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Active DNA demethylation by Gadd45 and DNA repair
Active DNA demethylation by Gadd45
and DNA repair
Christof Niehrs1,2 and Andrea Scha? fer1
1 Institute of Molecular Biology, 55128 Mainz, Germany
2 Division of Molecular Embryology, German Cancer Research Center (DKFZ)–Center for Molecular Biology of the University of
Heidelberg (ZMBH) Alliance, 69120 Heidelberg, Germany
ReviewHow DNA methylation patterns are established, main-
tained and remodeled is incompletely understood, how-
ever, it has become clear that DNA methylation is
reversible and dynamic as a result of enzymatic DNA
demethylation. Several different mechanisms that may
account for demethylation have recently been put for-
ward and all seem to involve DNA repair. Here, we
review DNA demethylation mediated by multifunctional
growth arrest and DNA damage 45 (Gadd45) protein
family members which mediate DNA demethylation
during cell differentiation and stress response. Gadd45
recruits nucleotide and/or base excision repair factors to
gene-specific loci and acts as an adapter between repair
factors and chromatin, thereby creating a nexus be-
tween epigenetics and DNA repair.
Dynamics of DNA methylation
DNA methylation is a common epigenetic mark conserved in
many eukaryotes including protists, fungi, plants and ani-
mals. Eukaryotic DNA methylation is primarily confined to
cytosine bases and is associated with transcriptional silenc-
ing. In animals, DNA methylation predominantly occurs at
CpG dinucleotides, but in embryonic stem cells it is also
found in non-CpG contexts [1–3]. In many promoters the
presence of 5-methylcytosine (5mC) promotes gene silenc-
ing, and hence DNA methylation is implicated in imprint-
ing, X chromosome inactivation, embryonic development,
tissue-specific gene expression, and cancer (reviewed in [4–
6]). More recently, oxidized derivatives of 5mC, including 5-
hydroxymethylcytosine, 5-formylcytosine and 5-carboxylcy-
tosine, have been recognized in a variety of mammalian cells
[7–11].
Methylation patterns are mitotically and meiot
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