Transcription Process: From DNA to RNA Explained

By Dr. Zubair Khalid, DVM, MS, PhD · 2026-08-11

Introduction to Transcription

Transcription is the enzymatic process by which the genetic information encoded in a DNA template is copied into a complementary RNA molecule. This process is the first step in gene expression and is fundamental to all living organisms. The term "transcription" is apt: the DNA sequence is transcribed—rewritten in the same language of nucleic acids—into RNA, as opposed to translation, where the language of nucleic acids is converted into the language of proteins.

Central Dogma Overview

The central dogma of molecular biology, first articulated by Francis Crick in 1957, describes the directional flow of genetic information: DNA → RNA → Protein. Transcription occupies the first arrow in this scheme. It is the point at which the genetic blueprint stored in the nucleus (in eukaryotes) or the nucleoid (in prokaryotes) is converted into a mobile, transient message—messenger RNA (mRNA)—that can be transported to the ribosome for protein synthesis. The transcription translation coupling is particularly notable in prokaryotes, where ribosomes can begin translating an mRNA molecule while it is still being transcribed, a phenomenon absent in eukaryotes due to nuclear compartmentalization.

Transcription is also responsible for producing non-coding RNAs: ribosomal RNA (rRNA), transfer RNA (tRNA), microRNAs, and long non-coding RNAs. These molecules are not translated into protein but serve structural, catalytic, or regulatory roles. Thus, transcription is not merely a precursor to translation; it is a major regulatory hub in its own right.

RNA vs DNA

Before examining the machinery, it is essential to understand the chemical differences between RNA and DNA that make transcription mechanistically distinct from DNA replication.

The 2'-hydroxyl group on ribose makes RNA more susceptible to alkaline hydrolysis and to attack by ribonucleases, which is why RNA is typically short-lived compared to DNA. The use of uracil instead of thymine is functionally significant: cytosine can undergo spontaneous deamination to uracil. In DNA, this mutation is recognized and repaired because thymine is the "correct" base; in RNA, there is no such repair system, and the transient nature of RNA makes this distinction less critical.

The Transcription Machinery

The core components of the transcription apparatus are the RNA polymerase enzyme, the DNA template, and a set of accessory proteins that direct the polymerase to the correct start site.

RNA Polymerase Structure

RNA polymerase (RNAP) is a large, multi-subunit enzyme that catalyzes the formation of phosphodiester bonds between ribonucleoside triphosphates (rNTPs). In prokaryotes, a single RNAP core enzyme (approximately 400 kDa) is responsible for all RNA synthesis. The E. coli core enzyme has five subunits: α₂ββ'ω. The two α subunits are involved in enzyme assembly and interaction with regulatory factors; the β and β' subunits form the catalytic cleft that accommodates the DNA template and the growing RNA chain; the ω subunit assists in assembly and stability.

The core enzyme alone can synthesize RNA but cannot initiate transcription at the correct sites. A sixth subunit, sigma (σ), associates with the core to form the holoenzyme. Sigma factors confer promoter specificity. The primary sigma factor in E. coli, σ⁷⁰, recognizes consensus sequences at the −35 and −10 positions relative to the transcription start site. Alternative sigma factors (e.g., σ³² for heat shock, σ⁵⁴ for nitrogen metabolism) redirect the polymerase to different promoter sets under stress conditions.

Eukaryotes have three nuclear RNA polymerases, each with distinct functions:

• RNA Polymerase I: Transcribes ribosomal RNA genes (28S, 18S, 5.8S rRNA).
• RNA Polymerase II: Transcribes all protein-coding genes (mRNA) and many non-coding RNAs. This is the enzyme most relevant to the transcription process described here.
• RNA Polymerase III: Transcribes tRNA, 5S rRNA, and other small RNAs.

RNA Polymerase II has 12 subunits (Rpb1–Rpb12) in yeast; the largest subunit, Rpb1, contains a C-terminal domain (CTD) consisting of heptapeptide repeats (Tyr-Ser-Pro-Thr-Ser-Pro-Ser). This CTD is phosphorylated during the transcription cycle and serves as a platform for recruiting RNA processing factors.

