CTCF also regulates transcription of certain imprinted genes, such as paternally imprinted Igf2 and H19. Expression and methylation status of many imprinted genes are affected by developmental alcohol exposure in both humans and rodent models (Laufer et al., 2017; Laufer et al., 2015; Laufer et al., 2013; Portales-Casamar et al., 2016), though additional work is needed to determine the full impact of developmental stress exposure. There is some evidence of overlap, as Igf2 is differentially methylated and downregulated in rodent models of prenatal alcohol exposure and in children with FASD (Downing et al., 2011; Laufer et al., 2015; Laufer et al., 2013) and decreased methylation of IGF2 was also found in individuals that had been exposed to famine conditions during periconception (Heijmans et al., 2008). Maternal diet has been shown to also alter methylation of imprinted genes in rodent models (Dunn and Bale, 2011; Zhang et al., 2010).
