chr5 : 35,317,354 35,318,124
770 bp 100 TFs 2 linked genes
This 770 bp open chromatin element is linked to PRLR and SPEF2 and is bound by 100 transcription factors.
Linked Genes
2 genes
Gene Expression Dist. to TSS Distance Link type
PRLR 87.3 kb Distal Multiome
SPEF2 300.1 kb Distal Multiome
Chromatin Accessibility

Genome Browser

Genomic view centred on this element. Tracks include ATAC-seq accessibility, RNA-seq expression, TF binding peaks, and element-to-gene loop arcs across the ESC → DE time course.

View limited to ±500 kb around this element.
For the full data, please see the UCSC track linked below.
chr5:35,312,354 – 35,323,124
This element Other elements Gene links ATAC-seq RNA-seq

Transcription factors with ChIP-seq peak overlap or TOBIAS motif footprint evidence at this element.

TF Binding
100 transcription factors
Source
Cell type
ASCL1 3 datasets
Motif DE_48h DE_48h-ASCL1_MA1631.2 9 bp overlap
Motif DE_60h DE_60h-ASCL1_MA1631.2 9 bp overlap
Motif DE_72h DE_72h-ASCL1_MA1631.2 9 bp overlap
ATF7 1 dataset
ChIP GM12878 ENCSR014YCR.ATF7.GM12878 136 bp overlap
BRD3 1 dataset
ChIP H-1_DE GSE126661.BRD3.H-1_DE 272 bp overlap
BRD4 1 dataset
ChIP HCC1395 GSE63581.BRD4.HCC1395 98 bp overlap
CRY1 2 datasets
ChIP U2OS GSE130602.CRY1.U2OS 278 bp overlap
ChIP U2OS_DMSO GSE130506.CRY1.U2OS_DMSO 277 bp overlap
Cebpa 1 dataset
ChIP BLaER1 ENCFF896HSY 251 bp overlap
Crx 3 datasets
Motif DE_36h DE_36h-Crx_MA0467.3 6 bp overlap
Motif DE_48h DE_48h-Crx_MA0467.3 6 bp overlap
Motif DE_60h DE_60h-Crx_MA0467.3 6 bp overlap
DPF2 2 datasets
ChIP GM12878 ENCFF681AJV 75 bp overlap
ChIP GM12878 ENCSR509FWH.DPF2.GM12878 69 bp overlap
Dmbx1 3 datasets
Motif DE_36h DE_36h-Dmbx1_MA0883.2 10 bp overlap
Motif DE_48h DE_48h-Dmbx1_MA0883.2 10 bp overlap
Motif DE_60h DE_60h-Dmbx1_MA0883.2 10 bp overlap
E2F6 1 dataset
Motif DE_48h DE_48h-E2F6_MA0471.3 8 bp overlap
E2F8 1 dataset
Motif DE_48h DE_48h-E2F8_MA0865.3 9 bp overlap
ELF3 1 dataset
Motif DE_48h DE_48h-ELF3_MA0640.3 9 bp overlap
EOMES 1 dataset
ChIP hESC GSE26097.EOMES.hESC 337 bp overlap
ERF::FOXI1 3 datasets
Motif DE_36h DE_36h-ERFFOXI1_MA1935.2 10 bp overlap
Motif DE_48h DE_48h-ERFFOXI1_MA1935.2 10 bp overlap
Motif DE_60h DE_60h-ERFFOXI1_MA1935.2 10 bp overlap
ERF::FOXO1 3 datasets
Motif DE_36h DE_36h-ERFFOXO1_MA1936.2 12 bp overlap
Motif DE_48h DE_48h-ERFFOXO1_MA1936.2 12 bp overlap
Motif DE_60h DE_60h-ERFFOXO1_MA1936.2 12 bp overlap
