are RNA molecules (the product of transcription), RNA polymerases (RNAP) which control transcription, and Gyrases (G) which regulate PSB. Finally, there needs to be a means to quantify PSB on the DNA (i.e. the promoter) at any given moment. This can be done by having some component in the system that is produced over time (e.g., during transcription events) to represent positive supercoils, and that is removed by the action of Gyrases. The amount of this component can then be set to affect the rate of transcription. Effects on sedimentation coefficient The topological properties of circular DNA are complex. In standard texts, these properties are invariably explained in terms of a helical model for DNA, but in 2008 it was noted that each topoisomer, negative or positive, adopts a unique and surprisingly wide distribution of three-dimensional conformations. When the sedimentation coefficient, s, of circular DNA is ascertained over a large range of pH, the following curves are seen. Three curves are shown here, representing three species of DNA. From top-to-bottom they are: "Form IV" (green), "Form I" (blue) and "Form II" (red). "Form I" (blue curve) is the traditional nomenclature used for the native form of duplex circular DNA, as recovered from viruses and intracellular plasmids. Form I is covalently closed, and any plectonemic winding which may be present is therefore locked in. If one or more nicks are introduced to Form I, free rotation of one strand with respect to the other becomes possible, and Form II (red curve) is seen. Form IV (green curve) is the product of alkali denaturation of Form I. Its structure is unknown, except that it is persistently duplex, and extremely dense. Between pH 7 and pH 11.5, the sedimentation coefficient s, for Form I, is constant. Then it dips, and at a pH just below 12, reaches a minimum. With further increases in pH, s then returns to its former value. It doesn’t stop there, however, but continues to increase relentlessly. By pH 13, the value of s has risen to nearly 50, two to three times its value at pH 7, indicating an extremely compact structure. If the pH is then lowered, the s value is not restored. Instead, one sees the upper, green curve. The DNA, now in the state known as Form IV, remains extremely dense, even if the pH is restored to the original physiologic range. As stated previously,