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The transcription cycle. The transcription cycle can be described in six steps: (1) Template binding and closed RNA polymerase-promoter complex formation: RNAP binds to DNA and then locates a promoter (P), (2) Open promoter complex formation: once bound to the promoter, RNAP melts the two DNA strands to form an open promoter complex; this complex is also referred to as the preinitiation complex or PIC. Strand separation allows the polymerase to access the coding information in the template strand of DNA (3) Chain initiation: using the coding information of the template RNAP catalyzes the coupling of the first base (often a purine) to the second, template-directed ribonucleoside triphosphate to form a dinucleotide (in this example forming the dinucleotide 5′ pppApNOH 3′). (4) Promoter clearance: after RNA chain length reaches ~10–20 nt, the polymerase undergoes a conformational Source: RNA Synthesis, Processing, & Modification, Harper's Illustrated Biochemistry, 30e change and then is able to move away from the promoter, transcribing down the transcription unit. (5) Chain elongation: Successive residues are added to Citation: VW,RNA Bender DA, Botham Kennellytermination PJ, Weil P. signal Harper's Biochemistry, 30e; 2015 Available at: release: Upon the 3′-OH terminus of Rodwell the nascent molecule until a KM, transcription (T) Illustrated is encountered. (6) Chain termination and RNAP http://mhmedical.com/ Accessed: August 03, 2017 encountering the transcription termination site RNAP undergoes an additional conformational change that leads to release of the completed RNA chain, Copyright © 2017 RNAP McGraw-Hill Education. rights reserved the DNA template and RNAP. can rebind to DNAAllbeginning the promoter search process and the cycle is repeated. Note that all of the steps in the transcription cycle are facilitated by additional proteins, and indeed are often subjected to regulation by positive and/or negative-acting factors.