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Promoter (genetics)

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415: 426:. In that study, two main forms of interference were measured. One is when an RNAP is on the downstream promoter, blocking the movement of RNAPs elongating from the upstream promoter. The other is when the two promoters are so close that when an RNAP sits on one of the promoters, it blocks any other RNAP from reaching the other promoter. These events are possible because the RNAP occupies several nucleotides when bound to the DNA, including in transcription start sites. Similar events occur when the promoters are in divergent and convergent formations. The possible events also depend on the distance between them. 644: 411:
divergent, tandem, and convergent directions. They can also be regulated by transcription factors and differ in various features, such as the nucleotide distance between them, the two promoter strengths, etc. The most important aspect of two closely spaced promoters is that they will, most likely, interfere with each other. Several studies have explored this using both analytical and stochastic models. There are also studies that measured gene expression in synthetic genes or from one to a few genes controlled by bidirectional promoters.
751:), with one member of the dimer anchored to its binding motif on the enhancer and the other member anchored to its binding motif on the promoter (represented by the red zigzags in the illustration). Several cell function specific transcription factors (there are about 1,600 transcription factors in a human cell) generally bind to specific motifs on an enhancer and a small combination of these enhancer-bound transcription factors, when brought close to a promoter by a DNA loop, govern the level of transcription of the target gene. 878:, and ACTACAnnTCCC are represented in bidirectional promoters at significantly higher rates than in unidirectional promoters. The absence of TATA boxes in bidirectional promoters suggests that TATA boxes play a role in determining the directionality of promoters, but counterexamples of bidirectional promoters do possess TATA boxes and unidirectional promoters without them indicates that they cannot be the only factor. 20: 588: 759:
may activate it and that activated transcription factor may then activate the enhancer to which it is bound (see small red star representing phosphorylation of transcription factor bound to enhancer in the illustration). An activated enhancer begins transcription of its RNA before activating a promoter to initiate transcription of messenger RNA from its target gene.
114:: 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. 995:. Importantly, intervention in the number or structure of promoter-bound proteins is one key to treating a disease without affecting expression of unrelated genes sharing elements with the target gene. Some genes whose change is not desirable are capable of influencing the potential of a cell to become cancerous. 953:
The initiation of the transcription is a multistep sequential process that involves several mechanisms: promoter location, initial reversible binding of RNA polymerase, conformational changes in RNA polymerase, conformational changes in DNA, binding of nucleoside triphosphate (NTP) to the functional
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The above consensus sequences, while conserved on average, are not found intact in most promoters. On average, only 3 to 4 of the 6 base pairs in each consensus sequence are found in any given promoter. Few natural promoters have been identified to date that possess intact consensus sequences at both
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of bidirectional promoter regions has been shown to downregulate both genes, and demethylation to upregulate both genes. There are exceptions to this, however. In some cases (about 11%), only one gene of a bidirectional pair is expressed. In these cases, the promoter is implicated in suppression of
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are regions of the genome that are major gene-regulatory elements. Enhancers control cell-type-specific gene expression programs, most often by looping through long distances to come in physical proximity with the promoters of their target genes. In a study of brain cortical neurons, 24,937 loops
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Enhancers, when active, are generally transcribed from both strands of DNA with RNA polymerases acting in two different directions, producing two eRNAs as illustrated in the Figure. An inactive enhancer may be bound by an inactive transcription factor. Phosphorylation of the transcription factor
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Gene promoters are typically located upstream of the gene and can have regulatory elements several kilobases away from the transcriptional start site (enhancers). In eukaryotes, the transcriptional complex can cause the DNA to bend back on itself, which allows for placement of regulatory sequences
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in a bidirectional gene pair. A "bidirectional gene pair" refers to two adjacent genes coded on opposite strands, with their 5' ends oriented toward one another. The two genes are often functionally related, and modification of their shared promoter region allows them to be co-regulated and thus
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in CpG islands of promoters causes stable silencing of genes. However, the presence or absence of the other elements have relatively small effects on gene expression in experiments. Two sequences, the TATA box and Inr, caused small but significant increases in expression (45% and 28% increases,
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Promoters can be very closely located in the DNA. Such "closely spaced promoters" have been observed in the DNAs of all life forms, from humans to prokaryotes and are highly conserved. Therefore, they may provide some (presently unknown) advantages. These pairs of promoters can be positioned in
1081:. The presence of multiple methylated CpG sites in CpG islands of promoters causes stable silencing of genes. Silencing of a gene may be initiated by other mechanisms, but this is often followed by methylation of CpG sites in the promoter CpG island to cause the stable silencing of the gene. 188:
Artificial promoters with conserved -10 and -35 elements transcribe more slowly. All DNAs have "Closely spaced promoters". Divergent, tandem, and convergent orientations are possible. Two closely spaced promoters will likely interfere. Regulatory elements can be several kilobases away from the
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Hypermethylation downregulates both genes, while demethylation upregulates them. Non-coding RNAs are linked to mRNA promoter regions. Subgenomic promoters range from 24 to 100 nucleotides (Beet necrotic yellow vein virus). Gene expression depends on promoter binding. Unwanted gene changes can
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Most diseases are heterogeneous in cause, meaning that one "disease" is often many different diseases at the molecular level, though symptoms exhibited and response to treatment may be identical. How diseases of different molecular origin respond to treatments is partially addressed in the
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MicroRNA promoters often contain CpG islands. DNA methylation forms 5-methylcytosines at the 5' pyrimidine ring of CpG cytosine residues. Some cancer genes are silenced by mutation, but most are silenced by DNA methylation. Others are regulated promoters. Selection may favor less energetic
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methods. A promoter region is located before the -35 and -10 Consensus sequences. The closer the promoter region is to the consensus sequences the more often transcription of that gene will take place. There is not a set pattern for promoter regions as there are for consensus sequences.
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However, natural selection may favor less energetic binding as a way of regulating transcriptional output. In this case, we may call the most common sequence in a population the wild-type sequence. It may not even be the most advantageous sequence to have under prevailing conditions.
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were found, bringing enhancers to promoters. Multiple enhancers, each often at tens or hundred of thousands of nucleotides distant from their target genes, loop to their target gene promoters and coordinate with each other to control expression of their common target gene.
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shows high affinity to non-specific sites of the DNA, this characteristic does not allow us to clarify the process of promoter location. This process of promoter location has been attributed to the structure of the holoenzyme to DNA and sigma 4 to DNA complexes.
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usually begins bidirectionally, but divergent transcription is halted at a checkpoint later during elongation. Possible mechanisms behind this regulation include sequences in the promoter region, chromatin modification, and the spatial orientation of the DNA.
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that are localized in DNA regions distant from the promoters of genes can have very large effects on gene expression, with some genes undergoing up to 100-fold increased expression due to such a cis-regulatory module. These cis-regulatory modules include
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In eukaryotes, the transcriptional complex can bend DNA, allowing regulatory sequences to be placed far from the transcription site. The distal promoter is upstream of the gene and may contain additional regulatory elements with a weaker influence.
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Copland JA, Sheffield-Moore M, Koldzic-Zivanovic N, Gentry S, Lamprou G, Tzortzatou-Stathopoulou F, et al. (June 2009). "Sex steroid receptors in skeletal differentiation and epithelial neoplasia: is tissue-specific intervention possible?".
704:(present in about 49% of promoters), upstream and downstream TFIIB recognition elements (BREu and BREd) (present in about 22% of promoters), and downstream core promoter element (DPE) (present in about 12% of promoters). The presence of multiple 109:: The transcription of the gene is 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 the mRNA encoding the lactase gene. 391:← upstream downstream β†’ 5'-XXXXXXXPPPPPPXXXXXXPPPPPPXXXXGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGGXXXX-3' -35 -10 Gene to be transcribed 295:) to direct the level of transcription of a given gene. A promoter is induced in response to changes in abundance or conformation of regulatory proteins in a cell, which enable activating transcription factors to recruit RNA polymerase. 683:
shown by a small red star on a transcription factor on the enhancer) the enhancer is activated and can now activate its target promoter. The active enhancer is transcribed on each strand of DNA in opposite directions by bound RNAP IIs.
755:(a complex usually consisting of about 26 proteins in an interacting structure) communicates regulatory signals from enhancer DNA-bound transcription factors directly to the RNA polymerase II (pol II) enzyme bound to the promoter. 811:
Some functional classes of genes are more likely to be bidirectionally paired than others. Genes implicated in DNA repair are five times more likely to be regulated by bidirectional promoters than by unidirectional promoters.
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The schematic illustration in this section shows an enhancer looping around to come into close physical proximity with the promoter of a target gene. The loop is stabilized by a dimer of a connector protein (e.g. dimer of
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The promoter binding process is crucial in the understanding of the process of gene expression. Tuning synthetic genetic systems relies on precisely engineered synthetic promoters with known levels of transcription rates.
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analysis has shown bidirectionally paired genes to be co-expressed to a higher degree than random genes or neighboring unidirectional genes. Although co-expression does not necessarily indicate co-regulation,
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of the gene. The loop is stabilized by one architectural protein anchored to the enhancer and one anchored to the promoter and these proteins are joined to form a dimer (red zigzags). Specific regulatory
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Koyanagi KO, Hagiwara M, Itoh T, Gojobori T, Imanishi T (July 2005). "Comparative genomics of bidirectional gene pairs and its implications for the evolution of a transcriptional regulation system".
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Sneppen K, Dodd IB, Shearwin KE, Palmer AC, Schubert RA, Callen BP, Egan JB (February 2005). "A mathematical model for transcriptional interference by RNA polymerase traffic in Escherichia coli".