ESR1 4 datasets
ChIP Ishikawa_C2-3 GSE147141.ESR1.Ishikawa_C2-3 60 bp overlap
ChIP Ishikawa_E2_D538G_clone2 GSE132426.ESR1.Ishikawa_E2_D538G_clone2 128 bp overlap
ChIP MCF-7_estradiol-Dex_4h GSE99626.ESR1.MCF-7_estradiol-Dex_4h 89 bp overlap
ChIP MCF-7_estradiol_4h GSE99626.ESR1.MCF-7_estradiol_4h 73 bp overlap
ETV1 3 datasets
Motif DE_36h DE_36h-ETV1_MA0761.3 9 bp overlap
Motif DE_48h DE_48h-ETV1_MA0761.3 9 bp overlap
Motif DE_60h DE_60h-ETV1_MA0761.3 9 bp overlap
ETV2::FOXI1 3 datasets
Motif DE_36h DE_36h-ETV2FOXI1_MA1942.2 11 bp overlap
Motif DE_48h DE_48h-ETV2FOXI1_MA1942.2 11 bp overlap
Motif DE_60h DE_60h-ETV2FOXI1_MA1942.2 11 bp overlap
ETV5::FOXI1 3 datasets
Motif DE_36h DE_36h-ETV5FOXI1_MA1946.2 12 bp overlap
Motif DE_48h DE_48h-ETV5FOXI1_MA1946.2 12 bp overlap
Motif DE_60h DE_60h-ETV5FOXI1_MA1946.2 12 bp overlap
ETV5::FOXO1 3 datasets
Motif DE_36h DE_36h-ETV5FOXO1_MA1947.2 10 bp overlap
Motif DE_48h DE_48h-ETV5FOXO1_MA1947.2 10 bp overlap
Motif DE_60h DE_60h-ETV5FOXO1_MA1947.2 10 bp overlap
FIGLA 3 datasets
Motif DE_48h DE_48h-FIGLA_MA0820.2 6 bp overlap
Motif DE_60h DE_60h-FIGLA_MA0820.2 6 bp overlap
Motif DE_72h DE_72h-FIGLA_MA0820.2 6 bp overlap
FOXA2 2 datasets
ChIP DE DE-FOXA2-1 346 bp overlap
ChIP DE DE-FOXA2-2 310 bp overlap
FOXJ2::ELF1 3 datasets
Motif DE_36h DE_36h-FOXJ2ELF1_MA1952.2 11 bp overlap
Motif DE_48h DE_48h-FOXJ2ELF1_MA1952.2 11 bp overlap
Motif DE_60h DE_60h-FOXJ2ELF1_MA1952.2 11 bp overlap
FOXO1::ELF1 3 datasets
Motif DE_36h DE_36h-FOXO1ELF1_MA1953.2 13 bp overlap
Motif DE_48h DE_48h-FOXO1ELF1_MA1953.2 13 bp overlap
Motif DE_60h DE_60h-FOXO1ELF1_MA1953.2 13 bp overlap
FOXO1::ELK1 3 datasets
Motif DE_36h DE_36h-FOXO1ELK1_MA1954.2 13 bp overlap
Motif DE_48h DE_48h-FOXO1ELK1_MA1954.2 13 bp overlap
Motif DE_60h DE_60h-FOXO1ELK1_MA1954.2 13 bp overlap
FOXO1::ELK3 3 datasets
Motif DE_36h DE_36h-FOXO1ELK3_MA1955.2 13 bp overlap
Motif DE_48h DE_48h-FOXO1ELK3_MA1955.2 13 bp overlap
Motif DE_60h DE_60h-FOXO1ELK3_MA1955.2 13 bp overlap
GATA4 2 datasets
ChIP DE DE-GATA4-1 528 bp overlap
ChIP DE DE-GATA4-2 770 bp overlap
GATA6 11 datasets
ChIP DE DE-GATA6-1 524 bp overlap
ChIP DE DE-GATA6-2 770 bp overlap
ChIP DE_D1 S09-DE-d1-GATA6-exp1 66 bp overlap
ChIP DE_D1 S09-DE-d1-GATA6-exp1 525 bp overlap
ChIP DE_D1 S14-DE-d1-GATA6-exp1 514 bp overlap
ChIP DE_D1 S41-DE-d1-GATA6-exp2 428 bp overlap
ChIP DE_D2 S55-DE-d2-GATA6-exp2 517 bp overlap
