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Reprogramming of mouse somatic cells into induced pluripotent stem cells (iPSCs) often generates partially reprogrammed iPSCs (pre-iPSCs), low-grade chimera forming iPSCs (lg-iPSCs) and fully reprogrammed, high-grade chimera production competent iPSCs (hg-iPSCs). Lg-iPSC transcriptome analysis revealed misregulated  Dlk1-Dio3  cluster gene expression and subsequently the imprinting defect at the  Dlk1-Dio3  locus. Here, we show that germ-cell marker  Dppa3  is present only in lg-iPSCs and hg-iPSCs, and that induction with exogenous  Dppa3  enhances reprogramming kinetics, generating all hg-iPSCs, similar to vitamin C (Vc). Conversely,  Dppa3 -null fibroblasts show reprogramming block at pre-iPSCs state and  Dlk1-Dio3  imprinting defect. At the molecular level, we show that Dppa3 is associated with  Dlk1-Dio3  locus and identify that Dppa3 maintains imprinting by antagonizing Dnmt3a binding. Our results further show molecular parallels between Dppa3 and Vc in  Dlk1-Dio3  imprinting maintenance and suggest that early activation of  Dppa3  is one of the cascades through which Vc facilitates the generation of fully reprogrammed iPSCs.

Dppa3 expression is critical for generation of fully reprogrammed iPS cells and maintenance of Dlk1-Dio3 imprinting