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Ross W, Gosink KK, Salomon J, Igarashi K, Zou C, Ishihama A, et al. (November 1993). "A third recognition element in bacterial promoters: DNA binding by the alpha subunit of RNA polymerase".
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the -10 and -35; artificial promoters with complete conservation of the -10 and -35 elements have been found to transcribe at lower frequencies than those with a few mismatches with the consensus.
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Burchard EG, Silverman EK, Rosenwasser LJ, Borish L, Yandava C, Pillari A, et al. (September 1999). "Association between a sequence variant in the IL-4 gene promoter and FEV(1) in asthma".
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Martins L, MΓ€kelΓ€ J, HΓ€kkinen A, Kandhavelu M, Yli-Harja O, Fonseca JM, Ribeiro AS (May 2012). "Dynamics of transcription of closely spaced promoters in Escherichia coli, one event at a time".
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Up-regulated expression of genes in mammals is initiated when signals are transmitted to the promoters associated with the genes. Promoter DNA sequences may include different elements such as
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In the case of a transcription factor binding site, there may be a single sequence that binds the protein most strongly under specified cellular conditions. This might be called canonical.
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Petrij F, Giles RH, Dauwerse HG, Saris JJ, Hennekam RC, Masuno M, et al. (July 1995). "Rubinstein-Taybi syndrome caused by mutations in the transcriptional co-activator CBP".
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Korbel JO, Jensen LJ, von Mering C, Bork P (July 2004). "Analysis of genomic context: prediction of functional associations from conserved bidirectionally transcribed gene pairs".
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Eukaryotic promoter regulatory sequences typically bind proteins called transcription factors that are involved in the formation of the transcriptional complex. An example is the
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Hobbs K, Negri J, Klinnert M, Rosenwasser LJ, Borish L (December 1998). "Interleukin-10 and transforming growth factor-beta promoter polymorphisms in allergies and asthma".
1117: 1090: 688:(a complex consisting of about 26 proteins in an interacting structure) communicates regulatory signals from the enhancer DNA-bound transcription factors to the promoter. 824:
and cellular metabolic genes are regulated by bidirectional promoters. The overrepresentation of bidirectionally paired DNA repair genes associates these promoters with
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respectively). The BREu and the BREd elements significantly decreased expression by 35% and 20%, respectively, and the DPE element had no detected effect on expression.
858:, and a symmetry around the midpoint of dominant Cs and As on one side and Gs and Ts on the other. A motif with the consensus sequence of TCTCGCGAGA, also called the 1188:. For example, to overexpress an important gene in a network, to yield higher production of target protein, synthetic biologists design promoters to upregulate its 1136: 1980:
Bordoy AE, Varanasi US, Courtney CM, Chatterjee A (December 2016). "Transcriptional Interference in Convergent Promoters as a Means for Tunable Gene Expression".
918:(sgRNA) as one of the common infection techniques used by these viruses and generally transcribe late viral genes. Subgenomic promoters range from 24 nucleotide ( 1093:. Although silencing of some genes in cancers occurs by mutation, a large proportion of carcinogenic gene silencing is a result of altered DNA methylation (see 934:
A wide variety of algorithms have been developed to facilitate detection of promoters in genomic sequence, and promoter prediction is a common element of many
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5'-AAAAAARNR-3' when centered in the -42 region; consensus sequence 5'-AWWWWWTTTTT-3' when centered in the -52 region; W = A or T; R = A or G; N = any base).
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Variations in promoters or transcription factors cause some diseases. Misunderstandings can result from using a canonical sequence to describe a promoter.
1661:"Bacterial promoter architecture: subsite structure of UP elements and interactions with the carboxy-terminal domain of the RNA polymerase alpha subunit" 835:
seem to be regulated by bidirectional promoters – significantly more than non-cancer causing genes. Hypermethylation of the promoters between gene pairs
477:). Some promoters that are targeted by multiple transcription factors might achieve a hyperactive state, leading to increased transcriptional activity. 3254:"Genome-wide analysis of the transcription factor binding preference of human bi-directional promoters and functional annotation of related gene pairs" 2304: 1135:
Silencing of DNA repair genes through methylation of CpG islands in their promoters appears to be especially important in progression to cancer (see
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to refer to a promoter is often problematic, and can lead to misunderstandings about promoter sequences. Canonical implies perfect, in some sense.
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are frequently associated with the promoter regions of mRNA-encoding genes. It has been hypothesized that the recruitment and initiation of
1520:"Promoter organization of the interferon-A genes differentially affects virus-induced expression and responsiveness to TBK1 and IKKepsilon" 4655: 4162:– a research project with an aim to generate 160 fully characterized, human DNA promoters of less than 4 kb (MiniPromoters) to drive 4127: 2155:"New core promoter element in RNA polymerase II-dependent transcription: sequence-specific DNA binding by transcription factor IIB" 608:
Text about mammals highly duplicated among uses of the same picture -- can we make a "canonical" version and redirect people there?
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As promoters are typically immediately adjacent to the gene in question, positions in the promoter are designated relative to the
4200: 458:. The TATA element and BRE typically are located close to the transcriptional start site (typically within 30 to 40 base pairs). 568:
Anything further upstream (but not an enhancer or other regulatory region whose influence is positional/orientation independent)
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Many other elements/motifs may be present. There is no such thing as a set of "universal elements" found in every core promoter.
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Vlahopoulos SA, Logotheti S, Mikas D, Giarika A, Gorgoulis V, Zoumpourlis V (April 2008). "The role of ATF-2 in oncogenesis".
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system (Ptac). Notice how tac is written as a tac promoter, while in fact tac is actually both a promoter and an operator.
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to upregulate transcription of one gene, or remove bound transcription factors to downregulate transcription of one gene.
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Given the short sequences of most promoter elements, promoters can rapidly evolve from random sequences. For instance, in
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In humans, about 70% of promoters located near the transcription start site of a gene (proximal promoters) contain a
630: 555: 3111:"Comprehensive annotation of bidirectional promoters identifies co-regulation among breast and ovarian cancer genes" 1053:, where the CpG island-containing promoter is located about 5,400 nucleotides upstream of the coding region of the 4949: 4893: 4550: 3770:"A genome-wide analysis of CpG dinucleotides in the human genome distinguishes two distinct classes of promoters" 3646:
Borukhov S, Nudler E (April 2003). "RNA polymerase holoenzyme: structure, function and biological implications".
3004:"Enhancer RNAs predict enhancer-gene regulatory links and are critical for enhancer function in neuronal systems" 2906:"The degree of enhancer or promoter activity is reflected by the levels and directionality of eRNA transcription" 987:
Not listed here are the many kinds of cancers involving aberrant transcriptional regulation owing to creation of
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Proximal promoter – the proximal sequence upstream of the gene that tends to contain primary regulatory elements
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with only one mutation, and that ~10% of random sequences can serve as active promoters even without evolution.
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Spitz F, Furlong EE (September 2012). "Transcription factors: from enhancer binding to developmental control".
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assays have shown that over half of human genes do not have a strong directional bias. Research suggests that
197:(RNAP II) bound to the transcription start site promoter can start mRNA synthesis. It also typically contains 181:. RNA polymerase must attach to DNA near a gene for transcription to occur. Promoter DNA sequences provide an 4976: 4907: 4648: 4598: 3401:"A methyl-sensitive element induces bidirectional transcription in TATA-less CpG island-associated promoters" 1298: 1233: 1232:
Some promoters are called constitutive as they are active in all circumstances in the cell, while others are
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can be used to design neutral DNA or insulators that do not trigger gene expression of downstream sequences.
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binding site. The -10 sequence is TATAAT. -35 sequences are conserved on average, but not in most promoters.
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Schoenfelder S, Fraser P (August 2019). "Long-range enhancer-promoter contacts in gene expression control".
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In humans, DNA methylation occurs at the 5' position of the pyrimidine ring of the cytosine residues within
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long, the sequence of which is highly dependent on the gene and product of transcription, type or class of
1371:"Automated model-predictive design of synthetic promoters to control transcriptional profiles in bacteria" 1204:
Some cases of many genetic diseases are associated with variations in promoters or transcription factors.
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far from the actual site of transcription. Eukaryotic RNA-polymerase-II-dependent promoters can contain a
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The process is more complicated, and at least seven different factors are necessary for the binding of an
4193: 388:. RNA polymerase holoenzymes containing other sigma factors recognize different core promoter sequences. 790:
database shared at least one database-assigned functional category with their partners 47% of the time.
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sequences of corresponding nucleotides that attach to specific promoters and regulate gene expression.
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Carullo NV, Phillips Iii RA, Simon RC, Soto SA, Hinds JE, Salisbury AJ, et al. (September 2020).
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Liefke R, Windhof-Jaidhauser IM, Gaedcke J, Salinas-Riester G, Wu F, Ghadimi M, Dango S (June 2015).
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Mikhaylichenko O, Bondarenko V, Harnett D, Schor IE, Males M, Viales RR, Furlong EE (January 2018).
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Certain sequence characteristics have been observed in bidirectional promoters, including a lack of
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the non-expressed gene. The mechanism behind this could be competition for the same polymerases, or
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Depiction the phenomenon of interference between tandem promoters. Figure created with BioRender.com
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Weingarten-Gabbay S, Nir R, Lubliner S, Sharon E, Kalma Y, Weinberger A, Segal E (February 2019).
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Distal promoters also frequently contain CpG islands, such as the promoter of the DNA repair gene
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Chauhan V, Bahrudeen MN, Palma CS, Baptista IS, Almeida BL, Dash S, et al. (January 2022).