ChIP HUES-8_DE_JNKi GSE109524.GATA6.HUES-8_DE_JNKi 649 bp overlap
ChIP HUES-8_DE_ctrl GSE109524.GATA6.HUES-8_DE_ctrl 57 bp overlap
ChIP HUES-8_DE_ctrl GSE109524.GATA6.HUES-8_DE_ctrl 525 bp overlap
ChIP endoderm_KO GSE117136.GATA6.endoderm_KO 343 bp overlap
GSC 3 datasets
Motif DE_36h DE_36h-GSC_MA0648.2 6 bp overlap
Motif DE_48h DE_48h-GSC_MA0648.2 6 bp overlap
Motif DE_60h DE_60h-GSC_MA0648.2 6 bp overlap
GSC2 3 datasets
Motif DE_36h DE_36h-GSC2_MA0891.2 6 bp overlap
Motif DE_48h DE_48h-GSC2_MA0891.2 6 bp overlap
Motif DE_60h DE_60h-GSC2_MA0891.2 6 bp overlap
HNRNPK 2 datasets
ChIP Hep-G2 ENCSR519QAA.HNRNPK.Hep-G2 193 bp overlap
ChIP Hep-G2 GSE120104.HNRNPK.Hep-G2 325 bp overlap
Hnf1A 3 datasets
Motif DE_48h DE_48h-Hnf1A_MA1991.2 10 bp overlap
Motif DE_60h DE_60h-Hnf1A_MA1991.2 10 bp overlap
Motif DE_72h DE_72h-Hnf1A_MA1991.2 10 bp overlap
IKZF1 3 datasets
Motif DE_36h DE_36h-IKZF1_MA1508.2 8 bp overlap
Motif DE_48h DE_48h-IKZF1_MA1508.2 8 bp overlap
Motif DE_60h DE_60h-IKZF1_MA1508.2 8 bp overlap
IKZF2 5 datasets
Motif DE_36h DE_36h-IKZF2_MA2326.1 6 bp overlap
Motif DE_48h DE_48h-IKZF2_MA2326.1 6 bp overlap
Motif DE_60h DE_60h-IKZF2_MA2326.1 6 bp overlap
ChIP GM12878 ENCFF238LYK 116 bp overlap
ChIP GM12878 ENCFF918AID 53 bp overlap
Ikzf3 1 dataset
Motif DE_48h DE_48h-Ikzf3_MA1992.2 9 bp overlap
JUN 3 datasets
ChIP DE_D1 S40-DE-d1-JUN-exp2 68 bp overlap
ChIP DE_D2 S02-DE-d2-JUN-exp1 409 bp overlap
ChIP HUES-8_DE_ctrl GSE109524.JUN.HUES-8_DE_ctrl 64 bp overlap
KLF1 1 dataset
Motif DE_48h DE_48h-KLF1_MA0493.3 8 bp overlap
KLF2 1 dataset
Motif DE_48h DE_48h-KLF2_MA1515.2 8 bp overlap
KLF4 1 dataset
Motif DE_48h DE_48h-KLF4_MA0039.5 8 bp overlap
KLF5 2 datasets
Motif DE_48h DE_48h-KLF5_MA0599.1 10 bp overlap
Motif DE_48h DE_48h-KLF5_MA0599.1 10 bp overlap
Lef1 3 datasets
Motif DE_48h DE_48h-Lef1_MA0768.3 8 bp overlap
Motif DE_60h DE_60h-Lef1_MA0768.3 8 bp overlap
Motif DE_72h DE_72h-Lef1_MA0768.3 8 bp overlap
MSANTD3 3 datasets
Motif DE_36h DE_36h-MSANTD3_MA1523.2 7 bp overlap
Motif DE_48h DE_48h-MSANTD3_MA1523.2 7 bp overlap
Motif DE_60h DE_60h-MSANTD3_MA1523.2 7 bp overlap
MTA2 1 dataset
ChIP GM12878 ENCSR293QAR.MTA2.GM12878 231 bp overlap
MXI1 2 datasets
Motif DE_48h DE_48h-MXI1_MA1108.3 6 bp overlap
Motif DE_60h DE_60h-MXI1_MA1108.3 6 bp overlap
MYOD1 3 datasets
ChIP RD GSE137168.MYOD1.RD 208 bp overlap
ChIP RH4 GSE83726.MYOD1.RH4 202 bp overlap
ChIP rhabdomyosarcoma GSE50413.MYOD1.rhabdomyosarcoma 156 bp overlap
