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expression fragment b-globin level locus globin footprint region bm gene mouse promoter grosveld probe element activity hypersensitive transgene b-globin locus liver deletion apovldl figure control analysis binding result position chicken erythroid transcription effect sequence factor expression level erythroid cells mel cells tissue acids research core fragment nf-e 2 b-globin gene interaction chromatin number hss 4 pac 1 oligo property activation insulator binding sites hypersensitive sites addition position effects hypersensitive site b gene protein philipsen function assay /3- dnase border adult oligonucleotide chicken b-globin locus /3- globin gene chromatin structure / biochimica panel nature e-globin gene bamhi research 1-6 hss 2 protection g-globin dillon transcriptional