978-0-7167-3136-8
1120:. W. H. Freeman.
1100:978-1-4292-1962-4
742:Alternatively, a
617:General structure
475:
474:
137:T7 RNA polymerase
73:together into an
16:(Redirected from
3361:
3279:William McGinnis
3248:Richard Lewontin
3243:C. H. Waddington
3115:
3114:
3092:Neutral networks
2842:
2835:
2828:
2819:
2818:
2640:
2639:
2537:
2536:
2455:
2448:
2441:
2432:
2431:
2305:Response element
2288:Response element
2101:
2100:
2029:
2028:
2000:
1993:
1986:
1977:
1976:
1945:
1944:
1899:
1893:
1892:
1882:
1850:
1844:
1843:
1833:
1823:
1791:
1785:
1784:
1766:
1760:
1759:
1757:
1755:
1739:
1733:
1732:
1711:Lewin B (1990).
1708:
1699:
1698:
1688:
1678:
1654:
1648:
1647:
1637:
1605:
1599:
1598:
1588:
1553:
1547:
1546:
1534:
1524:
1518:
1517:
1507:
1483:
1477:
1476:
1474:
1473:
1467:
1461:. Archived from
1444:
1434:
1425:
1424:
1422:
1420:
1401:
1395:
1394:
1384:
1352:
1346:
1345:
1343:
1341:
1326:
1320:
1319:
1309:
1285:
1276:
1275:
1265:
1242:The EMBO Journal
1233:
1227:
1226:
1190:
1184:
1183:
1173:
1141:
1132:
1131:
1111:
1105:
1104:
1084:
1032:
1027:
1026:
1018:
1013:
1012:
996:L. monocytogenes
881:structural genes
876:Escherichia coli
718:Structural genes
607:Asgard (archaea)
605:; the closer an
599:Asgard (archaea)
590:structural genes
555:structural genes
511:structural genes
464:(blue) regulate
423:
422:
412:
405:
398:
391:
384:
377:
370:
363:
356:
349:
342:
335:
328:
321:
314:
307:
300:
293:
286:
279:
272:
265:
257:
249:
242:
235:
228:
221:
212:
201:
104:(which includes
69:. The genes are
45:A typical operon
21:
3369:
3368:
3364:
3363:
3362:
3360:
3359:
3358:
3349:Gene expression
3339:
3338:
3337:
3332:
3323:
3302:
3289:Sean B. Carroll
3227:
3159:
3106:
3070:
3022:
3003:Maternal effect
2986:
2919:
2856:
2846:
2816:
2811:
2790:
2725:Transcriptional
2695:
2664:
2625:
2616:Polyadenylation
2587:
2561:
2526:
2520:Protein→Protein
2471:
2464:
2462:Gene expression
2459:
2429:
2424:
2399:
2393:
2387:
2352:
2326:
2247:
2205:
2186:
2169:DNA methylation
2163:
2107:
2092:
2018:
2004:
1970:Operon database
1954:
1949:
1948:
1911:(7249): 950–6.
1900:
1896:
1851:
1847:
1792:
1788:
1781:
1767:
1763:
1753:
1751:
1740:
1736:
1729:
1709:
1702:
1655:
1651:
1614:Genome Research
1606:
1602:
1554:
1550:
1543:
1525:
1521:
1484:
1480:
1471:
1469:
1465:
1442:
1435:
1428:
1418:
1416:
1403:
1402:
1398:
1367:(13): 4369–75.