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RNA polymerase-promoter complex, and nonproductive and productive initiation of RNA synthesis.
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Core promoter – the minimal portion of the promoter required to properly initiate transcription
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Hossain A, Lopez E, Halper SM, Cetnar DP, Reis AC, Strickland D, et al. (December 2020).
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bind to the promoter. When a transcription factor is activated by a signal (here indicated as
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Promoters represent critical elements that can work in concert with other regulatory regions (
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Weintraub AS, Li CH, Zamudio AV, Sigova AA, Hannett NM, Day DS, et al. (December 2017).
2320:"Regulation of gene expression via the core promoter and the basal transcriptional machinery" 2298: 2090: 729: 728:
and tethering elements. Among this constellation of elements, enhancers and their associated
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transcript from the DNA downstream of the promoter. The RNA transcript may encode a protein (
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Vaishnav ED, de Boer CG, Molinet J, Yassour M, Fan L, Adiconis X, et al. (March 2022).
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Beagan JA, Pastuzyn ED, Fernandez LR, Guo MH, Feng K, Titus KR, et al. (June 2020).
2255:"The oxidative demethylase ALKBH3 marks hyperactive gene promoters in human cancer cells" 1283: 735: 721: 717: 651: 643: 284: 4178: 4088: 3785: 3416: 3126: 3060:
Trinklein ND, Aldred SF, Hartman SJ, Schroeder DI, Otillar RP, Myers RM (January 2004).
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Lambert SA, Jolma A, Campitelli LF, Das PK, Yin Y, Albu M, et al. (February 2018).
2215: 2032: 1899: 1766: 1629: 1576: 1443: 1386: 1336:"Analysis of Biological Networks: Transcriptional Networks – Promoter Sequence Analysis" 4920: 4803: 4266: 4108: 3946: 3853: 3828: 3804: 3769: 3750: 3707: 3484: 3459: 3435: 3400: 3376: 3351: 3280: 3253: 3199: 3145: 3110: 3028: 3003: 2930: 2905: 2881: 2856: 2832: 2797: 2732: 2707: 2688: 2642: 2594: 2567: 2543: 2518: 2491: 2466: 2393: 2368: 2344: 2319: 2281: 2254: 2235: 2051: 2016: 1957: 1930: 1833: 1593: 1560: 1460: 1427: 1403: 1370: 1256:; i.e., the lac promoter is IPTG inducible, meaning that besides the lac promoter, the 1253: 1244:
A tissue-specific promoter is a promoter that has activity in only certain cell types.
1148: 862:, was recently shown to drive PolII-driven bidirectional transcription in CpG islands. 817: 813: 516: 440: 373: 356: 232:, must attach to the DNA near a gene. Promoters contain specific DNA sequences such as 3659: 3623: 3606: 3582: 3557: 3533: 3508: 3186: 3169: 3086: 3061: 2979: 2954: 2179: 2154: 1785: 1750: 1726: 1709: 1685: 1660: 881:
Although the term "bidirectional promoter" refers specifically to promoter regions of
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that provide a secure initial binding site for RNA polymerase and for proteins called
4585: 4100: 4057: 4016: 3981: 3950: 3938: 3899: 3858: 3809: 3742: 3711: 3699: 3663: 3628: 3587: 3573: 3538: 3524: 3489: 3460:"Transcription factor binding and modified histones in human bidirectional promoters" 3440: 3381: 3329: 3285: 3234: 3191: 3170:"Bidirectional gene organization: a common architectural feature of the human genome" 3150: 3091: 3033: 2984: 2935: 2886: 2837: 2778: 2737: 2692: 2680: 2646: 2634: 2599: 2548: 2496: 2444: 2398: 2349: 2286: 2227: 2184: 2135: 2094: 2056: 1997: 1962: 1911: 1868: 1825: 1790: 1731: 1710:"Bidirectional gene organization: a common architectural feature of the human genome" 1690: 1641: 1598: 1541: 1500: 1465: 1408: 1185: 1181: 948: 894: 829: 821: 701: 663: 598: 507: 499: 276: 233: 194: 5004: 2130: 2113: 1496: 372:
Some promoters contain one or more upstream promoter element (UP element) subsites (
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Li QJ, Yang SH, Maeda Y, Sladek FM, Sharrocks AD, Martins-Green M (January 2003).
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gene, resulting in the formation of mRNA for that gene alone. Many positive-sense
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are common, as they are in many promoters that lack TATA boxes. In addition, the
705: 680: 562: 3558:"RNA polymerase-promoter interactions: the comings and goings of RNA polymerase" 3509:"A positive-strand RNA virus with three very different subgenomic RNA promoters" 2335: 2202:
Levine M, Tjian R (July 2003). "Transcription regulation and animal diversity".
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Estrem ST, Ross W, Gaal T, Chen ZW, Niu W, Ebright RH, Gourse RL (August 1999).
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Yaniv M (September 2014). "Chromatin remodeling: from transcription to cancer".
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regulatory region is enabled to interact with the promoter region of its target
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Proceedings of the National Academy of Sciences of the United States of America
3607:"Promoter search by Escherichia coli RNA polymerase on a circular DNA template" 3230: 2802:
Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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More recently, one study measured most genes controlled by tandem promoters in
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Grossman SR, Engreitz J, Ray JP, Nguyen TH, Hacohen N, Lander ES (July 2018).
2676: 2584: 2534: 2271: 2252: 1993: 1864: 1125: 1102: 5024: 5009: 4696: 4283: 3680: 1097:). DNA methylation causing silencing in cancer typically occurs at multiple 988: 919: 787: 659: 503: 494: 3794: 3458:
Lin JM, Collins PJ, Trinklein ND, Fu Y, Xi H, Myers RM, Weng Z (June 2007).
2822: 1637: 4883: 4813: 4682: 4020: 3942: 3903: 3862: 3813: 3746: 3703: 3695: 3667: 3542: 3493: 3444: 3385: 3333: 3306:
Shu J, Jelinek J, Chang H, Shen L, Qin T, Chung W, et al. (May 2006).
3289: 3238: 3195: 3154: 3095: 3037: 2988: 2970: 2939: 2921: 2890: 2841: 2782: 2741: 2684: 2638: 2603: 2552: 2500: 2448: 2402: 2353: 2290: 2231: 2139: 2098: 2060: 2001: 1966: 1947: 1915: 1872: 1829: 1775: 1751:"Overlapping promoters and their control in Escherichia coli: the gal case" 1735: 1694: 1676: 1602: 1545: 1536: 1519: 1504: 1469: 1412: 1265: 1169: 1033: 907: 520: 259: 228:
For transcription to take place, the enzyme that synthesizes RNA, known as
163: 4104: 4061: 3985: 3632: 3591: 3308:"Silencing of bidirectional promoters by DNA methylation in tumorigenesis" 3019: 2482: 2188: 1794: 1645: 1113: 484:
Includes the transcription start site (TSS) and elements directly upstream
402:
for -35 sequence T T G A C A 69% 79% 61% 56% 54% 54%
4986: 4915: 4470: 4438: 4341: 2153:
Lagrange T, Kapanidis AN, Tang H, Reinberg D, Ebright RH (January 1998).
805: 796: 470: 399:
for -10 sequence T A T A A T 77% 76% 60% 61% 56% 82%
340: 3844: 2568:"The Why of YY1: Mechanisms of Transcriptional Regulation by Yin Yang 1" 2223: 2170: 362:
The sequence at -35 (the -35 element) has the consensus sequence TTGACA.
4944: 4622: 4316: 4271: 4261: 4256: 4251: 4246: 3738: 3475: 3077: 2439: 2422: 1257: 1025: 967: 886: 791: 381: 305: 270: 240:
that recruit RNA polymerase. These transcription factors have specific
198: 167: 3894: 3877: 1929:
HΓ€kkinen A, Oliveira SM, Neeli-Venkata R, Ribeiro AS (December 2019).
1013:. CpG islands are generally 200 to 2000 base pairs long, have a C:G 339:, the promoter contains two short sequence elements approximately 10 ( 4278: 4096: 3399:
Mahpour A, Scruggs BS, Smiraglia D, Ouchi T, Gelman IH (2018-10-17).
2077:
Smale ST, Kadonaga JT (2003). "The RNA polymerase II core promoter".
1303: 1193: 1037: 1014: 911: 863: 832: 801: 245: 178: 154:. Promoters are located near the transcription start sites of genes, 45: 4034:
Kulozik AE, Bellan-Koch A, Bail S, Kohne E, Kleihauer E (May 1991).
2903: 2872: 2630: 1928: 1428:"The evolution, evolvability and engineering of gene regulatory DNA" 1236:, becoming active in the cell only in response to specific stimuli. 906:
A subgenomic promoter is a promoter added to a virus for a specific
767:
Bidirectional promoters are short (<1 kbp) intergenic regions of
4964: 4954: 4878: 4485: 4475: 2565: 1821: 1252:
When referring to a promoter some authors actually mean promoter +
1129: 1121: 1109: 1098: 1074: 1068: 1062: 1022: 855: 851: 697: 693: 533: 511: 436: 385: 380:
The above promoter sequences are recognized only by RNA polymerase
336: 253: 202: 119: 4633: 3998: 2464: 1199: 1112:
also silence or activate many genes in progression to cancer (see
4700: 4445: 4312: 1161:
Recent evidence also indicates that several genes (including the
1029: 131: 84: 3724: 2367:
Yang C, Bolotin E, Jiang T, Sladek FM, Martinez E (March 2007).
