# Promoter TATA Box: Definition, Role, and Mechanisms

## Introduction to the Promoter TATA Box

The TATA box is a conserved DNA sequence motif located within the core promoter region of many eukaryotic genes. It is defined as a cis-acting regulatory element that serves as the primary binding site for the general [transcription factor](/knowledge/molecular-biology/transcription-factor) TFIID, specifically through its TATA-binding protein (TBP) subunit. The TATA box is one of the most well-characterized core promoter elements, and its discovery in 1979 by David Hogness and colleagues in *Drosophila* genes marked a turning point in understanding how RNA polymerase II initiates transcription.

The name "TATA box" derives from its consensus sequence, which is rich in thymine and adenine residues. In most organisms, the canonical sequence is TATAWAWR, where W represents either adenine or thymine, and R represents either adenine or guanine. The most common form is TATAAAAG, though variations are tolerated to different degrees depending on the organism and the specific gene.

Functionally, the TATA box is not merely a passive sequence motif; it is the nucleation point for the assembly of the preinitiation complex (PIC). When TBP binds to the TATA box, it induces a sharp bend in the DNA helix, creating a platform for the recruitment of additional general [transcription factors](/knowledge/molecular-biology/transcription-factor) (TFIIA, TFIIB, TFIIE, TFIIF, TFIIH) and ultimately RNA polymerase II. This binding event is often described as the rate-limiting step in [transcription initiation](/knowledge/molecular-biology/transcription-initiation) for TATA-containing promoters.

It is essential to distinguish the TATA box from the broader [Promoter Region](/knowledge/molecular-biology/promoter-region). The promoter region encompasses all DNA sequences required for accurate and efficient [transcription initiation](/knowledge/molecular-biology/transcription-initiation), including the core promoter (where the TATA box resides) and proximal promoter elements such as GC boxes and CAAT boxes. The TATA box is thus a component of the core promoter, not the promoter itself.

## Location and Consensus Sequence of the TATA Box

The TATA box is located approximately 25 to 30 base pairs upstream of the transcription start site (TSS) in higher eukaryotes. In *Saccharomyces cerevisiae* (budding yeast), the distance is slightly more variable, typically ranging from 40 to 120 base pairs upstream of the TSS. This positional variability reflects differences in how yeast and metazoans assemble the PIC, but the fundamental role of the TATA box remains conserved.

The consensus sequence is most commonly written as TATAWAWR, where:

- W = A or T
- R = A or G

The most frequently observed sequence in human promoters is TATAAAAG, followed by TATATAAA and TATAAATA. The first four nucleotides (TATA) are the most highly conserved and are critical for TBP binding. The fifth position (W) is almost always A or T, and the sixth position (A) is invariant in nearly all functional TATA boxes. The seventh position (W) and eighth position (R) show more variability but still contribute to binding affinity.

The TATA box is almost always flanked by GC-rich sequences, which influence DNA bending and TBP binding kinetics. The flanking sequences are not part of the consensus but can modulate the affinity of TBP for the TATA box by up to 10-fold. For example, a TATA box preceded by a run of guanines binds TBP more weakly than one preceded by adenines, due to differences in DNA flexibility.

A common misconception is that the TATA box sequence is identical across all genes. In reality, the TATA box is a degenerate motif, and many functional variants exist. Some genes contain a "TATA-less" promoter that nevertheless has a functional TBP-binding site with a sequence that deviates significantly from the consensus. This degeneracy is biologically important because it allows for differential regulation of genes: promoters with high-affinity TATA boxes (e.g., TATAAAAG) tend to be constitutively active or strongly inducible, while those with weak TATA boxes are more likely to be regulated by developmental or tissue-specific signals.

## Role of the TATA Box in Transcription Initiation

### Binding of TATA-Binding Protein (TBP)

The TATA box is recognized by the TATA-binding protein (TBP), a 38 kDa protein in humans that is a subunit of the general transcription factor TFIID. TBP binds to the minor groove of the DNA helix at the TATA box, which is unusual because most DNA-binding proteins interact with the major groove. The minor groove binding is facilitated by the narrow width of the AT-rich sequence, which allows the protein's phenylalanine residues to intercalate between base pairs.