1353:
1349:
1339:
1337:
1328:
1327:
1323:
1286:
1279:
1234:
1230:
1191:
1187:
1142:
1135:
1128:
1112:
1108:
1101:
1085:
1081:
1076:
1028:
1021:
1014:
1007:
1004:
986:Pascale Cossart
963:
931:
925:
863:
857:
760:gene regulation
756:
726:regulatory gene
697:– a segment of
619:
595:gene clustering
586:
551:
471:
470:
469:
424:
416:
415:
414:
413:
408:
406:
401:
399:
394:
392:
387:
385:
380:
378:
373:
371:
366:
364:
359:
357:
352:
350:
345:
343:
338:
336:
331:
329:
324:
322:
317:
315:
310:
308:
303:
301:
296:
294:
289:
287:
282:
280:
275:
273:
268:
266:
260:
258:
252:
250:
245:
243:
238:
236:
231:
229:
224:
222:
217:
213:
199:
181:was awarded to
149:
131:. For example,
39:
28:
23:
22:
15:
12:
11:
5:
3367:
3357:
3356:
3351:
3334:
3333:
3328:
3325:
3324:
3322:
3321:
3316:
3310:
3308:
3304:
3303:
3301:
3300:
3299:
3298:
3286:
3281:
3276:
3271:
3266:
3265:
3264:
3253:François Jacob
3250:
3245:
3239:
3237:
3233:
3232:
3229:
3228:
3226:
3225:
3220:
3215:
3210:
3205:
3200:
3195:
3190:
3189:
3188:
3178:
3173:
3167:
3165:
3161:
3160:
3158:
3157:
3152:
3147:
3142:
3137:
3132:
3127:
3121:
3119:
3112:
3108:
3107:
3105:
3104:
3099:
3094:
3089:
3084:
3078:
3076:
3072:
3071:
3069:
3068:
3063:
3058:
3053:
3052:
3051:
3046:
3036:
3030:
3028:
3024:
3023:
3021:
3020:
3015:
3010:
3005:
3000:
2994:
2992:
2988:
2987:
2985:
2984:
2982:Sequence space
2979:
2974:
2969:
2964:
2959:
2950:
2945:
2940:
2935:
2929:
2927:
2921:
2920:
2918:
2917:
2912:
2911:
2910:
2900:
2895:
2890:
2885:
2880:
2875:
2870:
2864:
2862:
2858:
2857:
2845:
2844:
2837:
2830:
2822:
2813:
2812:
2810:
2809:
2804:
2802:François Jacob
2798:
2796:
2792:
2791:
2789:
2788:
2787:
2786:
2781:
2771:
2766:
2765:
2764:
2759:
2754:
2744:
2739:
2738:
2737:
2732:
2722:
2721:
2720:
2709:
2707:
2701:
2700:
2697:
2696:
2694:
2693:
2688:
2683:
2678:
2672:
2670:
2666:
2665:
2663:
2662:
2657:
2652:
2646:
2644:
2637:
2631:
2630:
2627:
2626:
2624:
2623:
2618:
2613:
2608:
2603:
2597:
2595:
2589:
2588:
2586:
2585:
2580:
2578:RNA polymerase
2575:
2569:
2567:
2563:
2562:
2560:
2559:
2554:
2549:
2543:
2541:
2534:
2528:
2527:
2525:
2524:
2523:
2522:
2517:
2512:
2502:
2501:
2500:
2482:
2476:
2474:
2466:
2465:
2458:
2457:
2450:
2443:
2435:
2426:
2425:
2423:
2422:
2417:
2412:
2406:
2404:
2389:
2388:
2386:
2385:
2379:RNA polymerase
2373:
2367:RNA polymerase
2360:
2358:
2354:
2353:
2351:
2350:
2345:
2340:
2334:
2332:
2328:
2327:
2325:
2324:
2319:
2314:
2309:
2308:
2307:
2302:
2292:
2291:
2290:
2285:
2280:
2275:
2270:
2259:
2257:
2253:
2252:
2249:
2248:
2246:
2245:
2240:
2239:
2238:
2233:
2228:
2217:
2215:
2211:
2210:
2207:
2206:
2204:
2203:
2197:
2195:
2188:
2187:
2185:
2184:
2183:
2182:
2174:
2172:
2165:
2164:
2162:
2161:
2160:
2159:
2154:
2141:
2136:
2135:
2134:
2119:
2117:
2098:
2094:
2093:
2091:
2090:
2089:
2088:
2083:
2073:
2072:
2071:
2066:
2061:
2056:
2051:
2046:
2035:
2033:
2026:
2020:
2019:
2003:
2002:
1995:
1988:
1980:
1974:
1973:
1963:
1953:
1952:External links
1950:
1947:
1946:
1894:
1865:(5): 1216–21.
1845:
1806:(12): 6652–7.
1786:
1779:
1761:
1734:
1727:
1700:
1649:
1600:
1548:
1541:
1519:
1478:
1426:
1396:
1347:
1321:
1300:(3): 199–211.
1277:
1248:(8): 2023–31.