2017:"Analytical kinetic model of native tandem promoters in E. coli" 1979: 1180:
Promoters are important gene regulatory elements used in tuning
1084: 4241: 3059: 1885: 1293: 1211: 844: 840: 825: 780: 777: 454:(BRE), which is recognized by the general transcription factor 394: 369:
The optimal spacing between the -35 and -10 sequences is 17 bp.
258:
The promoter is recognized by RNA polymerase and an associated
182: 71: 776:
co-expressed. Bidirectional promoters are a common feature of
4722: 4502: 4354: 3963: 3349: 3062:"An abundance of bidirectional promoters in the human genome" 2857:"The Mediator complex: a central integrator of transcription" 2152: 2014: 1518:
Civas A, GΓ©nin P, Morin P, Lin R, Hiscott J (February 2006).
1425: 1165: 1049: 974: 926:) and are usually found upstream of the transcription start. 871: 836: 772: 462: 455: 19: 3398: 3216: 3001: 2566:
Verheul TC, van Hijfte L, Perenthaler E, Barakat TS (2020).
1175: 465:(sequence CACGTG), which binds transcription factors in the 4573: 4403: 4334: 4033: 3350:
Wei W, Pelechano V, JΓ€rvelin AI, Steinmetz LM (July 2011).
1264:
would not have an inducible effect. Another example is the
1105:
that are present in the promoters of protein coding genes.
1089:
Generally, in progression to cancer, hundreds of genes are
882: 744: 655: 474: 151: 147: 143: 4172:
RNA and chromatin modification patterns around promoters.
4148: 4001:
American Journal of Respiratory and Critical Care Medicine
3966:
American Journal of Respiratory and Critical Care Medicine
3876:
Chen HY, Shao CJ, Chen FR, Kwan AL, Chen ZP (April 2010).
2708:"YY1 Is a Structural Regulator of Enhancer-Promoter Loops" 2516: 1807: 1260:
is also present. If the lac operator were not present the
1057:
gene. CpG islands also occur frequently in promoters for
732:
have a leading role in the regulation of gene expression.
189:
transcriptional start site in gene promoters (enhancers).
4692: 4208: 2795: 2754: 1850: 1615: 1018: 875: 768: 748: 139: 127: 4074: 3916: 2705: 1561:"Random sequences rapidly evolve into de novo promoters" 783:. About 11% of human genes are bidirectionally paired. 551:
Approximately 250 base pairs upstream of the start site
3555: 2366: 658:
by formation of a chromosome loop. This can initiate
311: 3457: 2317: 4154:
Identifying a Protein Binding Sites on DNA molecule
3352:"Functional consequences of bidirectional promoters" 2460: 2458: 1142: 1032:
nucleotide and this occurs frequently in the linear
146:), or can have a function in and of itself, such as 3305: 2111: 1368: 4166:in defined brain regions of therapeutic interests. 3767: 3108: 2662: 1658: 1517: 1369:LaFleur TL, Hossain A, Salis HM (September 2022). 804:modification. Divergent transcription could shift 177:Promoters control gene expression in bacteria and 3829:"CpG islands and the regulation of transcription" 3556:deHaseth PL, Zupancic ML, Record MT (June 1998). 2952: 2455: 532:General transcription factor binding sites, e.g. 5022: 3875: 2423:"DNA methylation patterns and epigenetic memory" 1069:Methylation of CpG islands stably silences genes 174:recruited to the site, and species of organism. 2658: 2656: 1200:Diseases that may be associated with variations 1116:). Altered microRNA expression occurs through 493:: transcribes genes encoding 18S, 5.8S and 28S 4149:ORegAnno – Open Regulatory Annotation Database 3768:Saxonov S, Berg P, Brutlag DL (January 2006). 3645: 3109:Yang MQ, Koehly LM, Elnitski LL (April 2007). 1748: 1128:in promoters controlling transcription of the 405: 355:The sequence at -10 (the -10 element) has the 4649: 4194: 2467:"Systematic interrogation of human promoters" 2303:: CS1 maint: DOI inactive as of April 2024 ( 2076: 1922: 1085:Promoter CpG hyper/hypo-methylation in cancer 675:bind to DNA sequence motifs on the enhancer. 3869: 3820: 3761: 3167: 3055: 3053: 3051: 3049: 3047: 2995: 2946: 2897: 2854: 2848: 2789: 2748: 2699: 2653: 2616: 2610: 2559: 2311: 2072: 2070: 2008: 1879: 1844: 1742: 1707: 1227: 762: 395:Probability of occurrence of each nucleotide 3598: 3251: 2572:Frontiers in Cell and Developmental Biology 2318:Juven-Gershon T, Kadonaga JT (March 2010). 2201: 1973: 1801: 1558: 1511: 1476: 1239: 998: 847:/BF195580 has been associated with tumors. 571:Specific transcription factor binding sites 4804:Precursor mRNA (pre-mRNA / hnRNA) 4656: 4642: 4201: 4187: 3826: 3506: 2246: 2112:Gershenzon NI, Ioshikhes IP (April 2005). 1701: 975:Diseases associated with aberrant function 820:are more than twice as likely. Many basic 4051: 3893: 3852: 3803: 3793: 3622: 3581: 3532: 3483: 3434: 3424: 3375: 3323: 3279: 3269: 3185: 3144: 3134: 3085: 3044: 3027: 2978: 2929: 2880: 2831: 2821: 2772: 2731: 2593: 2583: 2542: 2512: 2510: 2490: 2438: 2392: 2343: 2280: 2270: 2178: 2129: 2067: 2050: 2040: 1956: 1946: 1784: 1774: 1725: 1684: 1592: 1535: 1459: 1402: 1176:Synthetic promoter design and engineering 1137:methylation of DNA repair genes in cancer 631:Learn how and when to remove this message 3604: 2091:10.1146/annurev.biochem.72.121801.161520 1609: 1172:motifs as potential regulatory signals. 642: 413: 18: 2416: 2414: 2412: 1935:Journal of the Royal Society, Interface 696:(present in about 70% of promoters), a 666:(RNAP II) bound to the promoter at the 16:Region of DNA encouraging transcription 5023: 3301: 3299: 2861:Nature Reviews. Molecular Cell Biology 2507: 1559:Yona AH, Alm EJ, Gore J (April 2018). 1333: 648:Regulation of transcription in mammals 577: 4824:Histone acetylation and deacetylation 4637: 4347:Histone acetylation and deacetylation 4182: 3345: 3343: 1749:Herbert M, Kolb A, Buc H (May 1986). 1482: 1017:content >50%, and have regions of 1005:Regulation of transcription in cancer 700:(present in about 24% of promoters), 4889:Ribosome-nascent chain complex (RNC) 2420: 2409: 2360: 1364: 1362: 1360: 1358: 1356: 786:Bidirectionally paired genes in the 581: 443:TATAAA), which is recognized by the 4663: 3611:The Journal of Biological Chemistry 3296: 2855:Allen BL, Taatjes DJ (March 2015). 1524:The Journal of Biological Chemistry 312:Identification of relative location 166:). Promoters can be about 100–1000 13: 3340: 3252:Liu B, Chen J, Shen B (May 2011). 1247: 556:transcription factor binding sites 487:A binding site for RNA polymerase 14: 5042: 4142: 4125: 3168:Adachi N, Lieber MR (June 2002). 2757:"The Human Transcription Factors" 1708:Adachi N, Lieber MR (June 2002). 1353: 1289:Glossary of gene expression terms 1143:Canonical sequences and wild-type 816:are three times more likely, and 3605:Singer P, Wu CW (October 1987). 3574:10.1128/jb.180.12.3019-3025.1998 3525:10.1128/jvi.74.13.5988-5996.2000 586: 4894:Post-translational modification 4551:Archaeal transcription factor B 4119: 4068: 4027: 3992: 3957: 3910: 3882:International Journal of Cancer 3718: 3674: 3648:Current Opinion in Microbiology 3639: 3549: 3507:Koev G, Miller WA (July 2000). 