Promoters and Enhancers

A promoter is a DNA sequence, typically 100–1000 base pairs in eukaryotes, that directs RNA polymerase to the transcription start site. In prokaryotes, the promoter consists of two conserved hexamers: the −35 box (TTGACA) and the −10 box (TATAAT, also called the Pribnow box). The spacing between these elements is critical—usually 16–18 base pairs—and the TATA box transcription element is recognized by σ⁷⁰.

Eukaryotic RNA Polymerase II promoters are more complex. The core promoter typically contains a TATA box (consensus TATAAAA) located approximately 25–30 base pairs upstream of the start site, recognized by the TATA-binding protein (TBP). Other core promoter elements include the initiator (Inr) sequence, the downstream promoter element (DPE), and the TFIIB recognition element (BRE). Promoters can also contain upstream regulatory elements such as GC boxes (bound by Sp1) and CAAT boxes (bound by NF-Y).

Enhancers are distal regulatory DNA elements, often located thousands of base pairs away from the promoter, that increase transcription levels. They function by looping the DNA so that enhancer-bound activator proteins can contact the basal transcription machinery at the promoter. Enhancers are orientation- and position-independent, meaning they can function upstream, downstream, or even within the gene they regulate.

Initiation of Transcription

Initiation is the most highly regulated step of transcription. It involves promoter recognition, DNA unwinding, and the synthesis of the first few phosphodiester bonds. The details differ substantially between prokaryotes and eukaryotes, but the fundamental principles are conserved.

Promoter Binding

In prokaryotes, the σ⁷⁰ subunit of the RNAP holoenzyme scans the DNA for promoter sequences. The holoenzyme initially binds the DNA in a closed complex, where the DNA remains double-stranded. Upon recognition of the −35 and −10 elements, the polymerase undergoes a conformational change, and the DNA around the −10 region begins to melt. This transition from closed to open complex is driven by the unwinding of approximately 12–14 base pairs, creating a transcription bubble.

The open complex is highly stable at physiological temperatures (37°C for E. coli). However, the polymerase can abortively initiate: it synthesizes short RNA products (2–9 nucleotides) and releases them while remaining bound to the promoter. This abortive cycling continues until the polymerase successfully escapes the promoter, a process that requires the σ subunit to be released or to undergo a conformational change that weakens its promoter contacts.

In eukaryotes, promoter recognition is performed not by RNA Polymerase II itself but by a set of general transcription factors (GTFs). The order of assembly is:

1. TFIID binds to the TATA box via its TBP subunit. TFIID is a large complex containing TBP and 13–14 TBP-associated factors (TAFs).
2. TFIIA stabilizes TFIID binding.
3. TFIIB binds to the promoter, bridging TFIID and the polymerase.
4. The preformed RNA Polymerase II–TFIIF complex joins the assembly.
5. TFIIE and TFIIH bind, completing the preinitiation complex (PIC).

TFIIH is particularly important: it contains helicase subunits (XPB and XPD) that unwind the DNA around the start site, and a kinase subunit (CDK7) that phosphorylates the CTD of RNA Polymerase II. This phosphorylation triggers promoter escape and the transition to elongation.

Transcription Bubble Formation

The transcription bubble is a locally melted region of DNA where the template strand is exposed for base pairing with incoming rNTPs. In the open complex, the bubble spans approximately 13–17 base pairs, with the RNA-DNA hybrid occupying about 8–9 base pairs.

The formation of the bubble requires energy. In prokaryotes, this energy comes from the binding energy of the polymerase to the promoter and from thermal fluctuations. In eukaryotes, the ATP-dependent helicase activity of TFIIH drives bubble formation. The unwound region is stabilized by single-stranded DNA binding domains within the polymerase.

The first phosphodiester bond is formed between the two initiating rNTPs, typically a purine (ATP or GTP) at the +1 position. The initiating nucleotide remains in the active site, and the second nucleotide is added to its 3'-OH. After the synthesis of approximately 10 nucleotides, the polymerase breaks its contacts with the promoter and enters the elongation phase. This transition is called promoter escape and is a major kinetic bottleneck in transcription.

Elongation of the RNA Strand

During elongation, RNA polymerase moves processively along the DNA template, synthesizing RNA at a rate of approximately 20–50 nucleotides per second in prokaryotes and 20–30 nucleotides per second in eukaryotes. The enzyme maintains a transcription bubble of about 18 base pairs, with an 8–9 base pair RNA-DNA hybrid.

