We further randomly selected 26 and 15 untranscribed (negligible amount of overlapping FANTOM5 HeLa-S3 CAGE tags) 500bp regions centered on mid positions of HeLa-S3 E states and HeLa-S3 ENCODE DHSs. Two literature-derived44 HeLa-S3 positive enhancers and 4 random regions (see ‘Construction of random genomic background regions’) were used for comparison. For comparison, we also randomly selected 20 manually defined untranscribed HeLa-S3 chromatin-defined active enhancers (see ‘Proof of concept analysis’).