MZF1 3 datasets
Motif DE_36h DE_36h-MZF1_MA0056.3 8 bp overlap
Motif DE_48h DE_48h-MZF1_MA0056.3 8 bp overlap
Motif DE_60h DE_60h-MZF1_MA0056.3 8 bp overlap
Mafb 3 datasets
Motif DE_48h DE_48h-Mafb_MA0117.3 11 bp overlap
Motif DE_60h DE_60h-Mafb_MA0117.3 11 bp overlap
Motif DE_72h DE_72h-Mafb_MA0117.3 11 bp overlap
NANOG 1 dataset
ChIP WA01_3IL ERP004238.NANOG.WA01_3IL 337 bp overlap
NRL 3 datasets
Motif DE_48h DE_48h-NRL_MA0842.3 12 bp overlap
Motif DE_60h DE_60h-NRL_MA0842.3 12 bp overlap
Motif DE_72h DE_72h-NRL_MA0842.3 12 bp overlap
Nr2F6 1 dataset
Motif DE_48h DE_48h-Nr2F6_MA0728.1 15 bp overlap
OTX1 3 datasets
Motif DE_36h DE_36h-OTX1_MA0711.2 6 bp overlap
Motif DE_48h DE_48h-OTX1_MA0711.2 6 bp overlap
Motif DE_60h DE_60h-OTX1_MA0711.2 6 bp overlap
OTX2 3 datasets
Motif DE_36h DE_36h-OTX2_MA0712.3 7 bp overlap
Motif DE_48h DE_48h-OTX2_MA0712.3 7 bp overlap
Motif DE_60h DE_60h-OTX2_MA0712.3 7 bp overlap
PAX3-FOXO1 2 datasets
ChIP Hs-352-Sk_PAX3-FOXO1-vector GSE83725.PAX3-FOXO1.Hs-352-Sk_PAX3-FOXO1-vector 241 bp overlap
ChIP RH4 GSE140115.PAX3-FOXO1.RH4 166 bp overlap
PITX1 3 datasets
Motif DE_36h DE_36h-PITX1_MA0682.3 6 bp overlap
Motif DE_48h DE_48h-PITX1_MA0682.3 6 bp overlap
Motif DE_60h DE_60h-PITX1_MA0682.3 6 bp overlap
PITX3 3 datasets
Motif DE_36h DE_36h-PITX3_MA0714.2 6 bp overlap
Motif DE_48h DE_48h-PITX3_MA0714.2 6 bp overlap
Motif DE_60h DE_60h-PITX3_MA0714.2 6 bp overlap
RARA 1 dataset
Motif DE_48h DE_48h-RARA_MA0729.1 18 bp overlap
RELA 1 dataset
ChIP FaDu_LPS GSE132018.RELA.FaDu_LPS 203 bp overlap
RHOXF1 3 datasets
Motif DE_36h DE_36h-RHOXF1_MA0719.2 6 bp overlap
Motif DE_48h DE_48h-RHOXF1_MA0719.2 6 bp overlap
Motif DE_60h DE_60h-RHOXF1_MA0719.2 6 bp overlap
SATB1 2 datasets
ChIP MCF-10A_ICRF GSE123292.SATB1.MCF-10A_ICRF 151 bp overlap
ChIP MCF-10A_dHD GSE123292.SATB1.MCF-10A_dHD 184 bp overlap
SKIL 1 dataset
ChIP GM12878 ENCSR212YKD.SKIL.GM12878 115 bp overlap
SMAD2 1 dataset
ChIP endoderm GSE29422.SMAD2.endoderm 242 bp overlap
SMAD2-3 4 datasets
ChIP HUES-8_DE GSE109524.SMAD2-3.HUES-8_DE 638 bp overlap
ChIP HUES-8_DE_JNKi GSE109524.SMAD2-3.HUES-8_DE_JNKi 565 bp overlap
ChIP HUES-8_DE_ctrl GSE109524.SMAD2-3.HUES-8_DE_ctrl 226 bp overlap
ChIP HUES-8_DE_ctrl GSE109524.SMAD2-3.HUES-8_DE_ctrl 277 bp overlap
SMAD2_3 7 datasets
ChIP DE_D1 S07-DE-d1-SMAD2_3-exp1 60 bp overlap
ChIP DE_D1 S07-DE-d1-SMAD2_3-exp1 272 bp overlap