3500: 3451: 3392: 3245: 3210: 3161: 3102: 2195: 2146: 2105: 1497:10.1016/j.cancergen.2014.03.006 924:Beet necrotic yellow vein virus 213:increase a cell's cancer risk. 3827:Deaton AM, Bird A (May 2011). 1888:Journal of Theoretical Biology 1652: 1552: 1419: 1327: 828:. Forty-five percent of human 1: 4053:10.1182/blood.V77.9.2054.2054 3660:10.1016/s1369-5274(03)00036-5 3624:10.1016/S0021-9258(18)47921-5 3325:10.1158/0008-5472.CAN-05-2629 3187:10.1016/S0092-8674(02)00758-4 2131:10.1093/bioinformatics/bti172 2079:Annual Review of Biochemistry 1727:10.1016/S0092-8674(02)00758-4 1320: 1299:Regulation of gene expression 901: 677:General transcription factors 519:: transcribes genes encoding 502:: transcribes genes encoding 429: 4013:10.1164/ajrccm.160.3.9812024 3978:10.1164/ajrccm.158.6.9804011 3426:10.1371/journal.pone.0205608 3136:10.1371/journal.pcbi.0030072 2042:10.1371/journal.pcbi.1009824 1853:Journal of Molecular Biology 929: 445:general transcription factor 330: 7: 2336:10.1016/j.ydbio.2009.08.009 1334:Sharan R (4 January 2007). 1271: 961: 922:) to over 100 nucleotides ( 771:between the 5' ends of the 606:. The specific problem is: 406:Bidirectional (prokaryotic) 323: 223: 10: 5047: 4227:Transcriptional regulation 4132:San Diego State University 3231:10.1016/j.gene.2005.04.027 3115:PLOS Computational Biology 2774:10.1016/j.cell.2018.01.029 2724:10.1016/j.cell.2017.11.008 2385:10.1016/j.gene.2006.09.029 2021:PLOS Computational Biology 1908:10.1016/j.jtbi.2012.02.015 1585:10.1038/s41467-018-04026-w 1452:10.1038/s41586-022-04506-6 1395:10.1038/s41467-022-32829-5 1002: 946: 942: 318:transcriptional start site 207:TFIIB recognition elements 4997: 4906: 4871: 4845: 4836: 4794: 4768: 4742: 4733: 4671: 4594: 4559: 4533: 4458: 4424:Transcription coregulator 4416: 4393: 4370: 4360:Histone acetyltransferase 4330:Histone methyltransferase 4308:Histone-modifying enzymes 4306: 4299: 4234: 4225: 4160:Pleiades Promoter Project 3935:10.1038/s41587-020-0584-2 3368:10.1016/j.tig.2011.04.002 3271:10.1186/1752-0509-5-S1-S2 2677:10.1038/s41576-019-0128-0 2585:10.3389/fcell.2020.592164 2535:10.1038/s41593-020-0634-6 2272:10.1186/s13073-015-0180-0 1994:10.1021/acssynbio.5b00223 1865:10.1016/j.jmb.2004.11.075 1228:Constitutive vs regulated 1222:Rubinstein-Taybi syndrome 1095:DNA methylation in cancer 1059:functional noncoding RNAs 993:chromosomal translocation 763:Bidirectional (mammalian) 217:transcriptional binding. 4955:sequestration (P-bodies) 2665:Nature Reviews. Genetics 2619:Nature Reviews. Genetics 1240:Tissue-specific promoter 999:CpG islands in promoters 966:Although RNA polymerase 668:transcription start site 349:transcription start site 158:on the DNA (towards the 4933:Gene regulatory network 4525:Internal control region 4170:ENCODE threads Explorer 3833:Genes & Development 3795:10.1073/pnas.0510310103 3562:Journal of Bacteriology 2910:Genes & Development 2823:10.1073/pnas.1804663115 2427:Genes & Development 2421:Bird A (January 2002). 2275:(inactive 2024-04-27). 2159:Genes & Development 1665:Genes & Development 1638:10.1126/science.8248780 1108:Altered expressions of 4938:cis-regulatory element 4156:YouTube tutorial video 3696:10.1002/bies.200800138 3008:Nucleic Acids Research 2922:10.1101/gad.308619.117 1948:10.1098/rsif.2019.0507 1776:10.1073/pnas.83.9.2807 1677:10.1101/gad.13.16.2134 1537:10.1074/jbc.M506812200 1182:synthetically designed 1147:The usage of the term 1118:hyper/hypo-methylation 753:Mediator (coactivator) 713:Cis-regulatory modules 689: 686:Mediator (coactivator) 467:basic helix-loop-helix 419: 115: 4618:Intrinsic termination 4383:DNA methyltransferase 2483:10.1101/gr.236075.118 2324:Developmental Biology 1982:ACS Synthetic Biology 1565:Nature Communications 1375:Nature Communications 1341:. Tel Aviv University 1184:genetic circuits and 1091:silenced or activated 991:through pathological 730:transcription factors 673:transcription factors 646: 538:B recognition element 452:B recognition element 417: 343:) and 35 nucleotides 238:transcription factors 22: 4960:alternative splicing 4950:Post-transcriptional 4776:Transcription factor 4395:Chromatin remodeling 4128:"Expression vectors" 3923:Nature Biotechnology 2971:10.1093/emboj/cdg028 2718:(7): 1573–1588.e28. 1810:Nature Biotechnology 1309:Transcription factor 1279:Activator (genetics) 706:methylated CpG sites 662:(mRNA) synthesis by 613:improve this section 602:to meet Knowledge's 527:and other small RNAs 469:(bHLH) family (e.g. 448:TATA-binding protein 291:, boundary elements/ 4884:Transfer RNA (tRNA) 4352:Histone deacetylase 4342:Histone demethylase 4326:Histone methylation 4089:1995Natur.376..348P 3845:10.1101/gad.2037511 3786:2006PNAS..103.1412S 3617:(29): 14178–14189. 3513:Journal of Virology 3417:2018PLoSO..1305608M 3258:BMC Systems Biology 3127:2007PLSCB...3...72Y 3020:10.1093/nar/gkaa671 2814:2018PNAS..115E7222G 2808:(30): E7222–E7230. 2523:Nature Neuroscience 2224:10.1038/nature01763 2216:2003Natur.424..147L 2171:10.1101/gad.12.1.34 2033:2022PLSCB..18E9824C 1900:2012JThBi.301...83M 1767:1986PNAS...83.2807H 1630:1993Sci...262.1407R 1624:(5138): 1407–1413. 1577:2018NatCo...9.1530Y 1444:2022Natur.603..455V 1387:2022NatCo..13.5159L 1284:Enhancer (genetics) 1114:microRNAs in cancer 818:mitochondrial genes 578:Mammalian promoters 4998:Influential people 4977:Post-translational 4796:Post-transcription 3739:10.1002/bies.20734 3476:10.1101/gr.5623407 3356:Trends in Genetics 3078:10.1101/gr.1982804 2440:10.1101/gad.947102 1207:Examples include: 1186:metabolic networks 1149:canonical sequence 854:, an abundance of 814:Chaperone proteins 690: 517:RNA polymerase III 508:small nuclear RNAs 441:consensus sequence 420: 374:consensus sequence 357:consensus sequence 116: 5018: 5017: 4902: 4901: 4832: 4831: 4708:Special transfers 4631: 4630: 4586:RNA polymerase II 4454: 4453: 4412: 4411: 4083:(6538): 348–351. 3929:(12): 1466–1475. 3895:10.1002/ijc.24772 3839:(10): 1010–1022. 3568:(12): 3019–3025. 3519:(13): 5988–5996. 3318:(10): 5077–5084. 3014:(17): 9550–9570. 2210:(6945): 147–151. 1988:(12): 1331–1341. 1941:(161): 20190507. 1671:(16): 2134–2147. 1438:(7901): 455–463. 1079:5-methylcytosines 1042:5' β†’ 3' direction 1028:is followed by a 949:Promoter activity 895:RNA polymerase II 885:-encoding genes, 664:RNA polymerase II 641: 640: 633: 604:quality standards 595:This section may 525:5s ribosomal RNAs 500:RNA polymerase II 277:RNA polymerase II 234:response elements 195:RNA polymerase II 134:bind to initiate 126:is a sequence of 5038: 4843: 4842: 4740: 4739: 4658: 4651: 4644: 4635: 4634: 4508:Response element 4491:Response element 4304: 4303: 4232: 4231: 4203: 4196: 4189: 4180: 4179: 4174:Nature (journal) 4136: 4135: 4123: 4117: 4116: 4097:10.1038/376348a0 4072: 4066: 4065: 4055: 4046:(9): 2054–2058. 4031: 4025: 4024: 3996: 3990: 3989: 3972:(6): 1958–1962. 