Upon binding, TBP induces a dramatic conformational change in the DNA: the helix is bent by approximately 80 degrees, and the minor groove is widened. This bending is critical for the subsequent recruitment of other transcription factors. The bend creates a surface that is recognized by TFIIB, which then recruits RNA polymerase II and TFIIF. The TBP–TATA box interaction is also stabilized by TFIIA, which binds to the upstream side of TBP and protects it from proteolytic degradation.

The affinity of TBP for the TATA box is high, with a dissociation constant (Kd) in the nanomolar range (typically 1–10 nM for the consensus sequence). However, the binding is not static; TBP undergoes conformational changes upon binding, and the residence time on the DNA is influenced by the specific TATA box sequence and the presence of other factors. For example, the presence of TFIIA increases the half-life of the TBP–DNA complex from approximately 15 minutes to over 60 minutes in vitro.

### Formation of the Preinitiation Complex

The assembly of the preinitiation complex (PIC) proceeds in a defined order, although the exact sequence of events can vary depending on the promoter and the cellular context. The canonical pathway is as follows:

1. **TBP binding**: TBP (as part of TFIID) binds to the TATA box, bending the DNA and creating a platform for further assembly.
2. **TFIIA and TFIIB recruitment**: TFIIA binds to the upstream side of TBP, while TFIIB binds to the downstream side. TFIIB also makes sequence-specific contacts with the DNA just upstream of the TSS, helping to position RNA polymerase II correctly.
3. **RNA polymerase II and TFIIF loading**: TFIIF binds to RNA polymerase II and helps recruit the polymerase to the TFIIB–TBP–DNA complex. TFIIF also reduces the non-specific DNA-binding affinity of the polymerase, ensuring that it only initiates at the correct site.
4. **TFIIE and TFIIH association**: TFIIE recruits TFIIH, a multi-subunit complex with helicase and kinase activities. TFIIH unwinds the DNA at the TSS to form the open complex and phosphorylates the C-terminal domain (CTD) of RNA polymerase II at serine 5, which is required for promoter escape.
5. **Promoter clearance**: RNA polymerase II synthesizes a short RNA transcript (approximately 20–30 nucleotides) and then escapes the promoter, leaving the PIC behind. TFIIH remains associated with the polymerase during early elongation.

The TATA box is thus the nucleation site for the entire PIC. Without a functional TATA box, the assembly of the PIC is severely impaired, and transcription initiation is reduced by 10- to 100-fold in vitro. However, as discussed below, some promoters can bypass the TATA box requirement by using alternative core promoter elements.

## TATA Box vs. TATA-Less Promoters

Not all promoters contain a TATA box. In fact, it is estimated that only 10–20% of human promoters are TATA-dependent. The majority of promoters are TATA-less and rely on other core promoter elements to direct transcription initiation. Understanding the differences between these two classes is essential for interpreting gene expression data and designing experiments.

The table below summarizes the key differences between TATA-containing and TATA-less promoters:

| Feature | TATA-Containing Promoters | TATA-Less Promoters |
|---------|---------------------------|---------------------|
| Core promoter elements | TATA box, sometimes Inr | Inr, DPE, BRE, MTE, CpG islands |
| TBP binding | Direct, high affinity | Indirect, via tethering factors (e.g., NC2, TFTC) |
| Position of TSS | Focused, single or few start sites | Dispersed, multiple start sites |
| Gene classes | Highly regulated, tissue-specific, stress-responsive | Housekeeping, developmental, broadly expressed |
| Chromatin context | Often nucleosome-depleted but regulated | Often CpG-rich, constitutively open |
| Evolutionary conservation | High | Lower, more variable |

The initiator (Inr) element is a sequence that overlaps the TSS and has the consensus YYANWYY (where Y = C or T, W = A or T). The Inr is recognized by TFIID through its TAF1 and TAF2 subunits, and it can function independently of the TATA box. The downstream promoter element (DPE) is located approximately 28–32 nucleotides downstream of the TSS and is recognized by TAF6 and TAF9. DPE-dependent promoters often lack a TATA box and are common in *Drosophila*.

In vertebrates, many TATA-less promoters are associated with CpG islands, which are GC-rich regions that are often unmethylated and associated with active chromatin. These promoters typically have dispersed TSSs and are constitutively active. The transcription factor Sp1 binds to GC boxes within these promoters and helps recruit TFIID in a TATA-independent manner.

