Histones eukaryotic gene structure

Trievel, R.C. (2004) Structure and function of histone methyltransferases. Critical Reviews in Eukaryotic Gene Expression,... 

The genome is the total set of DNA-encoded genetic information in an organism. A chromosome is a DNA molecule, usually complexed with certain proteins. Chromatin is the partially decondensed form of eukaryotic chromosomes. Nucleosomes are the repeating structural units of eukaryotic chromosomes formed by the interaction of DNA with the histones. A gene is a DNA sequence that codes for a polypeptide or an RNA molecule. 

In the nuclei of all eukaryotic cells, DNA is tightly wrapped around an octamer of histone proteins and is compacted into a dense structure known as chromatin. In order to access the genetic information which is required in numerous essential cellular processes including DNA replication, gene expression and DNA repair, chromatin needs to be partially unwound. One important mechanism to regulate chromatin structure and thus to control the access of the genomic DNA is through histone modifications [1-6]. The histone octamer is composed of two copies of H2A, H2B, H3 and H4 core histone proteins. Their tails, that protrude out of the surface of the... 

Histone acetyltransferases (HATs) are enzymes that acetylate specific lysine residues in histones through the transfer of an acetyl group from an acetyl-coenzymeA (AcCoA) molecule, causing profound effects on chromatin structure and assembly as well as gene transcription. HATs are found in most, if not all, eukaryotic organisms as multiprotein complexes, some HAT catalytic subunits even being shared between various complexes that display different substrate specificities based on their subunit composition [12]. Despite their name, HATs do not restrict themselves to the acetylation of histones, since these enzymes have also been shown to act on nonhistone proteins, broadening their scope of action [13]. 

Histones are very basic proteins with an isoelectric point between 10.31 and 11.27 for human complement. They are present in virtually all eukaryotes (with the exception of dinoflagellates [14]) where they are associated with most of the nuclear DNA. The DNA is wrapped around an octamer formed by the four core histones H2A, H2B, H3 and H4 to build a nucleosome. This particle is the fundamental repeating unit of chromatin [15]. A string of nudeosomes can fold into a higher order structure, the exact molecular nature of which is still not fully understood but clearly has a strong influence on gene expression. 

Physical Structure of the Bacterial Chromosome The Genetic Map of Escherichia coli Eukaryotic DNA Is Complexed with Histones Organization of Genes within Eukaryotic... 

Transcriptional control in eukaryotes can be accomplished at several levels. Chromatin structure can control transcription. The formation of so-called hypersensitive sites (sites where the DNA is not bound into nucleosomes) allows protein factors and RNA polymerase to access the DNA. This is necessary for transcription to occur, but hypersensitive sites are not enough. The removal of histone HI allows transcription to occur from a chromatin domain. Some protein factors (for example, TBF) may be bound to a promoter region even if the gene is not being transcribed. TBF also is necessary but not sufficient for transcription. 

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