SMAD Binding Resource

A curated collection of public datasets on SMAD binding, prepared as a resource for meta-analysis of signal-dependent transcription factor datasets.

The current focus is on the TGF-β family and SMAD signaling pathway. For signal-dependent transcription factors, interpretation requires more than listing binding sites. Binding and transcriptional responses need to be considered together with stimulation time course, cell type, and experimental context. Each dataset has therefore been examined carefully, with attention to features such as co-localizing factors and possible feed-forward mechanisms.

This page was prepared as a practical entry point for inspecting and comparing these datasets in the UCSC Genome Browser, and for organizing input data for future interpretation-support studies.


This table was originally prepared for the review article by Morikawa et al. (Oncogene, 2013; doi: 10.1038/onc.2012.191) and was later updated for the website of the Miyazono Laboratory, Department of Molecular Pathology, The University of Tokyo.

Masato Morikawa, E-mail: address



ChIP-chip, ChIP-seq, CUT&Tag, CUT&RUN

UCSC Genome Browser links are shown for entries with reprocessed bigWig tracks.

HaCaT keratinocytes
SMAD2/3 ChIP-chip, TGF-β3, 1.5 h GSE11710 Koinuma et al, Mol Cell Biol, 2009,
doi: 10.1128/MCB.01038-08
SMAD2 ChIP-seq, TGF-β3, 1.5 h, rep1
SMAD2 ChIP-seq, TGF-β3, 1.5 h, rep2
SMAD3 ChIP-seq, TGF-β3, 1.5 h, rep1
SMAD3 ChIP-seq, TGF-β3, 1.5 h, rep2
GSE180252 Ogami et al, J Biochem, 2022,
doi: 10.1093/jb/mvab141
p63 ChIP-seq, LacZ
p63 ChIP-seq, TGF-β3, 1.5 h/LacZ
p63 ChIP-seq, TGF-β3, 1.5 h/Ras(G12V)
p63 ChIP-seq, TGF-β3, 1.5 h/Ras(N116Y)
GSE60814 Vasilaki et al, Sci Signal, 2016,
doi: 10.1126/scisignal.aag3232
Differentiated human progenitor epidermal keratinocytes (HPEK)
SMAD3 ChIP-seq, TGF-β3, 1.5 h GSE180252 Ogami et al, J Biochem, 2022,
doi: 10.1093/jb/mvab141
HepG2
SMAD2/3 ChIP-chip, TGF-β3, 1.5 h GSE28798 Mizutani et al, J Biol Chem, 2011,
doi: 10.1074/jbc.M110.217745
Hep3B
SMAD2/3 ChIP-chip, TGF-β3, 1.5 h GSE28798 Mizutani et al, J Biol Chem, 2011,
doi: 10.1074/jbc.M110.217745
Human intestinal organoids
SMAD3 CUT&RUN, APC-KO/ARID1A-KO
SMAD3 CUT&RUN, APC-KO/ARID1A-KO, TGF-β1, 30 min
SMAD3 CUT&RUN, APC-KO/SMARCA4-KO
SMAD3 CUT&RUN, APC-KO/SMARCA4-KO, TGF-β1, 30 min
SMAD3 CUT&RUN, APC-KO
SMAD3 CUT&RUN, APC-KO, TGF-β1, 30 min
GSE145183 Ringel et al, Cell Stem Cell, 2020,
doi: 10.1016/j.stem.2020.02.007
MCF10A MII (Human breast epithelial cell line)
SMAD2/3 ChIP-seq, TGF-β3, 1.5 h
SMAD2/3 ChIP-seq, TGF-β3, 16 h
GSE83787 Sundqvist et al, Nucleic Acids Res, 2018,
doi: 10.1093/nar/gkx1190
Hs578T (Human breast cancer cell line, classified as triple-negative)
SMAD2/3 ChIP-seq, TGF-β3, 1.5 h
SMAD2/3 ChIP-seq, TGF-β3, 16 h
GSE104352 Sundqvist et al, Nucleic Acids Res, 2018,
doi: 10.1093/nar/gkx1190
SMAD2 ChIP-seq, Activin A, 1.5 h, rep1
SMAD2 ChIP-seq, Activin A, 1.5 h, rep2
SMAD2 ChIP-seq, Activin A, 1.5 h/Palbociclib, rep1
SMAD2 ChIP-seq, Activin A, 1.5 h/Palbociclib, rep2
GSE117502 Harada et al, Cancer Sci, 2019,
doi: 10.1111/cas.13841
BT549 (Human breast cancer cell line, classified as triple-negative)
SMAD2/3 ChIP-seq, TGF-β3, 1.5 h
SMAD2/3 ChIP-seq, TGF-β3, 16 h
GSE104352 Sundqvist et al, Nucleic Acids Res, 2018,
doi: 10.1093/nar/gkx1190
T47D (Human breast cancer cell line, classified as Luminal A)
SMAD2 ChIP-seq, Activin A, 1.5 h, rep1
SMAD2 ChIP-seq, Activin A, 1.5 h, rep2
SMAD2 ChIP-seq, Activin A, 1.5 h/Palbociclib, rep1