ChIP DE_D1 S12-DE-d1-SMAD2_3-exp1 511 bp overlap
ChIP DE_D1 S39-DE-d1-SMAD2_3-exp2 70 bp overlap
ChIP DE_D2 S01-DE-d2-SMAD2_3-exp1 623 bp overlap
ChIP DE_D2 S53-DE-d2-SMAD2_3-exp2 75 bp overlap
ChIP DE_D2 S53-DE-d2-SMAD2_3-exp2 519 bp overlap
SMAD3 7 datasets
ChIP BG03_DIFF_48H GSE36578.SMAD3.BG03_DIFF_48H 235 bp overlap
ChIP HCC1954 GSE104760.SMAD3.HCC1954 71 bp overlap
ChIP HCC1954_TGFb GSE104760.SMAD3.HCC1954_TGFb 273 bp overlap
ChIP HMLE_Doxycicline_TGFb GSE104760.SMAD3.HMLE_Doxycicline_TGFb 60 bp overlap
ChIP NCI-H441 GSE51509.SMAD3.NCI-H441 404 bp overlap
ChIP NCI-H441_SIITTF1 GSE51509.SMAD3.NCI-H441_SIITTF1 383 bp overlap
ChIP endoderm GSE29422.SMAD3.endoderm 217 bp overlap
SMAD4 1 dataset
ChIP endoderm GSE29422.SMAD4.endoderm 220 bp overlap
SNAI1 3 datasets
Motif DE_48h DE_48h-SNAI1_MA1558.2 7 bp overlap
Motif DE_60h DE_60h-SNAI1_MA1558.2 7 bp overlap
Motif DE_72h DE_72h-SNAI1_MA1558.2 7 bp overlap
SNAI2 1 dataset
ChIP RD GSE137168.SNAI2.RD 219 bp overlap
SOX17 1 dataset
ChIP DE_D2 DED2-SOX17_Batch_II 431 bp overlap
SOX17_M 1 dataset
ChIP DE_D2 DED2-SOX17-M_Batch_II 550 bp overlap
SP5 3 datasets
Motif DE_48h DE_48h-SP5_MA1965.2 6 bp overlap
Motif DE_48h DE_48h-SP5_MA1965.2 6 bp overlap
Motif DE_60h DE_60h-SP5_MA1965.2 6 bp overlap
SREBF2 1 dataset
Motif DE_48h DE_48h-SREBF2_MA0596.1 10 bp overlap
Spz1 1 dataset
Motif DE_48h DE_48h-Spz1_MA0111.1 11 bp overlap
TBX21 1 dataset
ChIP GM12878 ENCSR739IHN.TBX21.GM12878 269 bp overlap
TCF12 3 datasets
Motif DE_48h DE_48h-TCF12_MA1648.2 7 bp overlap
Motif DE_60h DE_60h-TCF12_MA1648.2 7 bp overlap
Motif DE_72h DE_72h-TCF12_MA1648.2 7 bp overlap
TCF3 3 datasets
Motif DE_48h DE_48h-TCF3_MA0522.4 7 bp overlap
Motif DE_60h DE_60h-TCF3_MA0522.4 7 bp overlap
Motif DE_72h DE_72h-TCF3_MA0522.4 7 bp overlap
TCF4 3 datasets
Motif DE_48h DE_48h-TCF4_MA0830.3 8 bp overlap
Motif DE_60h DE_60h-TCF4_MA0830.3 8 bp overlap
Motif DE_72h DE_72h-TCF4_MA0830.3 8 bp overlap
TCF7 3 datasets
Motif DE_48h DE_48h-TCF7_MA0769.3 7 bp overlap
Motif DE_60h DE_60h-TCF7_MA0769.3 7 bp overlap
Motif DE_72h DE_72h-TCF7_MA0769.3 7 bp overlap
Tbx6 1 dataset
Motif DE_48h DE_48h-Tbx6_MA1567.3 9 bp overlap
ZBTB24 1 dataset
Motif DE_48h DE_48h-ZBTB24_MA2330.1 10 bp overlap
ZEB1 3 datasets
Motif DE_48h DE_48h-ZEB1_MA0103.4 6 bp overlap
Motif DE_60h DE_60h-ZEB1_MA0103.4 6 bp overlap
Motif DE_72h DE_72h-ZEB1_MA0103.4 6 bp overlap
ZNF143 1 dataset
ChIP FLP143HA_T4 GSE39263.ZNF143.FLP143HA_T4 145 bp overlap
