Affinage

SMAD1

SMAD family member 1 · UniProt Q15797

Length
465 aa
Mass
52.3 kDa
Annotated
2026-06-10
100 papers in source corpus 34 papers cited in narrative 34 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SMAD1 is a receptor-activated transcription factor that transduces BMP signals into nuclear transcriptional programs governing skeletal, hematopoietic, vascular, neuronal, and iron-homeostatic processes (PMID:8653785, PMID:28438754, PMID:38632412). BMP2 stimulation, but not TGF-β or activin, rapidly induces SMAD1 phosphorylation that is required for activity and drives its nuclear accumulation (PMID:8653785); a parallel non-canonical route operates downstream of TGF-β, in which TGFBR1 phosphorylates and activates ACVR1, which in turn phosphorylates SMAD1/5, accounting for a substantial fraction of the TGF-β transcriptome and being essential for ID1 induction and EMT (PMID:29376829). Once activated, SMAD1 engages DNA through an MH1 domain whose N-terminal helix adopts a domain-swapped conformation that lowers intrinsic affinity but increases cooperativity on palindromic Smad binding elements (PMID:20147459), and it recruits the coactivators p300/CBP in a phosphorylation-enhanced manner (PMID:10673036). Transcriptional specificity is achieved through cell-type-restricted DNA-binding partners and cofactors—Hoxc-8, GATA factors, PU.1, and Runx2—that direct SMAD1 to context-specific target genes (PMID:10224145, PMID:12944489, PMID:21571370, PMID:23387849), and SMAD1 both activates targets such as CCN2/CTGF, Wif1, and Id genes and represses others such as SOX2 via H3K27me3 and DNA methylation (PMID:18668566, PMID:21270055, PMID:25010525, PMID:35905726). Signal output is tuned by multiple negative regulators: sequential MAPK then GSK3 linker phosphorylation routes SMAD1 to centrosomal proteasomal degradation (PMID:18590818), calmodulin binding modulates linker phosphorylation (PMID:11007779), and phosphatase balance (PPM1A, SCP1/2 counteracted by DUSP5) and m6A mRNA methylation/demethylation (METTL14/IGF2BP1, ALKBH5) further set SMAD1 levels (PMID:22588298, PMID:34169608, PMID:36319624, PMID:35724807). Upstream, SMAD1 activation is gated at the receptor and membrane level by CD44, SnoN, endoglin, SANE, and shear-stress–responsive KLF2-BMPER and integrin inputs (PMID:12393873, PMID:15452148, PMID:24019535, PMID:21344387, PMID:39196179), and SMAD1 is integrated with the DNA-damage response through ATM-mediated Ser239 phosphorylation that stabilizes p53 (PMID:22588298). Physiologically, SMAD1/5 redundantly control hepcidin-mediated iron regulation (PMID:28438754), and SMAD1 in parvalbumin interneurons maintains cortical excitation-inhibition balance, its loss causing epileptic seizures (PMID:38632412).

Mechanistic history

Synthesis pass · year-by-year structured walk · 27 steps
  1. 1996 High

    Established SMAD1 as a dedicated, phosphorylation-dependent intracellular effector of BMP receptor signaling, distinguishing it from TGF-β/activin responses.

    Evidence Phosphorylation assays, loss-of-function point mutagenesis, and subcellular fractionation comparing BMP2 vs TGF-β vs activin

    PMID:8653785

    Open questions at the time
    • Did not identify the receptor kinase responsible
    • Did not map the phosphoacceptor SSXS motif residues directly
  2. 1999 High

    Defined how phosphorylated SMAD1 achieves transcriptional output by recruiting p300/CBP coactivators and by de-repressing Hoxc-8-occupied promoters, giving the first mechanistic basis for SMAD1-driven gene activation.

    Evidence Yeast two-hybrid, GST pull-down, Co-IP, EMSA, and reporter assays for Hoxc-8 and p300/CBP interactions

    PMID:10224145 PMID:10673036

    Open questions at the time
    • Genome-wide target scope not addressed
    • Stoichiometry of SMAD1-coactivator-partner complexes unresolved
  3. 1999 Medium

    Revealed that SMAD1 activation is not purely receptor-driven but is modulated by RTK-linked Ras/MEK signaling, opening the concept of cross-pathway input on SMAD1 phosphorylation.

    Evidence Dominant-negative Ras and MEK inhibitor PD98059 with endogenous phosphorylation and reporter readouts

    PMID:10208426

    Open questions at the time
    • Whether MEK acts on the C-terminal or linker phosphosites not distinguished
    • Single cell-type context (intestinal epithelium)
  4. 2000 Medium

    Showed calmodulin directly binds SMAD1 and antagonizes ERK-dependent linker phosphorylation, integrating Ca2+ signaling into the regulation of SMAD1 activity.