Nucleotide Addition

The catalytic mechanism of nucleotide addition is conserved across all RNA polymerases. The active site contains two Mg²⁺ ions (metal A and metal B) coordinated by conserved aspartate residues. The incoming rNTP enters the active site through a secondary channel and base-pairs with the template strand. Metal A activates the 3'-OH of the growing RNA chain for nucleophilic attack on the α-phosphate of the incoming rNTP. Metal B stabilizes the pentacovalent transition state and facilitates pyrophosphate release.

The reaction proceeds as follows:

1. The template base at position +1 (the next base to be copied) is positioned in the active site.
2. The incoming rNTP forms a Watson-Crick base pair with the template base.
3. The 3'-OH of the growing RNA chain attacks the α-phosphate of the rNTP.
4. Pyrophosphate (PPi) is released, and the RNA chain is extended by one nucleotide.
5. The polymerase translocates one base pair downstream, and the cycle repeats.

The RNA chain grows strictly in the 5' to 3' direction. The template strand is read in the 3' to 5' direction. This directionality is a common source of confusion; the polymerase moves along the template strand in the 3'→5' direction while synthesizing RNA in the 5'→3' direction.

Proofreading and Correction

RNA polymerases are less accurate than DNA polymerases, with an error rate of approximately 10⁻⁴ to 10⁻⁵ per base incorporated. This is acceptable because RNA is transient and errors in mRNA do not permanently alter the genome. However, RNA polymerase does possess two proofreading mechanisms:

1. Pyrophosphorolytic editing: The polymerase reverses the polymerization reaction, re-adding PPi to remove the misincorporated nucleotide and regenerate the rNTP.
2. Hydrolytic editing: The polymerase backtracks by one or more nucleotides and cleaves the RNA chain endonucleolytically, removing the erroneous segment. This mechanism is stimulated by the transcription factor GreA/GreB in prokaryotes and TFIIS in eukaryotes.

The backtracking mechanism is also important for resolving transcriptional pauses. RNA polymerase frequently pauses during elongation, particularly at sequences that form RNA hairpins or at sites of DNA damage. Pausing provides opportunities for regulatory factors to interact with the polymerase and for the enzyme to correct errors. The transcription error rate is thus kept within tolerable limits, though errors in mRNA can still lead to the production of mutant proteins.

Termination of Transcription

Termination is the process by which RNA polymerase stops transcription and releases both the RNA product and the DNA template. The mechanisms differ markedly between prokaryotes and eukaryotes.

Rho-Dependent Termination

In prokaryotes, rho-dependent termination requires the hexameric ATP-dependent helicase Rho. Rho binds to a rut (rho utilization) site on the nascent RNA, typically a C-rich, G-poor sequence of approximately 70–80 nucleotides. Rho moves along the RNA in the 5'→3' direction, tracking the polymerase. When the polymerase pauses at a downstream termination site, Rho catches up and uses its ATPase activity to translocate along the RNA, pulling it out of the polymerase active site and disrupting the RNA-DNA hybrid. This leads to the release of the transcript.

Rho-dependent terminators are less common than rho-independent ones and are often found in genes where rapid regulation is required, such as in bacteriophage λ genes.

Rho-Independent Termination

Rho-independent (intrinsic) termination is more common in E. coli and does not require accessory proteins. The terminator sequence contains two elements:

1. A GC-rich inverted repeat that forms a stable stem-loop (hairpin) structure in the nascent RNA.
2. A downstream run of 4–8 adenine residues in the template strand, which results in a run of uracils in the RNA.

The mechanism is as follows: as the polymerase transcribes the inverted repeat, the RNA folds into a hairpin. The hairpin formation destabilizes the elongation complex by disrupting the RNA-DNA hybrid at the upstream edge of the transcription bubble. The weak A-U base pairs in the hybrid (which have lower thermal stability than G-C pairs) facilitate the dissociation of the RNA from the template. The polymerase then releases the transcript and dissociates from the DNA.

The efficiency of intrinsic termination depends on the stability of the hairpin and the length of the U-run. Mutations that destabilize the hairpin or shorten the U-run reduce termination efficiency. This mechanism is described in detail under Transcription Termination.

Eukaryotic Termination

Eukaryotic termination is more complex and is coupled to RNA processing. For RNA Polymerase II, termination is linked to the cleavage and polyadenylation of the pre-mRNA. The key sequence elements are:

• The polyadenylation signal AAUAAA, located 10–30 nucleotides upstream of the cleavage site.
• A downstream GU-rich element.