It is important to note that the distinction between TATA-containing and TATA-less promoters is not absolute. Some promoters contain both a TATA box and an Inr, while others contain weak TATA boxes that are not essential for basal transcription. The presence or absence of a TATA box is therefore a useful but not definitive predictor of promoter behavior. For a more detailed comparison of promoter elements, see the [Promoter Sequence](/knowledge/molecular-biology/promoter-sequence) article.

## Experimental Methods to Study the TATA Box

### Mutational Analysis

The most direct way to study the TATA box is to mutate it and measure the effect on transcription. This is typically done using a reporter assay, where the promoter of interest is cloned upstream of a reporter gene such as luciferase or green fluorescent protein (GFP). The TATA box is mutated by site-directed mutagenesis, and the mutant promoter is compared to the wild-type promoter in a transient transfection assay.

A typical experiment might involve the following steps:

1. Clone the wild-type promoter (e.g., 500 base pairs upstream of the TSS) into a luciferase reporter vector.
2. Generate a series of mutants: a complete deletion of the TATA box, a single-base substitution (e.g., TATAAAAG → TATAAACG), and a scrambled sequence (e.g., TATAAAAG → AAGATATA).
3. Transfect each construct into a relevant cell line (e.g., HeLa or HEK293) using a lipid-based transfection reagent.
4. After 24–48 hours, lyse the cells and measure luciferase activity using a luminometer. Normalize to a co-transfected control (e.g., Renilla luciferase under a constitutive promoter).
5. Compare the activity of the mutants to the wild-type. A reduction of >80% in luciferase activity indicates that the TATA box is essential for promoter function.

Mutational analysis can also be combined with *in vitro* transcription assays, where nuclear extracts are used to transcribe a DNA template containing the promoter. This approach allows for more precise control of the reaction conditions and can be used to measure the kinetics of PIC assembly.

### Electrophoretic Mobility Shift Assay (EMSA)

The electrophoretic mobility shift assay (EMSA), also known as a gel shift assay, is used to detect protein–DNA interactions. In the context of the TATA box, EMSA is used to measure the binding of TBP to the TATA box sequence.

The procedure is as follows:

1. Synthesize a double-stranded DNA oligonucleotide (approximately 30–40 base pairs) containing the TATA box sequence. Label the 5' end with a radioactive isotope (e.g., ³²P) or a fluorescent dye.
2. Incubate the labeled DNA with recombinant TBP (or nuclear extract) in a binding buffer containing 10 mM Tris-HCl (pH 7.5), 50 mM KCl, 5 mM MgCl₂, 1 mM DTT, and 5% glycerol. The reaction is typically incubated at room temperature for 20–30 minutes.
3. Load the reaction onto a native polyacrylamide gel (4–6% acrylamide) and run at 100 V for 1–2 hours at 4°C.
4. Visualize the gel by autoradiography or fluorescence imaging. The free DNA migrates faster than the TBP–DNA complex, which appears as a slower-migrating band.

To confirm specificity, a competition assay can be performed: add an excess of unlabeled TATA box DNA to the reaction. If the binding is specific, the unlabeled DNA will compete with the labeled DNA, and the shifted band will disappear. A mutant TATA box (e.g., TATAAAAG → TATCCCCG) should not compete.

EMSA can also be used to measure the dissociation constant (Kd) of TBP for different TATA box variants. By titrating increasing amounts of TBP and quantifying the fraction of bound DNA, one can calculate the Kd using a binding isotherm.

### Chromatin Immunoprecipitation (ChIP)

Chromatin immunoprecipitation (ChIP) is used to determine whether TBP (or other transcription factors) is bound to the TATA box in living cells. This technique provides information about the occupancy of the TATA box under different conditions.

The ChIP protocol involves the following steps:

1. Crosslink proteins to DNA by adding formaldehyde to the cell culture medium at a final concentration of 1% and incubating for 10 minutes at room temperature. Quench the crosslinking by adding glycine to a final concentration of 125 mM.
2. Lyse the cells and sonicate the chromatin to shear the DNA into fragments of approximately 200–600 base pairs. This is typically done using a probe sonicator at 30% amplitude for 10–15 cycles of 30 seconds on/30 seconds off.
3. Incubate the sheared chromatin with an antibody specific to TBP (e.g., anti-TBP, clone 1TBP18) overnight at 4°C with rotation.
4. Add protein A/G agarose beads to capture the antibody–protein–DNA complexes. Wash the beads extensively with a buffer containing 150 mM NaCl, 0.1% SDS, 1% Triton X-100, and 10 mM Tris-HCl (pH 8.0).
5. Elute the DNA from the beads by heating at 65°C for 4–6 hours to reverse the crosslinks. Purify the DNA using a column-based kit.
6. Analyze the purified DNA by quantitative PCR (qPCR) using primers that flank the TATA box region. The enrichment of TBP at the TATA box is calculated as the fold enrichment over a negative control region (e.g., an intergenic region).