1228:
1185:
1133:
1126:
1106:
1099:
1078:
1077:
1075:
1072:
1071:
1070:
1068:Umu Chromotest
1065:
1060:
1055:
1050:
1045:
1040:
1034:
1033:
1030:Biology portal
1019:
1003:
1000:
962:
959:
927:Main article:
924:
917:
869:operon of the
859:Main article:
856:
849:
848:
847:
836:
816:
815:
801:regulator gene
793:
755:
752:
722:
721:
715:
692:
689:RNA polymerase
618:
615:
585:
582:
574:RNA polymerase
550:
547:
473:
472:
418:
417:
215:
214:
207:
206:
205:
204:
198:
195:
183:François Jacob
148:
145:
129:bacteriophages
95:co-transcribed
26:
9:
6:
4:
3:
2:
3366:
3355:
3352:
3350:
3347:
3346:
3344:
3331:
3326:
3320:
3317:
3315:
3312:
3311:
3309:
3305:
3297:
3296:
3292:
3291:
3290:
3287:
3285:
3282:
3280:
3277:
3275:
3272:
3270:
3267:
3263:
3260:
3259:
3258:
3257:Jacques Monod
3254:
3251:
3249:
3246:
3244:
3241:
3240:
3238:
3234:
3224:
3221:
3219:
3216:
3214:
3211:
3209:
3206:
3204:
3201:
3199:
3196:
3194:
3191:
3187:
3184:
3183:
3182:
3179:
3177:
3174:
3172:
3171:Homeotic gene
3169:
3168:
3166:
3162:
3156:
3153:
3151:
3148:
3146:
3143:
3141:
3138:
3136:
3133:
3131:
3128:
3126:
3123:
3122:
3120:
3116:
3113:
3109:
3103:
3100:
3098:
3095:
3093:
3090:
3088:
3085:
3083:
3080:
3079:
3077:
3073:
3067:
3064:
3062:
3059:
3057:
3054:
3050:
3047:
3045:
3042:
3041:
3040:
3039:Morphogenesis
3037:
3035:
3032:
3031:
3029:
3025:
3019:
3016:
3014:
3011:
3009:
3006:
3004:
3001:
2999:
2996:
2995:
2993:
2989:
2983:
2980:
2978:
2975:
2973:
2970:
2968:
2965:
2963:
2960:
2958:
2954:
2951:
2949:
2946:
2944:
2941:
2939:
2936:
2934:
2931:
2930:
2928:
2926:
2922:
2916:
2913:
2909:
2906:
2905:
2904:
2901:
2899:
2896:
2894:
2891:
2889:
2886:
2884:
2881:
2879:
2876:
2874:
2873:Reaction norm
2871:
2869:
2866:
2865:
2863:
2859:
2855:
2851:
2843:
2838:
2836:
2831:
2829:
2824:
2823:
2820:
2808:
2807:Jacques Monod
2805:
2803:
2800:
2799:
2797:
2793:
2785:
2782:
2780:
2777:
2776:
2775:
2772:
2770:
2769:Translational
2767:
2763:
2760:
2758:
2755:
2753:
2750:
2749:
2748:
2745:
2743:
2740:
2736:
2733:
2731:
2728:
2727:
2726:
2723:
2719:
2716:
2715:
2714:
2711:
2710:
2708:
2706:
2702:
2692:
2689:
2687:
2684:
2682:
2679:
2677:
2674:
2673:
2671:
2667:
2661:
2658:
2656:
2653:
2651:
2648:
2647:
2645:
2641:
2638:
2636:
2632:
2622:
2619:
2617:
2614:
2612:
2609:
2607:
2604:
2602:
2599:
2598:
2596:
2594:
2590:
2584:
2581:
2579:
2576:
2574:
2571:
2570:
2568:
2564:
2558:
2555:
2553:
2550:
2548:
2545:
2544:
2542:
2538:
2535:
2533:
2532:Transcription
2529:
2521:
2518:
2516:
2513:
2511:
2508:
2507:
2506:
2503:
2499:
2495:
2491:
2488:
2487:
2486:
2485:Central dogma
2483:
2481:
2478:
2477:
2475:
2473:
2467:
2463:
2456:
2451:
2449:
2444:
2442:
2437:
2436:
2433:
2421:
2418:
2416:
2413:
2411:
2408:
2407:
2405:
2402:
2397:
2390:
2384:
2380:
2377:
2374:
2372:
2368:
2365:
2362:
2361:
2359:
2355:
2349:
2346:
2344:
2341:
2339:
2336:
2335:
2333:
2329:
2323:
2320:
2318:
2315:
2313:
2310:
2306:
2303:
2301:
2298:
2297:
2296:
2293:
2289:
2286:
2284:
2281:
2279:
2276:
2274:
2271:
2269:
2266:
2265:
2264:
2261:
2260:
2258:
2254:
2244:
2241:
2237:
2234:
2232:
2229:
2227:
2224:
2223:
2222:
2219:
2218:
2216:
2212:
2202:
2199:
2198:
2196:
2193:
2189:
2181:
2178:
2177:
2176:
2175:
2173:
2170:
2166:
2158:
2155:
2153:
2150:
2147:
2146:
2145:
2142:
2140:
2137:
2133:
2130:
2129:
2128:
2124:
2121:
2120:
2118:
2115:
2111:
2106:
2102:
2099:
2095:
2087:
2086:trp repressor
2084:
2082:
2081:lac repressor
2079:
2078:
2077:
2074:
2070:
2067:
2065:
2062:
2060:
2057:
2055:
2052:
2050:
2047:
2045:
2042:
2041:
2040:
2037:
2036:
2034:
2030:
2027:
2025:
2021:
2016:
2012:
2008:
2007:Transcription
2001:
1996:
1994:
1989:
1987:
1982:
1981:
1978:
1971:
1967:
1964:
1962:
1960:
1956:
1955:
1942:
1938:
1934:
1930:
1926:
1922:
1918:
1914:
1910:
1906:
1898:
1890:
1886:
1881:
1876:
1872:
1868:
1864:
1860:
1856:
1849:
1841:
1837:
1832:
1827:
1822:
1817:
1813:
1809:
1805:
1801:
1797:
1790:
1782:
1776:
1772:
1765:
1749:
1745:
1738:
1730:
1724:
1720:
1716:
1715:
1707:
1705:
1696:
1692:
1687:
1682:
1677:
1672:
1668:
1664:
1660:
1653:
1645:
1641:
1636:
1631:
1627:
1623:
1620:(5): 875–82.
1619:
1615:
1611:
1604:
1596:
1592:
1587:
1582:
1578:
1574:
1571:(6031): 767.
1570:
1566:
1562:
1558:
1552:
1544:
1538:
1533:
1532:
1523:
1515:
1511:
1506:
1501:
1497:
1493:
1489:
1482:
1468:on 2016-03-04
1464:
1460:
1456:
1453:(6): 1727–9.
1452:
1448:
1440:
1433:
1431:
1414:
1410:
1406:
1400:
1392:
1388:
1383:
1378:
1374:
1370:
1366:
1362:
1358:
1351:
1335:
1331:
1325:
1317:
1313:
1308:
1303:
1299:
1295:
1291:
1284:
1282:
1273:
1269:
1264:
1259:
1255:
1251:
1247:
1243:
1239:
1232:
1224:
1220:
1216:
1212:
1208:
1204:
1201:(3): 521–32.
1200:
1196:
1189:
1181:
1177:
1172:
1167:
1163:
1159:
1155:
1151:
1147:
1140:
1138:
1129:
1123:
1119:
1118:
1110:
1102:
1096:
1092:
1091:
1083:
1079:
1069:
1066:
1064:
1061:
1059:
1056:
1054:
1051:
1049:
1046:
1044:
1041:
1039:
1036:
1035:
1031:
1025:
1020:
1017:
1011:
1006:
999:
997:
993:
992:
987:
983:
979:
976:
972:
970:
969:
958:
956:
955:corepressible
952:
948:
944:
940:
936:
935:Jacques Monod
930:
922:
916:
914:
913:derepressible
910:
906:
902:
898:
894:
890:
886:
882:
878:
877:
872:
868:
862:
854:
845:
841:
837:
833:
829:
825:
824:
823:
821:
813:
809:
807:
802:
798:
794:
791:
787:
785:
780:
776:
772:
771:
770:
768:
763:
761:
751:
749:
745:
740:
738:
733:
731:
727:
719:
716:
712:
708:
704:
700:
696:
693:
690:
686:
682:
678:
675:
674:
673:
667:
663:
660:
658:
654:
650:
646:
642:
638:
635:: Repressor,
634:
630:
625:
621:
614:
612:
608:
604:
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581:
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559:polycistronic
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459:
458:transcription
455:
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411:
404:
397:
390:
383:
376:
375:RBS
369:
368:RBS
362:
355:
354:Transcription
348:
341:
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327:
320:
313:
306:
299:
292:
285:
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264:
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191:Jacques Monod
188:
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130:
125:
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122:polycistronic
119:
115:
111:
107:
103:
98:
96:
92:
88:
87:monocistronic
84:
80:
76:
72:
68:
64:
60:
56:
52:
43:
37:
33:
19:
3293:
3186:eyeless gene
3082:Evolvability
3056:Segmentation
2933:Canalisation
2903:Heterochrony
2893:Heritability
2887:
2861:Key concepts
2784:irreversible
2669:Key elements
2566:Key elements
2480:Genetic code
2470:Introduction
2038:
1958:
1908:
1904:
1897:
1862:
1858:
1848:
1803:
1799:
1789:
1770:
1764:
1752:. Retrieved
1747:
1737:
1713:
1666:
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1652:
1617:
1613:
1603:
1568:
1564:
1559:(May 2011).