As the polymerase transcribes past the polyadenylation signal, the CPSF (cleavage and polyadenylation specificity factor) and CstF (cleavage stimulation factor) complexes recognize these sequences on the nascent RNA. The RNA is cleaved at the polyadenylation site, and the polymerase continues transcribing. The "torpedo" model proposes that the 5'→3' exonuclease Rat1/Xrn2 degrades the RNA remaining attached to the polymerase, and upon catching up to the polymerase, triggers its release. The "allosteric" model proposes that passage through the polyadenylation signal induces a conformational change in the polymerase that destabilizes the elongation complex.

RNA Polymerase I terminates via a specific termination factor (Reb1 in yeast) that binds to a terminator sequence downstream of the rRNA genes. RNA Polymerase III terminates at a run of 4–5 thymine residues, similar to intrinsic termination in prokaryotes.

Post-Transcriptional Modifications in Eukaryotes

Eukaryotic pre-mRNA undergoes three major processing events before it is exported to the cytoplasm as mature mRNA. These modifications are essential for mRNA stability, nuclear export, and efficient translation.

5' Capping

The 5' cap is added co-transcriptionally, when the nascent RNA is only 20–30 nucleotides long. The cap is a 7-methylguanosine linked to the first transcribed nucleotide via a 5'→5' triphosphate bridge (m⁷GpppN). The capping reaction occurs in three steps:

1. RNA triphosphatase removes the γ-phosphate from the 5' end.
2. Guanylyltransferase adds a GMP moiety from GTP, forming the 5'→5' linkage.
3. Methyltransferase adds a methyl group to the N7 position of the guanine.

The cap serves several functions: it protects the mRNA from 5'→3' exonucleases, it is recognized by the cap-binding complex (CBC) for nuclear export, and it is required for ribosome binding during translation initiation. The cap also plays a role in splicing by recruiting the U1 snRNP to the first exon.

3' Polyadenylation

The 3' end of most eukaryotic mRNAs is modified by the addition of a poly(A) tail of 50–250 adenine residues. This process is coupled to transcription termination and occurs in two steps:

1. Endonucleolytic cleavage of the pre-mRNA at a site 10–30 nucleotides downstream of the AAUAAA signal.
2. Processive addition of adenines by poly(A) polymerase (PAP).

The poly(A) tail is bound by poly(A)-binding protein (PABP), which protects the mRNA from 3'→5' degradation and promotes translation initiation. The length of the poly(A) tail can be regulated; deadenylation is often the first step in mRNA decay.

RNA Splicing

Most eukaryotic genes contain introns—non-coding sequences that must be removed from the pre-mRNA. Splicing is catalyzed by the spliceosome, a large ribonucleoprotein complex composed of five small nuclear RNAs (U1, U2, U4, U5, U6) and over 100 proteins.

The splicing reaction occurs in two transesterification steps:

1. The 2'-OH of the branch point adenosine (located 18–40 nucleotides upstream of the 3' splice site) attacks the 5' splice site, cleaving the RNA and forming a lariat structure.
2. The 3'-OH of the 5' exon attacks the 3' splice site, joining the two exons and releasing the intron as a lariat.

Splicing is guided by consensus sequences at the 5' splice site (GU), the 3' splice site (AG), and the branch point. Alternative splicing allows a single gene to produce multiple mRNA isoforms, greatly expanding the proteomic diversity of eukaryotes. It is estimated that over 95% of human multi-exon genes undergo alternative splicing.

Regulation of Transcription

Transcription is the primary point of regulation for most genes. The regulation occurs at multiple levels, from the accessibility of the DNA to the activity of the polymerase itself.

Transcription Factors

Transcription factors are proteins that bind to specific DNA sequences and modulate transcription. They are classified into two broad categories:

• General transcription factors (GTFs): Required for transcription of all genes (TFIIA, TFIIB, TFIID, TFIIE, TFIIF, TFIIH in eukaryotes).
• Specific transcription factors: Regulate subsets of genes by binding to enhancers or silencers.

Specific transcription factors typically contain a DNA-binding domain (e.g., zinc finger, helix-turn-helix, leucine zipper) and an activation domain that recruits coactivators or the basal machinery. The Transcription Factor family includes well-studied examples such as p53, NF-κB, and steroid hormone receptors.