3961: 3955: 3954: 3914: 3908: 3907: 3897: 3888:(8): 1944–1954. 3873: 3867: 3866: 3856: 3824: 3818: 3817: 3807: 3797: 3780:(5): 1412–1417. 3765: 3759: 3758: 3722: 3716: 3715: 3678: 3672: 3671: 3643: 3637: 3636: 3626: 3602: 3596: 3595: 3585: 3553: 3547: 3546: 3536: 3504: 3498: 3497: 3487: 3455: 3449: 3448: 3438: 3428: 3411:(10): e0205608. 3396: 3390: 3389: 3379: 3347: 3338: 3337: 3327: 3303: 3294: 3293: 3283: 3273: 3249: 3243: 3242: 3214: 3208: 3207: 3189: 3165: 3159: 3158: 3148: 3138: 3106: 3100: 3099: 3089: 3057: 3042: 3041: 3031: 2999: 2993: 2992: 2982: 2959:The EMBO Journal 2950: 2944: 2943: 2933: 2901: 2895: 2894: 2884: 2852: 2846: 2845: 2835: 2825: 2793: 2787: 2786: 2776: 2752: 2746: 2745: 2735: 2703: 2697: 2696: 2660: 2651: 2650: 2614: 2608: 2607: 2597: 2587: 2563: 2557: 2556: 2546: 2514: 2505: 2504: 2494: 2462: 2453: 2452: 2442: 2418: 2407: 2406: 2396: 2364: 2358: 2357: 2347: 2315: 2309: 2308: 2302: 2294: 2284: 2274: 2250: 2244: 2243: 2199: 2193: 2192: 2182: 2150: 2144: 2143: 2133: 2124:(8): 1295–1300. 2109: 2103: 2102: 2074: 2065: 2064: 2054: 2044: 2012: 2006: 2005: 1977: 1971: 1970: 1960: 1950: 1926: 1920: 1919: 1883: 1877: 1876: 1848: 1842: 1841: 1805: 1799: 1798: 1788: 1778: 1761:(9): 2807–2811. 1746: 1740: 1739: 1729: 1705: 1699: 1698: 1688: 1656: 1650: 1649: 1613: 1607: 1606: 1596: 1556: 1550: 1549: 1539: 1515: 1509: 1508: 1480: 1474: 1473: 1463: 1423: 1417: 1416: 1406: 1366: 1351: 1350: 1348: 1346: 1340: 1331: 1314:Promoter bashing 1217:Beta thalassemia 982:pharmacogenomics 916:subgenomic mRNAs 636: 629: 625: 622: 616: 590: 589: 582: 491:RNA polymerase I 279:to the promoter. 264:DNA binding site 91: 78: 65: 52: 39: 26: 5046: 5045: 5041: 5040: 5039: 5037: 5036: 5035: 5031:Gene expression 5021: 5020: 5019: 5014: 4993: 4928:Transcriptional 4898: 4867: 4828: 4819:Polyadenylation 4790: 4764: 4729: 4723:Proteinβ†’Protein 4674: 4667: 4665:Gene expression 4662: 4632: 4627: 4602: 4596: 4590: 4555: 4529: 4450: 4408: 4389: 4372:DNA methylation 4366: 4310: 4295: 4221: 4207: 4164:gene expression 4145: 4140: 4139: 4124: 4120: 4073: 4069: 4032: 4028: 3997: 3993: 3962: 3958: 3915: 3911: 3874: 3870: 3825: 3821: 3766: 3762: 3723: 3719: 3679: 3675: 3644: 3640: 3603: 3599: 3554: 3550: 3505: 3501: 3464:Genome Research 3456: 3452: 3397: 3393: 3348: 3341: 3312:Cancer Research 3304: 3297: 3264:(Suppl 1): S2. 3250: 3246: 3215: 3211: 3166: 3162: 3107: 3103: 3066:Genome Research 3058: 3045: 3000: 2996: 2951: 2947: 2902: 2898: 2873:10.1038/nrm3951 2853: 2849: 2794: 2790: 2753: 2749: 2704: 2700: 2661: 2654: 2631:10.1038/nrg3207 2615: 2611: 2564: 2560: 2515: 2508: 2471:Genome Research 2463: 2456: 2419: 2410: 2365: 2361: 2316: 2312: 2296: 2295: 2259:Genome Medicine 2251: 2247: 2200: 2196: 2151: 2147: 2110: 2106: 2075: 2068: 2027:(1): e1009824. 2013: 2009: 1978: 1974: 1927: 1923: 1884: 1880: 1849: 1845: 1806: 1802: 1747: 1743: 1706: 1702: 1657: 1653: 1614: 1610: 1557: 1553: 1516: 1512: 1485:Cancer Genetics 1481: 1477: 1424: 1420: 1367: 1354: 1344: 1342: 1338: 1332: 1328: 1323: 1318: 1274: 1250: 1248:Use of the term 1242: 1230: 1202: 1178: 1145: 1087: 1071: 1007: 1001: 977: 964: 951: 945: 936:gene prediction 932: 904: 891:non-coding RNAs 843:/BC040563, and 765: 702:initiator (Inr) 681:phosphorylation 637: 626: 620: 617: 610: 591: 587: 580: 563:Distal promoter 432: 408: 403: 400: 397: 392: 333: 326: 314: 226: 110: 105: 89: 87: 76: 74: 63: 61: 50: 48: 37: 35: 24: 17: 12: 11: 5: 5044: 5034: 5033: 5016: 5015: 5013: 5012: 5007: 5005:FranΓ§ois Jacob 5001: 4999: 4995: 4994: 4992: 4991: 4990: 4989: 4984: 4974: 4969: 4968: 4967: 4962: 4957: 4947: 4942: 4941: 4940: 4935: 4925: 4924: 4923: 4912: 4910: 4904: 4903: 4900: 4899: 4897: 4896: 4891: 4886: 4881: 4875: 4873: 4869: 4868: 4866: 4865: 4860: 4855: 4849: 4847: 4840: 4834: 4833: 4830: 4829: 4827: 4826: 4821: 4816: 4811: 4806: 4800: 4798: 4792: 4791: 4789: 4788: 4783: 4781:RNA polymerase 4778: 4772: 4770: 4766: 4765: 4763: 4762: 4757: 4752: 4746: 4744: 4737: 4731: 4730: 4728: 4727: 4726: 4725: 4720: 4715: 4705: 4704: 4703: 4685: 4679: 4677: 4669: 4668: 4661: 4660: 4653: 4646: 4638: 4629: 4628: 4626: 4625: 4620: 4615: 4609: 4607: 4592: 4591: 4589: 4588: 4582:RNA polymerase 4576: 4570:RNA polymerase 4563: 4561: 4557: 4556: 4554: 4553: 4548: 4543: 4537: 4535: 4531: 4530: 4528: 4527: 4522: 4517: 4512: 4511: 4510: 4505: 4495: 4494: 4493: 4488: 4483: 4478: 4473: 4462: 4460: 4456: 4455: 4452: 4451: 4449: 4448: 4443: 4442: 4441: 4436: 4431: 4420: 4418: 4414: 4413: 4410: 4409: 4407: 4406: 4400: 4398: 4391: 4390: 4388: 4387: 4386: 4385: 4377: 4375: 4368: 4367: 4365: 4364: 4363: 4362: 4357: 4344: 4339: 4338: 4337: 4322: 4320: 4301: 4297: 4296: 4294: 4293: 4292: 4291: 4286: 4276: 4275: 4274: 4269: 4264: 4259: 4254: 4249: 4238: 4236: 4229: 4223: 4222: 4206: 4205: 4198: 4191: 4183: 4177: 4176: 4167: 4157: 4151: 4144: 4143:External links 4141: 4138: 4137: 4118: 4067: 4026: 4007:(3): 919–922. 3991: 3956: 3909: 3868: 3819: 3760: 3733:(4): 314–327. 3717: 3690:(6): 629–641. 3673: 3638: 3597: 3548: 3499: 3470:(6): 818–827. 3450: 3391: 3362:(7): 267–276. 3339: 3295: 3244: 3225:(2): 169–176. 3209: 3180:(7): 807–809. 3160: 3101: 3043: 2994: 2965:(2): 281–291. 2945: 2896: 2867:(3): 155–166. 2847: 2788: 2767:(4): 650–665. 2747: 2698: 2671:(8): 437–455. 2652: 2625:(9): 613–626. 2609: 2558: 2529:(6): 707–717. 2506: 2477:(2): 171–183. 2454: 2408: 2359: 2330:(2): 225–229. 2310: 2245: 2194: 2145: 2118:Bioinformatics 2104: 2066: 2007: 1972: 1921: 1878: 1859:(2): 399–409. 1843: 1822:10.1038/nbt988 1816:(7): 911–917. 1800: 1741: 1720:(7): 807–809. 1700: 1651: 1608: 1551: 1530:(8): 4856–66. 1510: 1475: 1418: 1352: 1325: 1324: 1322: 1319: 1317: 1316: 1311: 1306: 1301: 1296: 1291: 1286: 1281: 1275: 1273: 1270: 1249: 1246: 1241: 1238: 1229: 1226: 1225: 1224: 1219: 1214: 1201: 1198: 1177: 1174: 1163:proto-oncogene 1144: 1141: 1086: 1083: 1070: 1067: 1003:Main article: 1000: 997: 989:chimeric genes 980:discipline of 976: 973: 963: 960: 944: 941: 931: 928: 914:produce these 903: 900: 764: 761: 639: 638: 621:September 2021 594: 592: 585: 579: 576: 575: 574: 573: 572: 569: 560: 559: 558: 552: 546: 545: 544: 541: 530: 529: 528: 514: 497: 495:ribosomal RNAs 485: 431: 428: 407: 404: 401: 398: 396: 393: 390: 378: 377: 370: 367: 363: 360: 332: 329: 325: 322: 313: 310: 281: 280: 273: 267: 256: 230:RNA polymerase 225: 222: 172:RNA polymerase 88: 75: 62: 49: 36: 33:RNA Polymerase 23: 15: 9: 6: 4: 3: 2: 5043: 5032: 5029: 5028: 5026: 5011: 5010:Jacques Monod 5008: 5006: 5003: 5002: 5000: 4996: 4988: 4985: 4983: 4980: 4979: 4978: 4975: 4973: 4972:Translational 4970: 4966: 4963: 4961: 4958: 4956: 4953: 4952: 4951: 4948: 4946: 4943: 4939: 4936: 4934: 4931: 4930: 4929: 4926: 4922: 4919: 4918: 4917: 4914: 4913: 4911: 4909: 4905: 4895: 4892: 4890: 4887: 4885: 4882: 4880: 4877: 4876: 4874: 4870: 4864: 4861: 4859: 4856: 4854: 4851: 4850: 