    Evidence Structure-function binding assays, Xenopus embryo gain/loss-of-function, and ERK2 phosphorylation assays

    PMID:11007779

    Open questions at the time
    • Physiological Ca2+ stimulus that engages this in mammalian cells not defined
    • Mechanism of how calmodulin occludes linker sites structurally unknown
  5. 2002 High

    Identified SANE as a receptor-level inhibitor that binds both SMAD1/5 and BMP type I receptors to block phosphorylation and nuclear translocation, defining negative control upstream of SMAD1.

    Evidence Reciprocal Co-IP, binding-mutant analysis, nuclear translocation, and in vivo Xenopus/bone-formation rescue

    PMID:12393873

    Open questions at the time
    • Endogenous tissue contexts where SANE limits BMP signaling not mapped
    • Structural basis of dual receptor/SMAD binding unresolved
  6. 2003 High

    Demonstrated that combinatorial co-occupancy with GATA factors sensitizes SMAD1 target elements to low BMP doses, showing how partner transcription factors tune the quantitative BMP response.

    Evidence ChIP in living cells plus binding-site mutagenesis and reporter assays on the Smad7 I-BRE

    PMID:12944489

    Open questions at the time
    • Generality across other BMP targets not tested
    • Direct SMAD1-GATA physical contact not structurally defined
  7. 2004 High

    Connected extracellular matrix sensing to SMAD1 activation by showing the CD44 cytoplasmic domain and hyaluronan-CD44 engagement are required for BMP-7-induced SMAD1 phosphorylation and nuclear translocation.

    Evidence Yeast two-hybrid, Co-IP, nuclear translocation, reporter assays, and hyaluronidase treatment

    PMID:15452148

    Open questions at the time
    • How CD44 couples mechanistically to the receptor complex unclear
    • Whether CD44 dependence is cell-type-specific not resolved
  8. 2005 Low

    Provided first evidence for E3-ligase-mediated turnover of SMAD1 via CHIP binding, pointing toward ubiquitin-dependent control of SMAD1 abundance.

    Evidence Surface plasmon resonance binding assay only

    PMID:15708501

    Open questions at the time
    • Single binding method without functional degradation assay
    • No demonstration of in-cell ubiquitination or turnover
  9. 2005 High

    Established a cytoprotective transcriptional role for SMAD1 in cardiomyocytes by linking it to Bcl-xL/β-catenin upregulation and reduced ischemic apoptosis.

    Evidence Cardiac-specific transgenic mice, adenoviral transfer, TUNEL apoptosis, and western blots for survival mediators

    PMID:15911698

    Open questions at the time
    • Whether SMAD1 directly binds Bcl-xL/β-catenin promoters not shown
    • Loss-of-function endogenous requirement not tested
  10. 2006 High

    Extended non-BMP activation of SMAD1 by showing angiotensin II drives SMAD1 phosphorylation through Src kinase to promote collagen IV synthesis, linking SMAD1 to fibrotic signaling.

    Evidence Src inhibitor PP2, dominant-negative Src, and Src siRNA with p-SMAD1 and collagen IV readouts in mesangial cells

    PMID:16767106

    Open questions at the time
    • Whether Src acts on receptor or directly on SMAD1 not resolved
    • Phosphosite targeted by Src not mapped
  11. 2006 Medium

    Showed that pluripotency factor Nanog restrains BMP-driven mesoderm differentiation by binding SMAD1 and blocking coactivator recruitment, defining a stem-cell-specific brake on SMAD1 output.

    Evidence Co-IP, ES cell differentiation, and reporter assays

    PMID:16801560

    Open questions at the time
    • Direct competition with p300/CBP not biochemically demonstrated
    • Interaction interface unmapped
  12. 2008 Medium

    Defined the linker-phosphorylation code (MAPK then GSK3) that targets SMAD1 to centrosomal proteasomal degradation and showed Wnt prolongs BMP signaling by inhibiting GSK3, establishing duration control of SMAD1 signaling.

    Evidence Review synthesizing Xenopus epistasis, phospho-resistant mutants, proteasome inhibition, and centrosome fractionation

    PMID:18590818

    Open questions at the time
    • Review rather than primary data
    • Identity of the centrosomal E3 ligase not specified here
  13. 2010 High

    Provided the structural explanation for SMAD1's distinctive DNA-binding behavior, showing a domain-swapped MH1 conformation that lowers affinity but raises cooperativity on palindromic elements relative to SMAD3.

