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Chromatin modification and remodeling

Chromatin is refractory to the process of transcription and must be modified to allow access to the DNA. The usual initial step of this modification is the binding of “pioneering” transcription factors to their cognate sequences. These factors then recruit chromatin remodeling and modifying complexes. Nucleosomes can be displaced, or their association with DNA can be altered by multiprotein complexes that hydrolyze ATP to generate the energy necessary for their functions. Other complexes replace certain canonical histones with specific histone variants. Histones can undergo acetylation, methylation, phosphorylation, ubiquitination, sumoylation, glycosylation, ADP ribosylation or hydroxyisobutyrylation of particular amino acids along their molecules. DNA nucleotides can be modified by methylation or hydroxymethylation, without altering the coding sequences. All of these chromatin changes are associated with gene function, with sets of modifications usually present on active or repressed genes. Not surprisingly, mutations that affect the timely occurrence or removal of these modifications result in particular diseases.