Collectively, the nine mutations have a graded impact and can act asCTCF perturbations, providing an elegant system with which to investigate, (i) therelationship between WT and mutant CTCF and chromatin, and (ii) site specific mutant effectson accessibility, TF binding, cohesin overlap, chromatin interactivity and gene expression,both of which offer insight into CTCF’s biology and its cancer and brain relatedeffects. Here, CTCF is linked to cancer.