4848: 4844: 4841: 4839: 4835: 4825: 4822: 4820: 4817: 4815: 4812: 4810: 4807: 4805: 4802: 4801: 4799: 4797: 4793: 4787: 4784: 4782: 4779: 4777: 4774: 4773: 4771: 4767: 4761: 4758: 4756: 4753: 4751: 4748: 4747: 4745: 4741: 4738: 4736: 4735:Transcription 4732: 4724: 4721: 4719: 4716: 4714: 4711: 4710: 4709: 4706: 4702: 4698: 4694: 4691: 4690: 4689: 4688:Central dogma 4686: 4684: 4681: 4680: 4678: 4676: 4670: 4666: 4659: 4654: 4652: 4647: 4645: 4640: 4639: 4636: 4624: 4621: 4619: 4616: 4614: 4611: 4610: 4608: 4605: 4600: 4593: 4587: 4583: 4580: 4577: 4575: 4571: 4568: 4565: 4564: 4562: 4558: 4552: 4549: 4547: 4544: 4542: 4539: 4538: 4536: 4532: 4526: 4523: 4521: 4518: 4516: 4513: 4509: 4506: 4504: 4501: 4500: 4499: 4496: 4492: 4489: 4487: 4484: 4482: 4479: 4477: 4474: 4472: 4469: 4468: 4467: 4464: 4463: 4461: 4457: 4447: 4444: 4440: 4437: 4435: 4432: 4430: 4427: 4426: 4425: 4422: 4421: 4419: 4415: 4405: 4402: 4401: 4399: 4396: 4392: 4384: 4381: 4380: 4379: 4378: 4376: 4373: 4369: 4361: 4358: 4356: 4353: 4350: 4349: 4348: 4345: 4343: 4340: 4336: 4333: 4332: 4331: 4327: 4324: 4323: 4321: 4318: 4314: 4309: 4305: 4302: 4298: 4290: 4289:trp repressor 4287: 4285: 4284:lac repressor 4282: 4281: 4280: 4277: 4273: 4270: 4268: 4265: 4263: 4260: 4258: 4255: 4253: 4250: 4248: 4245: 4244: 4243: 4240: 4239: 4237: 4233: 4230: 4228: 4224: 4219: 4215: 4211: 4210:Transcription 4204: 4199: 4197: 4192: 4190: 4185: 4184: 4181: 4175: 4171: 4168: 4165: 4161: 4158: 4155: 4152: 4150: 4147: 4146: 4133: 4129: 4122: 4114: 4110: 4106: 4102: 4098: 4094: 4090: 4086: 4082: 4078: 4071: 4063: 4059: 4054: 4049: 4045: 4041: 4037: 4030: 4022: 4018: 4014: 4010: 4006: 4002: 3995: 3987: 3983: 3979: 3975: 3971: 3967: 3960: 3952: 3948: 3944: 3940: 3936: 3932: 3928: 3924: 3920: 3913: 3905: 3901: 3896: 3891: 3887: 3883: 3879: 3872: 3864: 3860: 3855: 3850: 3846: 3842: 3838: 3834: 3830: 3823: 3815: 3811: 3806: 3801: 3796: 3791: 3787: 3783: 3779: 3775: 3771: 3764: 3756: 3752: 3748: 3744: 3740: 3736: 3732: 3728: 3721: 3713: 3709: 3705: 3701: 3697: 3693: 3689: 3685: 3677: 3669: 3665: 3661: 3657: 3654:(2): 93–100. 3653: 3649: 3642: 3634: 3630: 3625: 3620: 3616: 3612: 3608: 3601: 3593: 3589: 3584: 3579: 3575: 3571: 3567: 3563: 3559: 3552: 3544: 3540: 3535: 3530: 3526: 3522: 3518: 3514: 3510: 3503: 3495: 3491: 3486: 3481: 3477: 3473: 3469: 3465: 3461: 3454: 3446: 3442: 3437: 3432: 3427: 3422: 3418: 3414: 3410: 3406: 3402: 3395: 3387: 3383: 3378: 3373: 3369: 3365: 3361: 3357: 3353: 3346: 3344: 3335: 3331: 3326: 3321: 3317: 3313: 3309: 3302: 3300: 3291: 3287: 3282: 3277: 3272: 3267: 3263: 3259: 3255: 3248: 3240: 3236: 3232: 3228: 3224: 3220: 3213: 3205: 3201: 3197: 3193: 3188: 3183: 3179: 3175: 3171: 3164: 3156: 3152: 3147: 3142: 3137: 3132: 3128: 3124: 3120: 3116: 3112: 3105: 3097: 3093: 3088: 3083: 3079: 3075: 3071: 3067: 3063: 3056: 3054: 3052: 3050: 3048: 3039: 3035: 3030: 3025: 3021: 3017: 3013: 3009: 3005: 2998: 2990: 2986: 2981: 2976: 2972: 2968: 2964: 2960: 2956: 2949: 2941: 2937: 2932: 2927: 2923: 2919: 2915: 2911: 2907: 2900: 2892: 2888: 2883: 2878: 2874: 2870: 2866: 2862: 2858: 2851: 2843: 2839: 2834: 2829: 2824: 2819: 2815: 2811: 2807: 2803: 2799: 2792: 2784: 2780: 2775: 2770: 2766: 2762: 2758: 2751: 2743: 2739: 2734: 2729: 2725: 2721: 2717: 2713: 2709: 2702: 2694: 2690: 2686: 2682: 2678: 2674: 2670: 2666: 2659: 2657: 2648: 2644: 2640: 2636: 2632: 2628: 2624: 2620: 2613: 2605: 2601: 2596: 2591: 2586: 2581: 2577: 2573: 2569: 2562: 2554: 2550: 2545: 2540: 2536: 2532: 2528: 2524: 2520: 2513: 2511: 2502: 2498: 2493: 2488: 2484: 2480: 2476: 2472: 2468: 2461: 2459: 2450: 2446: 2441: 2436: 2432: 2428: 2424: 2417: 2415: 2413: 2404: 2400: 2395: 2390: 2386: 2382: 2378: 2374: 2370: 2363: 2355: 2351: 2346: 2341: 2337: 2333: 2329: 2325: 2321: 2314: 2306: 2300: 2292: 2288: 2283: 2278: 2273: 2268: 2264: 2260: 2256: 2249: 2241: 2237: 2233: 2229: 2225: 2221: 2217: 2213: 2209: 2205: 2198: 2190: 2186: 2181: 2176: 2172: 2168: 2164: 2160: 2156: 2149: 2141: 2137: 2132: 2127: 2123: 2119: 2115: 2108: 2100: 2096: 2092: 2088: 2084: 2080: 2073: 2071: 2062: 2058: 2053: 2048: 2043: 2038: 2034: 2030: 2026: 2022: 2018: 2011: 2003: 1999: 1995: 1991: 1987: 1983: 1976: 1968: 1964: 1959: 1954: 1949: 1944: 1940: 1936: 1932: 1925: 1917: 1913: 1909: 1905: 1901: 1897: 1893: 1889: 1882: 1874: 1870: 1866: 1862: 1858: 1854: 1847: 1839: 1835: 1831: 1827: 1823: 1819: 1815: 1811: 1804: 1796: 1792: 1787: 1782: 1777: 1772: 1768: 1764: 1760: 1756: 1752: 1745: 1737: 1733: 1728: 1723: 1719: 1715: 1711: 1704: 1696: 1692: 1687: 1682: 1678: 1674: 1670: 1666: 1662: 1655: 1647: 1643: 1639: 1635: 1631: 1627: 1623: 1619: 1612: 1604: 1600: 1595: 1590: 1586: 1582: 1578: 1574: 1570: 1566: 1562: 1555: 1547: 1543: 1538: 1533: 1529: 1525: 1521: 1514: 1506: 1502: 1498: 1494: 1490: 1486: 1479: 1471: 1467: 1462: 1457: 1453: 1449: 1445: 1441: 1437: 1433: 1429: 1422: 1414: 1410: 1405: 1400: 1396: 1392: 1388: 1384: 1380: 1376: 1372: 1365: 1363: 1361: 1359: 1357: 1337: 1330: 1326: 1315: 1312: 1310: 1307: 1305: 1302: 1300: 1297: 1295: 1292: 1290: 1287: 1285: 1282: 1280: 1277: 1276: 1269: 1267: 1263: 1259: 1255: 1245: 1237: 1235: 1223: 1220: 1218: 1215: 1213: 1210: 1209: 1208: 1205: 1197: 1195: 1191: 1187: 1183: 1173: 1171: 1167: 1164: 1159: 1155: 1152: 1150: 1140: 1138: 1133: 1131: 1127: 1123: 1119: 1115: 1111: 1106: 1104: 1100: 1096: 1092: 1082: 1080: 1076: 1066: 1064: 1060: 1056: 1052: 1051: 1045: 1043: 1039: 1035: 1031: 1027: 1024: 1020: 1016: 1012: 1006: 996: 994: 990: 985: 983: 972: 969: 959: 955: 950: 940: 937: 927: 925: 921: 920:Sindbis virus 917: 913: 909: 899: 896: 892: 888: 884: 879: 877: 873: 869: 865: 861: 857: 853: 848: 846: 842: 838: 834: 831: 827: 823: 819: 815: 809: 807: 803: 798: 793: 789: 788:Gene Ontology 784: 782: 779: 774: 770: 760: 756: 754: 750: 746: 740: 737: 733: 731: 727: 723: 719: 714: 710: 707: 703: 699: 695: 687: 682: 678: 674: 669: 665: 661: 660:messenger RNA 657: 653: 649: 645: 635: 632: 624: 614: 609: 605: 601: 600: 593: 584: 583: 570: 567: 566: 564: 561: 557: 553: 550: 549: 547: 542: 539: 535: 531: 526: 522: 518: 515: 513: 509: 505: 504:messenger RNA 501: 498: 496: 492: 489: 488: 486: 483: 482: 480: 479: 478: 476: 472: 468: 464: 459: 457: 453: 450:(TBP); and a 449: 446: 442: 438: 427: 425: 416: 412: 389: 387: 383: 375: 371: 368: 364: 361: 358: 354: 353: 352: 350: 346: 342: 338: 328: 321: 319: 309: 307: 303: 302: 296: 294: 290: 286: 278: 274: 272: 268: 265: 261: 257: 255: 251: 250: 249: 247: 243: 239: 235: 231: 221: 218: 214: 210: 208: 204: 200: 196: 190: 186: 184: 180: 175: 173: 169: 165: 161: 157: 153: 149: 145: 141: 137: 136:transcription 133: 129: 125: 121: 113: 108: 103: 100:: lacY, 99: 96:: lacZ, 95: 86: 82: 73: 69: 60: 56: 47: 43: 34: 30: 21: 4987:irreversible 4872:Key elements 4785: 4769:Key elements 4683:Genetic code 4673:Introduction 4465: 4131: 4121: 4080: 4076: 4070: 4043: 4039: 4029: 4004: 4000: 3994: 3969: 3965: 3959: 3926: 3922: 3912: 3885: 3881: 3871: 3836: 3832: 3822: 3777: 3773: 3763: 3730: 3726: 3720: 3687: 3683: 3676: 3651: 3647: 3641: 3614: 3610: 3600: 3565: 3561: 3551: 3516: 3512: 3502: 3467: 3463: 3453: 3408: 3404: 3394: 3359: 3355: 3315: 3311: 3261: 3257: 3247: 3222: 3218: 3212: 3177: 3173: 3163: 3118: 3114: 3104: 3072:(1): 62–66. 