Although CM germ cells exhibit only 25% of CTCF WT expression, chromatin binding of CTCF is preferentially lost from CTCF-BORIS heterodimeric sites. Testes with combined depletion of both CTCF and BORIS show reduced size, defective meiotic recombination, increased apoptosis, and malformed spermatozoa. Whereas Ctcf +/+ Boris +/+, Ctcf +/− Boris +/+, and Ctcf +/+ Boris −/− males are fertile, Ctcf +/− Boris −/− (Compound Mutant CM) males are sterile. To understand the CTCF-BORIS relationship, mouse models with varied CTCF and BORIS levels were generated. Unexpectedly, BORIS-null mice have only minimal germ cell defects. Its specialized paralog, BORIS, heterodimerizes with CTCF but is expressed only in male germ cells and in cancer states. CTCF is a key organizer of the 3D genome.
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