ZNF157 3 datasets
Motif DE_48h DE_48h-ZNF157_MA2331.1 21 bp overlap
Motif DE_60h DE_60h-ZNF157_MA2331.1 21 bp overlap
Motif DE_72h DE_72h-ZNF157_MA2331.1 21 bp overlap
ZNF175 3 datasets
Motif DE_36h DE_36h-ZNF175_MA2332.1 9 bp overlap
Motif DE_48h DE_48h-ZNF175_MA2332.1 9 bp overlap
Motif DE_60h DE_60h-ZNF175_MA2332.1 9 bp overlap
ZNF189 6 datasets
Motif DE_36h DE_36h-ZNF189_MA1725.2 9 bp overlap
Motif DE_48h DE_48h-ZNF189_MA1725.2 9 bp overlap
Motif DE_48h DE_48h-ZNF189_MA1725.2 9 bp overlap
Motif DE_60h DE_60h-ZNF189_MA1725.2 9 bp overlap
Motif DE_60h DE_60h-ZNF189_MA1725.2 9 bp overlap
Motif DE_72h DE_72h-ZNF189_MA1725.2 9 bp overlap
ZNF19 1 dataset
ChIP HEK293T GSE78099.ZNF19.HEK293T 333 bp overlap
ZNF24 3 datasets
Motif DE_36h DE_36h-ZNF24_MA1124.1 13 bp overlap
Motif DE_48h DE_48h-ZNF24_MA1124.1 13 bp overlap
Motif DE_60h DE_60h-ZNF24_MA1124.1 13 bp overlap
ZNF274 3 datasets
Motif DE_36h DE_36h-ZNF274_MA1592.2 12 bp overlap
Motif DE_48h DE_48h-ZNF274_MA1592.2 12 bp overlap
Motif DE_60h DE_60h-ZNF274_MA1592.2 12 bp overlap
ZNF317 3 datasets
Motif DE_48h DE_48h-ZNF317_MA1593.2 8 bp overlap
Motif DE_60h DE_60h-ZNF317_MA1593.2 8 bp overlap
Motif DE_72h DE_72h-ZNF317_MA1593.2 8 bp overlap
ZNF341 3 datasets
Motif DE_36h DE_36h-ZNF341_MA1655.2 8 bp overlap
Motif DE_48h DE_48h-ZNF341_MA1655.2 8 bp overlap
Motif DE_60h DE_60h-ZNF341_MA1655.2 8 bp overlap
ZNF549 1 dataset
Motif DE_48h DE_48h-ZNF549_MA1728.2 8 bp overlap
ZNF677 3 datasets
Motif DE_36h DE_36h-ZNF677_MA2101.1 12 bp overlap
Motif DE_48h DE_48h-ZNF677_MA2101.1 12 bp overlap
Motif DE_60h DE_60h-ZNF677_MA2101.1 12 bp overlap
ZNF687 1 dataset
ChIP GM12878 ENCFF233SGE 457 bp overlap
ZNF701 3 datasets
Motif DE_36h DE_36h-ZNF701_MA1987.2 17 bp overlap
Motif DE_48h DE_48h-ZNF701_MA1987.2 17 bp overlap
Motif DE_60h DE_60h-ZNF701_MA1987.2 17 bp overlap
ZNF816 1 dataset
Motif DE_48h DE_48h-ZNF816_MA1719.2 15 bp overlap
Zic1::Zic2 3 datasets
Motif DE_36h DE_36h-Zic1Zic2_MA1628.2 7 bp overlap
Motif DE_48h DE_48h-Zic1Zic2_MA1628.2 7 bp overlap
Motif DE_60h DE_60h-Zic1Zic2_MA1628.2 7 bp overlap
Zic2 3 datasets
Motif DE_36h DE_36h-Zic2_MA1629.2 9 bp overlap
Motif DE_48h DE_48h-Zic2_MA1629.2 9 bp overlap
Motif DE_60h DE_60h-Zic2_MA1629.2 9 bp overlap
Zic3 3 datasets
Motif DE_36h DE_36h-Zic3_MA0697.3 7 bp overlap
Motif DE_48h DE_48h-Zic3_MA0697.3 7 bp overlap
Motif DE_60h DE_60h-Zic3_MA0697.3 7 bp overlap