1551:
1530:
1522:
1495:
1491:
1481:
1470:. Retrieved
1463:the original
1450:
1446:
1417:. Retrieved
1413:the original
1408:
1399:
1364:
1360:
1350:
1338:. Retrieved
1333:
1324:
1297:
1293:
1245:
1241:
1231:
1198:
1194:
1188:
1153:
1149:
1116:
1109:
1089:
1082:
1043:Genetic code
995:
989:
984:
980:
977:
973:
966:
964:
938:
932:
920:
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874:
866:
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852:
839:
827:
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805:
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783:
774:
764:
757:
741:
734:
723:
710:
694:
671:
665:
661:
656:
652:
648:
644:
643:: Operator,
640:
639:: Promoter,
636:
632:
628:
627:
620:
587:
577:
552:
532:
524:corepressors
497:
491:
485:, including
476:
429:
218:
177:. The 1965
172:
166:
150:
126:
99:
94:
54:
48:
3284:Mike Levine
3193:Distal-less
3018:Polyphenism
2998:Epigenetics
2850:development
2635:Translation
2472:to genetics
2392:Termination
2268:Pribnow box
2236:Corepressor
2231:Coactivator
2032:prokaryotic
1754:30 December
1419:30 December
1340:30 December
943:allolactose
909:allolactose
835:activators.
790:allolactose
744:corepressor
701:to which a
685:transcribed
647:: Lactose,
479:prokaryotes
466:translation
427:prokaryotic
403:Translation
102:prokaryotes
71:transcribed
3343:Categories
3262:Lac operon
3087:Robustness
3066:Modularity
3061:Metamerism
2967:Plasticity
2962:Pleiotropy
2915:Heterotopy
2779:reversible
2742:lac operon
2718:imprinting
2713:Epigenetic
2705:Regulation
2660:Eukaryotic
2606:5' capping
2557:Eukaryotic
2420:Rho factor
2410:Terminator
2401:eukaryotic
2376:eukaryotic
2357:Elongation
2343:Eukaryotic
2331:Initiation
2114:nucleosome
2097:eukaryotic
2069:gal operon
2064:ara operon
2059:Gua Operon
2054:gab operon
2049:trp operon
2044:lac operon
2015:Eukaryotic
1472:2015-08-27
1074:References
951:attenuator
929:trp operon
889:terminator
873:bacterium
861:lac operon
812:tryptophan
754:Regulation
748:trp operon
707:lac operon
681:nucleotide
611:eukaryotes
609:is to the
528:activators
520:repressors
492:C. elegans
483:eukaryotes
347:Terminator
340:Stop
333:Stop
326:Start
319:Start
118:eukaryotes
106:organelles
85:to create
79:translated
3213:Morphogen
3198:Engrailed
3181:Pax genes
3102:Tinkering
2948:Epistasis
2943:Dominance
2854:phenotype
2650:Bacterial
2547:Bacterial
2396:bacterial
2364:bacterial
2338:Bacterial
2312:Insulator
2256:Promotion
2226:Activator
2076:Repressor
2011:Bacterial
1742:Mayer G.
1514:2002-4436
891:, and an
844:inhibitor
820:activator
767:repressor
703:repressor
539:stimulons
507:chordates
487:nematodes
157:repressor
133:T7 phages
91:expressed
3176:Hox gene
3164:Elements
3145:Homeobox
2762:microRNA
2676:Ribosome
2655:Archaeal
2611:Splicing
2583:Promoter
2552:Archaeal
2496: →
2492: →
2317:Silencer
2295:Enhancer
2283:CAAT box
2273:TATA box
2263:Promoter
1933:19448609
1889:11222772
1840:10823905
1714:Genes IV
1695:38314098
1686:10833476
1669:: 1–16.