ChIP can be combined with high-throughput sequencing (ChIP-seq) to map TBP binding across the entire genome. This approach has revealed that TBP binds to many TATA-less promoters, indicating that TBP can be recruited to promoters without directly binding to a TATA box.

## TATA Box in Different Organisms

The TATA box is found across all eukaryotes, but its prevalence and sequence preferences vary significantly among organisms.

In *Saccharomyces cerevisiae*, the TATA box is present in approximately 20% of promoters. Yeast TATA boxes are typically located 40–120 base pairs upstream of the TSS, which is farther than in metazoans. Yeast also has a higher tolerance for TATA box sequence variants; for example, TATATA and TATAA are both functional. The yeast TBP is a single polypeptide that can bind to the TATA box independently of other TFIID subunits, whereas in higher eukaryotes, TBP is always associated with TAFs (TBP-associated factors) as part of the TFIID complex.

In plants, the TATA box is found in approximately 30% of promoters, and its position is similar to that in animals (approximately 30 base pairs upstream of the TSS). Plant TATA boxes are often flanked by a pyrimidine-rich region, and the consensus sequence is TATATA or TATAA. The plant TBP is highly conserved, and the mechanism of PIC assembly is similar to that in animals.

In humans, the TATA box is present in only 10–20% of promoters. TATA-containing promoters are enriched for genes that are tightly regulated, such as those involved in development, stress response, and tissue-specific functions. In contrast, housekeeping genes (e.g., *GAPDH*, *ACTB*) typically lack a TATA box and instead have CpG island promoters. The TATA box is also more common in genes with a focused TSS (a single or few start sites) compared to genes with dispersed TSSs.

The evolutionary conservation of the TATA box is striking. The TBP protein itself is one of the most conserved proteins in eukaryotes, with the C-terminal core domain sharing >80% sequence identity between yeast and humans. This conservation underscores the fundamental importance of the TATA box–TBP interaction in transcription.

## Regulation of TATA Box Activity

The activity of the TATA box is not static; it is regulated by a variety of mechanisms that control the accessibility of the TATA box to TBP and the stability of the PIC.

One major regulatory mechanism is [chromatin structure](/knowledge/molecular-biology/chromatin-structure). In eukaryotic cells, DNA is packaged into nucleosomes, which can occlude the TATA box and prevent TBP binding. Nucleosome remodeling complexes such as SWI/SNF and RSC can reposition or evict nucleosomes to expose the TATA box. For example, the yeast *PHO5* promoter contains a TATA box that is initially buried in a nucleosome; upon phosphate starvation, the SWI/SNF complex remodels the chromatin to expose the TATA box, allowing transcription to occur.

Histone modifications also play a role. Acetylation of histone H3 and H4 lysine residues is associated with open chromatin and increased TBP binding. Conversely, methylation of histone H3 at lysine 9 (H3K9me) or lysine 27 (H3K27me) is associated with closed chromatin and reduced TBP occupancy. The [histone acetyltransferase](/knowledge/molecular-biology/histone-acetyltransferase) p300/CBP is often recruited to promoters by activators and acetylates histones, thereby facilitating TBP binding.

DNA methylation is another regulatory layer. In mammals, CpG dinucleotides within or near the TATA box can be methylated, which inhibits TBP binding. However, TATA-containing promoters are generally CpG-poor, so this mechanism is more relevant for TATA-less promoters.

The activity of the TATA box is also regulated by transcription factors that bind to upstream promoter elements or enhancers. Activators such as Gal4 (in yeast) or p53 (in mammals) can recruit TBP and TFIID to the TATA box through direct protein–protein interactions. For example, the acidic activation domain of Gal4 binds to the TBP-associated factor TAF9, stabilizing the binding of TFIID to the TATA box. Repressors can have the opposite effect: the yeast repressor Mot1 is an ATPase that removes TBP from the TATA box, while NC2 (negative cofactor 2) binds to TBP and prevents the recruitment of TFIIB.