SMAD2 ChIP-seq, Activin A, 1.5 h/Palbociclib, rep2
GSE117502 Harada et al, Cancer Sci, 2019,
doi: 10.1111/cas.13841
A549 (Human lung adenocarcinoma)
SMAD3 ChIP-seq, TGF-β3, 1.5h GSE51510 Isogaya et al, Cell Res, 2014,
doi: 10.1038/cr.2014.97
NCI-H441 (Human lung adenocarcinoma)
SMAD3 ChIP-seq, TGF-β3, 1.5 h/siCtrl
SMAD3 ChIP-seq, TGF-β3, 1.5 h/siTTF1
SMAD4 ChIP-seq, TGF-β3, 1.5 h/siCtrl
SMAD4 ChIP-seq, TGF-β3, 1.5 h/siTTF1
TTF1 ChIP-seq
TTF1 ChIP-seq, TGF-β3, 1.5 h
GSE51510 Isogaya et al, Cell Res, 2014,
doi: 10.1038/cr.2014.97
NCI-H345-TbRII
SMAD2/3 ChIP-seq* GSE63871 Murai et al, Cell Discov, 2015,
doi: 10.1038/celldisc.2015.26
Human umbilical vein endothelial cells (HUVECs)
SMAD1/5 ChIP-seq, BMP-6, 1.5 h
SMAD1/5 ChIP-seq, BMP-9, 1.5 h
GSE27661 Morikawa et al, Nucleic Acids Res, 2011,
doi: 10.1093/nar/gkr572
Human pulmonary arterial endothelial cells (HPAECs)
SMAD1/5 ChIP-seq, BMP-9, 1.5 h, rep1
SMAD1/5 ChIP-seq, BMP-9, 1.5 h, rep2
FLAG-ATOH8 ChIP-seq, rep1
FLAG-ATOH8 ChIP-seq, rep2
GSE104685 Morikawa et al, Sci Signal, 2019,
doi: 10.1126/scisignal.aay4430
Human pulmonary arterial smooth muscle cells (PASMCs)
SMAD1/5 ChIP-seq, BMP-4, 1.5 h GSE27661 Morikawa et al, Nucleic Acids Res, 2011,
doi: 10.1093/nar/gkr572
C2C12 myotube
SMAD3 ChIP-seq, TGF-β1, 2 h, rep1
SMAD3 ChIP-seq, TGF-β1, 2 h, rep2
GSE21614 Mullen et al, Cell, 2011,
doi: 10.1016/j.cell.2011.08.050
Mouse molar tooth germ (dental mesenchyme)
pSMAD1/5 CUT&RUN, dental mesenchyme, rep1
pSMAD1/5 CUT&RUN, dental mesenchyme, rep2
pSMAD3 CUT&RUN, dental mesenchyme, rep1
pSMAD3 CUT&RUN, dental mesenchyme, rep2
GSE297033 Tang et al, Development, 2025,
doi: 10.1242/dev.204563
Human embryonic stem cells (hESCs)
SMAD2/3 ChIP-seq, Ab1
SMAD2/3 ChIP-seq, Ab2
SMAD3 ChIP-seq
GSE29422 Kim et al, Dev Biol, 2011,
doi: 10.1016/j.ydbio.2011.06.009
Mouse embryonic stem cells (mESCs)
SMAD1/5 ChIP-seq, naive mESC/serum
SMAD1/5 ChIP-seq, naive mESC/BMP-4
SMAD1/5 ChIP-seq, primed EpiLC/BMP-4, 1.5 h
SMAD1/5 ChIP-seq, primed EpiLC/BMP-4, 24 h
GSE70581 Morikawa et al, Stem Cell Reports, 2016,
doi: 10.1016/j.stemcr.2015.12.004
KDM1A ChIP-seq, mESC, rep1
KDM1A ChIP-seq, mESC, rep2
KDM1A ChIP-seq, SMAD1/5-dKO mESC, rep1
KDM1A ChIP-seq, SMAD1/5-dKO mESC, rep2
GSE268911 Morikawa et al, J Biol Chem, 2025,
doi: 10.1016/j.jbc.2025.110591
SMAD2/3 ChIP-seq, mESC GSE60286 Yoon et al, Nat Commun, 2015,
doi: 10.1038/ncomms7546
SMAD2/3 ChIP-seq, EB day 3 GSE118765 Su et al, Nature, 2020,
doi: 10.1038/s41586-019-1897-5
SMAD2/3 ChIP-seq, SMAD4-KO mESC
SMAD2/3 ChIP-seq, SMAD4-KO EB, SB431542
SMAD2/3 ChIP-seq, SMAD4-KO EB + Activin A 2 h
SMAD2/3 CUT&Tag, SMAD4-KO EB + Activin A
SMAD2/3 CUT&Tag, SMAD4-KO/Dnmt3b-KD EB + Activin A
SMAD2/3 CUT&Tag, WT mesendoderm + Activin A
SMAD2/3 CUT&Tag, SMAD4-KO mesendoderm + Activin A
GSE240786 /
GSE240787
Zhao et al, Nat Commun, 2024,
doi: 10.1038/s41467-024-54433-5
  • Public ChIP-seq, CUT&RUN, and CUT&Tag datasets were reprocessed.
  • Reads were downloaded with prefetch, converted with fasterq-dump, and aligned to hg38 or mm10 with Bowtie2.
  • BAM files were sorted, indexed, deduplicated, and filtered with SAMtools.
  • CPM-normalized bigWig files were generated with bamCoverage after blacklist filtering.
  • Peaks were called with MACS3 at q = 0.01, using matched controls when available.
  • Peak and fold-enrichment tracks were further processed with BEDTools and UCSC bedGraphToBigWig.
* Peaks were called with the ChIP data alone, without control sample(s).