Enhancers and Silencers

Enhancers are DNA elements that increase transcription from a linked promoter, while silencers decrease it. Both function through the binding of specific transcription factors and the looping of DNA to bring these factors into proximity with the promoter.

Enhancers can act over long distances (up to 1 Mb in mammals) and can be located upstream, downstream, or within introns of the genes they regulate. The looping model is supported by chromosome conformation capture (3C) experiments, which show physical interactions between enhancers and promoters. Insulator elements, bound by CTCF in vertebrates, demarcate the boundaries of enhancer-promoter interactions.

Chromatin Structure

In eukaryotes, DNA is packaged into chromatin, with 147 base pairs of DNA wrapped around a histone octamer to form a nucleosome. This packaging presents a barrier to transcription: RNA polymerase cannot easily access DNA that is tightly wrapped around histones.

Chromatin remodeling is achieved by two classes of enzymes:

1. ATP-dependent chromatin remodelers (e.g., SWI/SNF, ISWI, CHD): Use ATP hydrolysis to slide, eject, or restructure nucleosomes.
2. Histone-modifying enzymes: Covalently modify histone tails, including acetylation (by HATs), methylation (by HMTs), and phosphorylation.

Histone acetylation at lysine residues neutralizes the positive charge of histones, weakening their interaction with DNA and promoting a more open chromatin state. Histone methylation can either activate or repress transcription depending on the specific lysine residue modified (e.g., H3K4me3 is associated with active promoters, while H3K27me3 is associated with repressed genes).

Methods to Study Transcription

Several experimental approaches are used to study transcription, each providing different types of information.

Reporter Assays

Reporter assays measure the activity of a promoter by placing it upstream of a reporter gene whose product is easily quantifiable. Common reporters include:

• Luciferase: Produces light in the presence of its substrate luciferin.
• Green fluorescent protein (GFP): Fluoresces when excited by blue light.
• β-galactosidase (LacZ): Cleaves X-gal to produce a blue color.

In a typical experiment, the promoter of interest is cloned upstream of the luciferase gene, and the construct is transfected into cells. After 24–48 hours, cells are lysed, and luciferase activity is measured using a luminometer. The activity reflects the strength of the promoter under the tested conditions.

ChIP-seq

Chromatin immunoprecipitation followed by sequencing (ChIP-seq) identifies the genomic locations where a specific protein (e.g., a transcription factor or a histone modification) is bound. The protocol involves:

1. Crosslinking proteins to DNA with formaldehyde.
2. Shearing the chromatin by sonication into fragments of 200–600 base pairs.
3. Immunoprecipitating the protein of interest with a specific antibody.
4. Reversing the crosslinks and purifying the DNA.
5. Sequencing the DNA fragments and mapping them to the genome.

ChIP-seq can identify transcription factor binding sites genome-wide, reveal the distribution of histone modifications, and map the positions of RNA polymerase II. It is a powerful tool for understanding transcriptional regulation.

RNA-seq

RNA sequencing (RNA-seq) quantifies the transcriptome—the complete set of RNA molecules in a cell. The workflow involves:

1. Isolating total RNA.
2. Enriching for mRNA (typically by poly(A) selection).
3. Fragmenting the RNA and converting it to cDNA.
4. Adding sequencing adapters and amplifying.
5. Sequencing and mapping the reads to a reference genome or transcriptome.

RNA-seq can measure gene expression levels, identify differentially expressed genes between conditions, detect alternative splicing isoforms, and discover novel transcripts. It has largely replaced microarrays as the standard method for transcriptome analysis.

Common Pitfalls and Misconceptions

Students frequently encounter specific conceptual difficulties when learning about transcription. Addressing these directly can prevent persistent misunderstandings.

Template vs Coding Strand

The most common error is confusing which DNA strand is transcribed. The template strand (also called the antisense or minus strand) is read by RNA polymerase and is complementary to the RNA product. The coding strand (also called the sense or plus strand) has the same sequence as the RNA (with T instead of U) and is not transcribed.

A useful mnemonic: the coding strand "codes" for the protein—its sequence matches the mRNA. When reading a gene sequence in a textbook, the sequence shown is usually the coding strand (5'→3'), and the RNA produced will have the same sequence (with U replacing T). The template strand is the one that actually pairs with the incoming rNTPs.