Enhancers can also influence TATA box activity through [Enhancer Promoter Interaction](/knowledge/molecular-biology/enhancer-promoter-interaction). Enhancers are distal regulatory elements that can loop to the promoter and deliver additional transcription factors and coactivators. This looping is mediated by proteins such as CTCF and cohesin, and it can bring activators into proximity with the TATA box, increasing the local concentration of TBP and other PIC components. The distinction between enhancers and promoters is discussed in detail in the [Difference Between Enhancer and Promoter](/knowledge/molecular-biology/difference-between-enhancer-and-promoter) article.

## Common Misconceptions and Pitfalls

Several misconceptions about the TATA box are common among students and even some researchers. Being aware of these pitfalls will help you avoid errors in both understanding and experimental design.

**Misconception 1: The TATA box is the promoter.** The TATA box is a core promoter element, but the promoter is a much larger region that includes all sequences required for transcription initiation. The [Promoter Region](/knowledge/molecular-biology/promoter-region) includes the core promoter (TATA box, Inr, DPE) and proximal promoter elements (GC boxes, CAAT boxes). Saying "the TATA box is the promoter" is like saying "the ignition is the car."

**Misconception 2: All promoters have a TATA box.** As discussed, only 10–20% of human promoters contain a TATA box. Many genes, especially housekeeping genes, use TATA-less promoters with CpG islands or DPE elements. If you are analyzing a promoter and do not find a TATA box, this is not an error; it is a common and biologically meaningful finding.

**Misconception 3: The TATA box sequence is always TATAAA.** The consensus is TATAWAWR, and many functional variants exist. For example, the *Drosophila hsp70* promoter has a TATA box with the sequence TATAAATA, while the human *β-globin* promoter has TATAAAAG. A sequence that deviates from TATAAA is not necessarily non-functional.

**Misconception 4: TBP only binds to the TATA box.** TBP can bind to other sequences, especially in TATA-less promoters. It can be recruited to promoters through protein–protein interactions with tethering factors such as TFTC (TBP-free TAF-containing complex) or through binding to the Inr element. ChIP experiments often show TBP occupancy at promoters that lack a canonical TATA box.

**Misconception 5: The TATA box is always 25–30 base pairs upstream of the TSS.** This is true for most metazoan promoters, but in yeast, the distance can be 40–120 base pairs. Additionally, some promoters have a TATA box that is further upstream or downstream due to unusual PIC architecture. Always verify the position empirically rather than assuming a fixed distance.

**Pitfall in experimental design: Mutating the TATA box without considering flanking sequences.** The flanking sequences can influence TBP binding affinity. If you mutate the TATA box and observe no effect on transcription, it may be because the flanking sequences compensate for the loss of the TATA box. Conversely, if you observe a large effect, it may be due to disruption of a flanking transcription factor binding site rather than the TATA box itself. Always include a control with a scrambled TATA box sequence rather than a simple deletion.

**Pitfall in bioinformatics: Using a simple string search to identify TATA boxes.** A naive search for "TATAAA" will produce many false positives and false negatives. Use position weight matrices (PWMs) that account for the degeneracy of the TATA box, and consider the positional distribution relative to the TSS. Tools such as the Eukaryotic Promoter Database (EPD) provide curated TATA box annotations.

## Summary and Key Takeaways

The TATA box is a core promoter element that plays a central role in transcription initiation by serving as the binding site for TBP. Its location, consensus sequence, and function are well characterized, but its prevalence and regulation vary across organisms and gene classes. Understanding the TATA box is essential for interpreting promoter function, designing reporter assays, and analyzing gene expression data.

### Key Takeaways

- The TATA box is a conserved DNA motif (consensus TATAWAWR) located approximately 25–30 base pairs upstream of the transcription start site in metazoans.
- The TATA box is recognized by the TATA-binding protein (TBP), a subunit of TFIID, which bends the DNA and nucleates the assembly of the preinitiation complex.
- Only 10–20% of human promoters contain a TATA box; the majority are TATA-less and rely on Inr, DPE, or CpG islands.
- The TATA box is not the promoter; it is one element within the core promoter region.
- TATA box activity is regulated by [chromatin structure](/knowledge/molecular-biology/chromatin-structure), histone modifications, DNA methylation, and transcription factors that recruit or displace TBP.
- Experimental methods to study the TATA box include mutational analysis, EMSA, ChIP, and structural studies.
- Common pitfalls include assuming all promoters have a TATA box, misidentifying the TATA box sequence, and ignoring the role of flanking sequences.