Microarray

Expression datasets related to the signaling conditions and downstream responses shown above.

HaCaT keratinocytes
TGF-β3 (3 ng/ml, 1.5 h, 6 h, 24 h)
si-SMAD2/3, si-ETS1, si-TFAP2A
GSE11710 Koinuma et al, Mol Cell Biol, 2009,
doi: 10.1128/MCB.01038-08
TGF-β1 (5 ng/ml, 24 h)
Ras(G12V), Ras(N116Y), si-p53
GSE60814 Vasilaki et al, Sci Signal, 2016,
doi: 10.1126/scisignal.aag3232
HepG2
TGF-β3 (3 ng/ml, 1.5 h, 24 h)
si-HNF4α
GSE28798 Mizutani et al, J Biol Chem, 2011,
doi: 10.1074/jbc.M110.217745
NCI-H441
TGF-β3 (3 ng/ml, 1.5 h, 24 h)
si-TTF1
GSE51510 Isogaya et al, Cell Res, 2014,
doi: 10.1038/cr.2014.97
Human umbilical vein endothelial cells (HUVEC)
BMP-6 (50 ng/ml, 2 h, 24 h) / BMP-9 (1 ng/ml, 2 h, 24 h) GSE27661 Morikawa et al, Nucleic Acids Res, 2011,
doi: 10.1093/nar/gkr572
Human pulmonary arterial smooth muscle cells (PASMC)
BMP-4 (50 ng/ml, 2 h, 24 h) GSE27661 Morikawa et al, Nucleic Acids Res, 2011,
doi: 10.1093/nar/gkr572
C2C12 myotube
TGF-β1 (2.5 ng/ml, 12 h) GSE21608 Mullen et al, Cell, 2011,
doi: 10.1016/j.cell.2011.08.050

RNA-seq

Selected transcriptome datasets matched or related to the binding datasets shown above.

Human progenitor epidermal keratinocytes (HPEK)
TGF-β3 (1 ng/ml, 24 h)
differentiated and undifferentiated HPEK cells
GSE180251 Ogami et al, J Biochem, 2022,
doi: 10.1093/jb/mvab141
Human intestinal organoids
TGF-β1 (10 ng/ml, 24 h) without A83-01
APC-KO, APC-KO/ARID1A-KO, APC-KO/SMARCA4-KO
GSE145308 Ringel et al, Cell Stem Cell, 2020,
doi: 10.1016/j.stem.2020.02.007
MCF10A MII (Human breast epithelial cell line)
TGF-β3 (5 ng/ml, 1.5 h, 16 h)
si-JUNB
GSE83786 Sundqvist et al, Nucleic Acids Res, 2018,
doi: 10.1093/nar/gkx1190
A549 (Human lung adenocarcinoma)
TGF-β3 (1 ng/ml, 24 h)
Ad-LacZ, Ad-TTF1
GSE51510 Isogaya et al, Cell Res, 2014,
doi: 10.1038/cr.2014.97
NCI-H441 (Human lung adenocarcinoma)
TGF-β3 (3 ng/ml, 1.5 h, 24 h)
si-TTF1
GSE51510 Isogaya et al, Cell Res, 2014,
doi: 10.1038/cr.2014.97
T47D (Human breast cancer cell line, classified as Luminal A)
Activin A (50 ng/ml, 24 h)
Palbociclib (CDK4/6 inhibitor)
GSE117502 Harada et al, Cancer Sci, 2019,
doi: 10.1111/cas.13841
Mouse molar tooth germ (dental mesenchyme)
SB431542, SIS3 (pSMAD3 inhibitor) GSE297137 Tang et al, Development, 2025,
doi: 10.1242/dev.204563
Mouse embryonic stem cells (mESCs)
LIF+serum, LIF+BMP-4 (10 ng/ml) GSE70581 Morikawa et al, Stem Cell Reports, 2016,
doi: 10.1016/j.stemcr.2015.12.004