Directionality Errors

Another common error involves the direction of synthesis. RNA is always synthesized in the 5'→3' direction. This means:

• The template strand is read in the 3'→5' direction.
• The promoter is located upstream (5') of the coding sequence on the coding strand.
• The transcription start site is designated +1, with upstream sequences given negative numbers (−10, −35).

Students often mistakenly think that RNA polymerase moves 5'→3' along the template strand. In fact, it moves 3'→5' along the template, which results in 5'→3' RNA synthesis. The Transcription Diagram can help visualize this geometry.

RNA Polymerase vs DNA Polymerase

RNA polymerase and DNA polymerase share some features but differ in critical ways:

RNA polymerase does not require a primer because it can initiate de novo by bringing two rNTPs together in the active site. DNA polymerase requires a pre-existing 3'-OH to extend. This difference is fundamental to their respective roles: transcription needs to initiate at specific sites frequently, while replication must be highly accurate and processive.

Frequently Asked Questions

What is the transcription process?

Transcription is the synthesis of an RNA molecule from a DNA template. The enzyme RNA polymerase reads the template strand of DNA and produces a complementary RNA strand in the 5' to 3' direction. This process is the first step in gene expression and converts the genetic information stored in DNA into a form that can be used to direct protein synthesis.

What are the main steps of transcription?

Transcription occurs in three main stages: initiation (RNA polymerase binds to the promoter and unwinds the DNA), elongation (the polymerase moves along the template, adding nucleotides to the growing RNA chain), and termination (the polymerase recognizes a stop signal, releases the RNA, and dissociates from the DNA). The Transcription Initiation step is the most highly regulated.

Where does transcription occur in a cell?

In prokaryotes, transcription occurs in the cytoplasm, where the DNA is located. In eukaryotes, transcription occurs in the nucleus, and the mature mRNA must be exported to the cytoplasm for translation. Mitochondria and chloroplasts also contain their own transcription machinery.

What is the role of RNA polymerase in transcription?

RNA polymerase is the core enzyme that catalyzes RNA synthesis. It binds to the DNA template, unwinds the double helix to form a transcription bubble, adds ribonucleotides complementary to the template strand, and moves processively along the DNA. In prokaryotes, a single RNA polymerase synthesizes all RNAs; in eukaryotes, three different polymerases (I, II, and III) transcribe different classes of genes.

What is the difference between template and coding strand?

The template strand is the DNA strand that is read by RNA polymerase; it is complementary to the RNA product. The coding strand is the other DNA strand; it has the same sequence as the RNA (except that thymine in DNA corresponds to uracil in RNA). The coding strand is the one typically shown in gene diagrams and databases.

How is transcription terminated?

Termination mechanisms differ between organisms. In prokaryotes, termination can be rho-independent (intrinsic), involving a hairpin structure followed by a U-run, or rho-dependent, requiring the Rho helicase to pull the RNA out of the polymerase. In eukaryotes, RNA Polymerase II termination is coupled to polyadenylation, with the Rat1/Xrn2 exonuclease degrading the residual RNA to trigger polymerase release.

What are post-transcriptional modifications?

Post-transcriptional modifications are processing events that occur to pre-mRNA in eukaryotes before it becomes mature mRNA. These include 5' capping (addition of 7-methylguanosine), 3' polyadenylation (addition of a poly(A) tail), and RNA splicing (removal of introns). These modifications are essential for mRNA stability, nuclear export, and translation.

Key Takeaways

• Transcription is the synthesis of RNA from a DNA template, catalyzed by RNA polymerase, and is the first step in gene expression.
• RNA polymerase reads the template strand in the 3'→5' direction and synthesizes RNA in the 5'→3' direction, without requiring a primer.
• The three stages of transcription are initiation, elongation, and termination, each with distinct molecular mechanisms.
• Prokaryotes use a single RNA polymerase with sigma factors for promoter recognition; eukaryotes use three RNA polymerases with general transcription factors.
• Eukaryotic pre-mRNA undergoes 5' capping, 3' polyadenylation, and splicing to produce mature mRNA.
• Transcription is regulated at multiple levels, including transcription factor binding, enhancer/silencer elements, and chromatin structure.
• The template strand is complementary to the RNA, while the coding strand matches the RNA sequence (with T→U).

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