    Evidence X-ray crystallography of MH1-SBE complex with thermal melting and DNA binding assays

    PMID:20147459

    Open questions at the time
    • Structure of full-length SMAD1 or partner-bound complexes not solved
    • Functional consequence in cells of cooperativity not directly tested
  14. 2011 High

    Identified SnoN and endoglin as endothelial receptor-complex factors that selectively promote SMAD1/5 activation at ALK1 and balance it against SMAD2/3, linking SMAD1 to angiogenesis and fibrosis.

    Evidence Co-IP, direct binding assays, siRNA, SMAD1/5 phosphorylation, and in vivo angiogenesis phenotypes

    PMID:21344387 PMID:24019535

    Open questions at the time
    • How SnoN/endoglin discriminate SMAD1/5 from SMAD2/3 mechanistically unresolved
    • Quantitative contribution to AVM pathology incomplete
  15. 2011 High

    Used conditional knockouts to assign SMAD1 non-redundant developmental roles, including direct Wif1 activation in lung epithelium and biphasic control of hematopoietic progenitor expansion with reciprocal SMAD2 cross-regulation.

    Evidence Lung- and mesoderm-targeted conditional/temporal Smad1 depletion, microarray, ChIP-chip, and Wnt reporter assays

    PMID:21270055 PMID:21515822

    Open questions at the time
    • Why SMAD1 but not SMAD5 activates Wif1 not explained
    • Cross-pathway switch mechanism with SMAD2 incompletely defined
  16. 2012 High

    Placed SMAD1 within the DNA-damage response by showing ATM phosphorylates it at Ser239, disrupting PPM1A binding and enabling SMAD1 to stabilize p53 against MDM2-mediated degradation.

    Evidence ATM kinase assay, S239 mutagenesis, Co-IP of SMAD1-PPM1A and SMAD1-p53, ubiquitination and survival assays

    PMID:22588298

    Open questions at the time
    • Whether nuclear SMAD1 directly contacts MDM2 or acts via p53 only unclear
    • Generality across genotoxic stresses not fully tested
  17. 2012 Medium

    Expanded the partner repertoire to PU.1 and scleraxis/E12, showing SMAD1 cooperates with lineage transcription factors and integrates non-SMAD ERK/JNK input to drive context-specific genes (IL-1β, SMA).

    Evidence Co-IP, ChIP, promoter assays, knockdown, and pathway inhibitors in macrophages and mesangial cells

    PMID:21571370 PMID:22474292

    Open questions at the time
    • Direct co-binding on endogenous promoters not structurally resolved
    • Single-lab, single-context findings
  18. 2013 Medium

    Demonstrated mechanotransduction into SMAD1/5 via oscillatory shear stress through BMPRIB-integrin association and a Shc/FAK/ERK cascade feeding into SMAD1/5-Runx2-mTOR-driven endothelial proliferation.

    Evidence Co-IP, proximity ligation assay, shRNA, kinase-domain deletion, and in vivo lentiviral injection

    PMID:23387849

    Open questions at the time
    • Direct SMAD1-Runx2 contact not biochemically isolated here
    • Relative contribution of SMAD1 vs SMAD5 not separated
  19. 2013 Medium

    Identified KSHV LANA as a viral hijacker of SMAD1 that sustains nuclear pSMAD1 and enhances loading on Id promoters to drive oncogenic transformation, demonstrating disease exploitation of the BMP-SMAD1-Id axis.

    Evidence Co-IP, ChIP on Id promoters, siRNA/chemical inhibition, and in vivo xenograft

    PMID:25010525

    Open questions at the time
    • LANA-SMAD1 binding interface unmapped
    • Whether LANA blocks SMAD1 degradation or phosphatase access unresolved
  20. 2016 High

    Showed YAP physically stabilizes SMAD1 and is required for its BMP2-induced nuclear accumulation and astrocytic differentiation of neural stem cells, linking Hippo pathway output to SMAD1 protein levels.

    Evidence Yap conditional knockouts, Co-IP, western blot, and SMAD1 re-expression rescue

    PMID:27381227

    Open questions at the time
    • Mechanism of YAP-mediated stabilization (transcriptional vs post-translational) not fully separated
    • Direct binding interface unmapped
  21. 2017 High

    Established SMAD1/5 as the redundant transcriptional node through which erythropoietin and erythroferrone suppress hepcidin, placing SMAD1 at the center of systemic iron homeostasis.

    Evidence Hepatocyte-specific Smad1/Smad5 double-knockout mice, EPO/erythroferrone treatment, siRNA, and primary hepatocyte assays

    PMID:28438754

    Open questions at the time
    • How erythroferrone signal reaches SMAD1/5 at the receptor not fully defined
    • Individual SMAD1 vs SMAD5 contribution masked by redundancy
  22. 2018 High

    Discovered a non-canonical TGFBR1→ACVR1 relay that activates SMAD1/5 under TGF-β, revealing that a major share of TGF-β transcriptional and EMT output is SMAD1/5-dependent.