## Frequently Asked Questions

### Is the TATA box a promoter?

No. The TATA box is a core promoter element, but the promoter is the entire region of DNA required for transcription initiation. The promoter includes the core promoter (which contains the TATA box, Inr, and DPE) and proximal promoter elements (such as GC boxes and CAAT boxes). The TATA box is one component of the promoter, not the promoter itself.

### What is the TATA box in a promoter?

The TATA box is a conserved DNA sequence motif located within the core promoter of many eukaryotic genes. It is the binding site for the TATA-binding protein (TBP), which is a subunit of the general transcription factor TFIID. The TATA box is essential for the assembly of the preinitiation complex and the accurate initiation of transcription by RNA polymerase II.

### What is the function of the TATA box?

The primary function of the TATA box is to recruit TBP to the promoter and nucleate the assembly of the preinitiation complex. TBP binding to the TATA box bends the DNA and creates a platform for the recruitment of TFIIA, TFIIB, RNA polymerase II, TFIIE, TFIIF, and TFIIH. This assembly is required for the unwinding of the DNA at the transcription start site and the initiation of RNA synthesis.

### Do all promoters have a TATA box?

No. Only 10–20% of human promoters contain a TATA box. Many promoters, especially those of housekeeping genes, are TATA-less and rely on other core promoter elements such as the initiator (Inr), downstream promoter element (DPE), or CpG islands. The presence or absence of a TATA box is a useful predictor of promoter regulation, but it is not a universal feature.

### Where is the TATA box located?

In metazoans, the TATA box is typically located 25–30 base pairs upstream of the transcription start site. In yeast, it is often found 40–120 base pairs upstream. The exact position can vary between genes, and some promoters have TATA boxes at non-canonical positions.

### What is the consensus sequence of the TATA box?

The consensus sequence is TATAWAWR, where W is A or T, and R is A or G. The most common form is TATAAAAG. The first four nucleotides (TATA) are the most highly conserved and are critical for TBP binding.

### How is the TATA box recognized?

The TATA box is recognized by the TATA-binding protein (TBP), which binds to the minor groove of the DNA helix. TBP induces a bend of approximately 80 degrees in the DNA and widens the minor groove. This binding is stabilized by TFIIA and TFIIB, which help recruit RNA polymerase II and the other general transcription factors to form the preinitiation complex.

## Further Reading

- Zou Y et al. *Predominant gain of promoter TATA box after gene duplication associated with stress responses*. [Molecular biology](/blog/careers/molecular-biology) and evolution. 2011. [PubMed 21515810](https://doi.org/10.1093/molbev/msr116)
- Zhang Y et al. *Cellular microRNAs up-regulate transcription via interaction with promoter TATA-box motifs*. RNA (New York, N.Y.). 2014. [PubMed 25336585](https://doi.org/10.1261/rna.045633.114)
- Zhang Y et al. *A Cellular MicroRNA Facilitates Regulatory T Lymphocyte Development by Targeting the FOXP3 Promoter TATA-Box Motif*. Journal of immunology (Baltimore, Md. : 1950). 2018. [PubMed 29282311](https://doi.org/10.4049/jimmunol.1700196)
- Han SO, Miller WL. *Activin A induces ovine follicle stimulating hormone beta using -169/-58 bp of its promoter and a simple TATA box*. Reproductive biology and endocrinology : RB&E. 2009. [PubMed 19552818](https://doi.org/10.1186/1477-7827-7-66)
- Gui CY, Dean A. *A major role for the TATA box in recruitment of chromatin modifying complexes to a globin gene promoter*. Proceedings of the National Academy of Sciences of the United States of America. 2003. [PubMed 12773626](https://doi.org/10.1073/pnas.1236499100)
- Baliga NS, DasSarma S. *Saturation mutagenesis of the TATA box and upstream activator sequence in the haloarchaeal bop gene promoter*. Journal of bacteriology. 1999. [PubMed 10198017](https://doi.org/10.1128/JB.181.8.2513-2518.1999)

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