1644:12695325
1595:21566161
1459:14406329
1391:15205439
1316:15642184
1223:26918553
1180:30799038
1063:TATA box
1002:See also
893:operator
885:promoter
695:Operator
677:Promoter
659:: lacA.
655:: lacY,
651:: lacZ,
588:All the
578:operator
570:promoter
543:modulons
535:regulons
515:promoter
489:such as
454:enhancer
450:operator
446:Promoter
305:UTR
298:ORF
291:ORF
277:Promoter
270:Operator
263:silencer
255:silencer
247:Enhancer
240:Enhancer
197:Overview
161:operator
114:bacteria
110:plastids
83:splicing
67:promoter
51:genetics
3354:Operons
3307:Debates
3118:Systems
3044:Eyespot
2908:Neoteny
2515:RNA→DNA
2510:RNA→RNA
2498:Protein
2243:Inducer
2110:histone
1941:4341657
1913:Bibcode
1808:Bibcode
1663:Archaea
1573:Bibcode
1565:Science
1557:Jacob F
1272:9155028
1263:1169805
1215:8098272
1171:7295392
975:bases.
968:E. coli
957:model.
939:E. coli
905:lactose
901:glucose
832:inducer
779:inducer
737:inducer
603:Archaea
566:protein
444:(red).
174:E. coli
147:History
32:Opteron
3208:Ligand
2888:Operon
2039:Operon
1939:
1931:
1905:Nature
1887:
1877:
1838:
1828:
1777:
1725:
1719:243–58
1693:
1683:
1642:
1635:430880
1632:
1593:
1539:
1512:
1457:
1389:
1382:421614
1379:
1314:
1270:
1260:
1221:
1213:
1178:
1168:
1124:
1097:
923:operon
895:. The
855:operon
808:operon
786:operon
714:genes.
666:Bottom
526:, and
430:operon
169:operon
55:operon
36:Oberon
2643:Types
2540:Types
2300:E-box
2152:HDAC1
1937:S2CID
1880:29727
1831:18690
1498:(1).
1466:(PDF)
1443:(PDF)
1219:S2CID
871:model
434:genes
141:lysis
108:like
63:genes
53:, an
2848:The
2371:rpoB
2214:both
2201:CHD7
2132:EZH2
1929:PMID
1885:PMID
1836:PMID
1775:ISBN
1756:2012
1723:ISBN
1691:PMID
1667:2023
1640:PMID
1591:PMID
1537:ISBN
1510:ISSN
1455:PMID
1421:2012
1387:PMID
1342:2012
1312:PMID
1268:PMID
1211:PMID
1195:Cell
1176:PMID
1150:Cell
1122:ISBN
1095:ISBN
919:The
903:and
887:, a
883:, a
865:The
851:The
679:– a
562:mRNA
541:and
503:rRNA
452:and
189:and
75:mRNA
2852:of
2494:RNA
2490:DNA
2278:BRE
1966:OBD
1921:doi
1909:459
1875:PMC
1867:doi
1826:PMC
1816:doi
1681:PMC
1671:doi
1630:PMC
1622:doi
1581:doi
1569:332
1500:doi
1451:250
1377:PMC
1369:doi
1365:186
1302:doi
1258:PMC
1250:doi
1203:doi
1166:PMC
1158:doi
1154:176
921:trp
897:lac
867:lac
853:lac
838:In
826:In
806:trp
795:In
784:lac
773:In
735:An
711:lac
699:DNA
662:Top
171:in
167:lac
59:DNA
49:In
34:or
3345::
3255:+
2381::
2369::
2116:):
2013:,
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1935:.
1927:.
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1907:.
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1873:.
1863:29
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1857:.
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1824:.
1814:.
1804:97
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1798:.
1746:.
1721:.
1703:^
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1429:^
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1310:.
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1280:^
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1240:.
1217:.
1209:.
1199:73
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1174:.
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1152:.
1148:.
1136:^
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1673::
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1502::
1496:4
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1371::
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1318:.
1304::
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1160::
1130:.
1103:.
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637:3
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261:/
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