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In an equilibrium model, the state of a binding site on DNA depends on the concentration of the protein that binds to it. The Nucleosome Is the Subunit of All Chromatin The nucleosome may be a cylinder with DNA organized into two turns around the surface. Sequences on the DNA that lie on different turns around the nucleosome may be close together. Suppose that the DNA sequence is organized into nucleosomes in only one particular configuration, so that each site on the DNA always is located at a particular position on the nucleosome. This type of organization is called nucleosome positioning (or sometimes nucleosome phasing). Nucleosome positioning places restriction sites at unique positions relative to the linker sites cleaved by micrococcal nuclease. Chromatin Can Have Alternative States Chromatin Remodeling Is an Active Process The dynamic model for transcription of chromatin relies upon factors that can use energy provided by hydrolysis of ATP to displace nucleosomes from specific DNA sequences. Remodeling complexes can cause nucleosomes to slide along DNA, can displace nucleosomes from DNA, or can reorganize the spacing between nucleosomes. Nucleosome Organization May Be Changed at the Promoter Remodeling complexes are recruited to promoters by sequence-specific activators. The factor may be released once the remodeling complex has bound. The MMTV promoter requires a change in rotational positioning of a nucleosome to allow an activator to bind to DNA on the nucleosome. Histone Modification Is a Key Event Whether a gene is expressed depends on the structure of chromatin both locally (at the promoter) and in the surrounding domain. Chromatin structure correspondingly can be regulated by individual activation events or by changes that affect a wide chromosomal region. The most localized events concern an individual target gene, where changes in nucleosomal structure and organization occur in the immediate vicinity of the promoter. Histones
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