3069: 3065: 3011: 3007: 2997: 2962: 2958: 2948: 2916:(1): 42–57. 2913: 2909: 2899: 2864: 2860: 2850: 2805: 2801: 2791: 2764: 2760: 2750: 2715: 2711: 2701: 2668: 2664: 2622: 2618: 2612: 2575: 2571: 2561: 2526: 2522: 2474: 2470: 2430: 2426: 2379:(1): 52–65. 2376: 2372: 2362: 2327: 2323: 2313: 2299:cite journal 2262: 2258: 2248: 2207: 2203: 2197: 2165:(1): 34–44. 2162: 2158: 2148: 2121: 2117: 2107: 2082: 2078: 2024: 2020: 2010: 1985: 1981: 1975: 1938: 1934: 1924: 1891: 1887: 1881: 1856: 1852: 1846: 1813: 1809: 1803: 1758: 1754: 1744: 1717: 1713: 1703: 1668: 1664: 1654: 1621: 1617: 1611: 1568: 1564: 1554: 1527: 1523: 1513: 1491:(9): 352–7. 1488: 1484: 1478: 1435: 1431: 1421: 1378: 1374: 1343:. Retrieved 1329: 1266:Tac-Promoter 1251: 1243: 1231: 1206: 1203: 1192:. Automated 1179: 1170:G-quadruplex 1160: 1156: 1153: 1146: 1134: 1107: 1088: 1072: 1054: 1048: 1046: 1008: 986: 978: 965: 956: 952: 933: 908:heterologous 905: 880: 860:CGCG element 849: 822:housekeeping 810: 785: 766: 757: 741: 734: 711: 691: 650:. An active 647: 627: 618: 611:Please help 607: 596: 521:transfer RNA 506:and certain 460: 433: 423: 421: 409: 379: 344: 334: 327: 315: 306:lac promoter 300: 297: 282: 260:sigma factor 227: 219: 215: 211: 191: 187: 176: 164:sense strand 138:of a single 123: 117: 111: 106: 104:: lacA. 101: 97: 93: 80: 67: 58: 54: 41: 28: 4838:Translation 4675:to genetics 4595:Termination 4471:Pribnow box 4439:Corepressor 4434:Coactivator 4235:prokaryotic 2433:(1): 6–21. 2085:: 449–479. 1571:(1): 1530. 1381:(1): 5159. 1345:30 December 1126:CpG islands 1103:CpG islands 912:RNA viruses 864:CCAAT boxes 856:CpG islands 839:/CD558500, 806:nucleosomes 797:methylation 694:CpG islands 615:if you can. 471:BMAL1-Clock 384:containing 341:Pribnow Box 199:CpG islands 4982:reversible 4945:lac operon 4921:imprinting 4916:Epigenetic 4908:Regulation 4863:Eukaryotic 4809:5' capping 4760:Eukaryotic 4623:Rho factor 4613:Terminator 4604:eukaryotic 4579:eukaryotic 4560:Elongation 4546:Eukaryotic 4534:Initiation 4317:nucleosome 4300:eukaryotic 4272:gal operon 4267:ara operon 4262:Gua Operon 4257:gab operon 4252:trp operon 4247:lac operon 4218:Eukaryotic 3121:(4): e72. 2578:: 592164. 1321:References 1258:lac operon 1194:algorithms 1190:expression 1040:along its 1026:nucleotide 1011:CpG island 968:holoenzyme 947:See also: 902:Subgenomic 887:luciferase 852:TATA boxes 792:Microarray 726:insulators 430:Eukaryotic 382:holoenzyme 293:insulators 271:eukaryotes 179:eukaryotes 168:base pairs 4853:Bacterial 4750:Bacterial 4599:bacterial 4567:bacterial 4541:Bacterial 4515:Insulator 4459:Promotion 4429:Activator 4279:Repressor 4214:Bacterial 4126:Maloy S. 3951:220506228 3727:BioEssays 3712:205469320 3684:BioEssays 2693:152283312 2647:205485256 2265:(1): 66. 1894:: 83–94. 1304:Repressor 1234:regulated 1130:microRNAs 1122:CpG sites 1110:microRNAs 1099:CpG sites 1075:CpG sites 1063:microRNAs 1015:base pair 930:Detection 833:oncogenes 802:chromatin 778:mammalian 736:Enhancers 722:silencers 718:enhancers 554:Specific 347:from the 331:Bacterial 289:silencers 285:enhancers 246:repressor 242:activator 160:5' region 130:to which 46:Repressor 5025:Category 4965:microRNA 4879:Ribosome 4858:Archaeal 4814:Splicing 4786:Promoter 4755:Archaeal 4699: β†’ 4695: β†’ 4520:Silencer 4498:Enhancer 4486:CAAT box 4476:TATA box 4466:Promoter 4021:10471619 3943:32661437 3904:19626585 3863:21576262 3814:16432200 3747:18348191 3704:19382224 3668:12732296 3543:10846080 3494:17568000 3445:30332484 3405:PLOS ONE 3386:21601935 3334:16707430 3290:21689477 3239:15944140 3196:12110178 3155:17447839 3096:14707170 3038:32810208 2989:12514134 2940:29378788 2891:25693131 2842:29987030 2783:29425488 2742:29224777 2685:31086298 2639:22868264 2604:33102493 2553:32451484 2501:30622120 2449:11782440 2403:17123746 2354:19682982 2291:26221185 2232:12853946 2140:15572469 2099:12651739 2061:35100257 2002:27346626 1967:31822223 1916:22370562 1873:15670592 1830:15229555 1736:12110178 1695:10465790 1603:29670097 1546:16380379 1505:24825771 1470:35264797 1413:36056029 1272:See also 1254:operator 1077:to form 1061:such as 1034:sequence 1023:cytosine 1021:where a 962:Location 698:TATA box 652:enhancer 597:require 534:TATA box 512:microRNA 437:TATA box 386:sigma-70 345:upstream 337:bacteria 324:Elements 254:bacteria 224:Overview 203:TATA box 156:upstream 132:proteins 124:promoter 120:genetics 72:Operator 59:Promoter 4718:RNAβ†’DNA 4713:RNAβ†’RNA 4701:Protein 4446:Inducer 4313:histone 4113:4254507 4105:7630403 4085:Bibcode 4062:2018842 3986:9847292 3854:3093116 3805:1345710 3782:Bibcode 3633:3308887 3592:9620948 3485:1891341 3436:6192621 3413:Bibcode 3377:3123404 3281:3121118 3204:8556921 3146:1853124 3123:Bibcode 3029:7515708 2931:5828394 2882:4963239 2833:6065035 2810:Bibcode 2733:5785279 2595:7554316 2544:7558717 2492:6360817 2394:1955227 2345:2830304 2282:4517488 2240:4373712 2212:Bibcode 2189:9420329 2052:8830795 2029:Bibcode 1958:6936044 1896:Bibcode 1838:3546895 1795:3010319 1763:Bibcode 1646:8248780 1626:Bibcode 1618:Science 1594:5906472 1573:Bibcode 1461:8934302 1440:Bibcode 1404:9440211 1383:Bibcode 1168:) have 1101:in the 1030:guanine 943:Binding 870:NRF-1, 830:somatic 781:genomes 599:cleanup 424:E. coli 359:TATAAT. 301:E. coli 266:nearby. 162:of the 85:Lactose 83::  70::  57::  44::  31::  4242:Operon 4111:  4103:  4077:Nature 4060:  4019:  3984:  3949:  3941:  3902:  3861:  3851:  3812:  3802:  3755:678541 3753:  3745:  3710:  3702:  3666:  3631:  3590:  3583:107799 3580:  3541:  3534:112095 3531:  3492:  3482:  3443:  3433:  3384:  3374:  3332:  3288:  3278:  3237:  3202:  3194:  3153:  3143:  3094:  3087:314279 3084:  3036:  3026:  2987:  2980:140103 2977:  2938:  2928:  2889:  2879:  2840:  2830:  2781:  2740:  2730:  2691:  2683:  2645:  2637:  2602:  2592:  2551:  2541:  2499:  2489:  2447:  2401:  2391:  2352:  2342:  2289:  2279:  2238:  2230:  2204:Nature 2187:  2180:316406 2177:  2138:  2097:  2059:  2049:  2000:  1965:  1955:  1914:  1871:  1836:  1828:  1793:  1786:323395 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Index


RNA Polymerase
Repressor
Promoter
Operator
Lactose
genetics
DNA
proteins
transcription
RNA
mRNA
tRNA
rRNA
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5' region
sense strand
base pairs
RNA polymerase
eukaryotes
enzyme
RNA polymerase II
CpG islands
TATA box
TFIIB recognition elements
RNA polymerase
response elements
transcription factors
activator
repressor

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