    Evidence Receptor knockdown/knockout, phosphorylation assays, RNA-seq, and EMT functional assays

    PMID:29376829

    Open questions at the time
    • Determinants of when TGF-β engages this relay vs SMAD2/3-only unclear
    • ACVR1 ligand-independence in this context not fully resolved
  23. 2019 Medium

    Defined phosphatase-level control of SMAD1, showing DUSP5 sustains SMAD1 activation by competitively sequestering SCP1/2 to promote osteogenic differentiation.

    Evidence Co-IP with domain mapping, in vitro dephosphorylation assays, osteogenic differentiation, and OVX mouse rescue

    PMID:34169608

    Open questions at the time
    • Direct SCP1/2 dephosphorylation site on SMAD1 not mapped here
    • Single-lab in vivo model
  24. 2022 Medium

    Established m6A mRNA modification as a layer of SMAD1 control, with METTL14/IGF2BP1 stabilizing SMAD1 transcripts to drive osteogenesis and hypoxia-induced ALKBH5 demethylation enhancing SMAD1 translation to promote trophoblast invasion.

    Evidence MeRIP/m6A-seq, RIP, reporter assays, siRNA, and in vivo OVX and trophoblast knockdown models

    PMID:35724807 PMID:36319624

    Open questions at the time
    • Which m6A sites on SMAD1 mRNA are functionally decisive not pinpointed
    • Single-lab findings per context
  25. 2022 Medium

    Revealed SMAD1 as a transcriptional repressor at chromatin, recruiting H3K27me3 and DNA methylation to silence SOX2 in opposition to SMAD3-driven activation, defining a reciprocal epigenetic axis.

    Evidence Expression profiling, ChIP for histone marks, DNA methylation assays, and in vivo tumor model

    PMID:35905726

    Open questions at the time
    • Which chromatin-modifying enzymes SMAD1 recruits not identified
    • Generality of SMAD1 repressive function beyond SOX2 untested
  26. 2024 High

    Defined an essential physiological role for SMAD1 in cortical parvalbumin interneurons, where BMP2-SMAD1 signaling controls glutamatergic synapse and perineuronal net proteins to maintain excitation-inhibition balance, its loss causing epileptic seizures.

    Evidence PV-interneuron-specific Smad1 conditional knockout, electrophysiology, synaptic proteomics, IHC, and in vivo EEG

    PMID:38632412

    Open questions at the time
    • Direct SMAD1 target genes among synapse/PNN proteins not enumerated
    • Whether other neuronal subtypes require SMAD1 untested
  27. 2024 Medium

    Showed high fluid shear stress suppresses endothelial SMAD1/5 via a KLF2-BMPER axis to enable outward vessel remodeling, integrating hemodynamic input into SMAD1 pathway control.

    Evidence High-FSS mouse model, endothelial BMPER conditional knockout, SMAD1/5 phosphorylation, and KLF2 reporter assays

    PMID:39196179

    Open questions at the time
    • Direct effect of BMPER on SMAD1 vs SMAD5 not separated
    • Downstream SMAD1 transcriptional targets in remodeling not defined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the diverse upstream inputs (canonical BMP receptors, the TGFBR1-ACVR1 relay, Src, integrin/shear, ATM, calmodulin) are integrated quantitatively at SMAD1 to specify distinct transcriptional programs across tissues remains unresolved.
  • No unified model linking phosphosite combinations to target-gene selection
  • Genome-wide SMAD1 cistrome across cell types incompletely defined
  • Structural basis of partner-directed activation vs repression unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 6 GO:0003677 DNA binding 3 GO:0060089 molecular transducer activity 2 GO:0140096 catalytic activity, acting on a protein 1
Localization
GO:0005634 nucleus 6 GO:0005829 cytosol 2 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-162582 Signal Transduction 4 R-HSA-74160 Gene expression (Transcription) 4 R-HSA-1266738 Developmental Biology 3 R-HSA-4839726 Chromatin organization 1 R-HSA-73894 DNA Repair 1
Partners
CD44HOXC-8P300/CBPPPM1APU.1RUNX2SMAD4YAP

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 MADR1 (SMAD1) is a downstream mediator of BMP2 receptor signaling: BMP2 (but not TGF-β or activin) rapidly induces SMAD1 phosphorylation, which is required for function (a point mutant that gives a null phenotype in Drosophila is not phosphorylated), and BMP2 treatment results in accumulation of SMAD1 in the nucleus. Phosphorylation assays, point mutagenesis, subcellular fractionation/immunofluorescence in response to BMP2 vs. TGF-β vs. activin Cell High 8653785
1999 SMAD1 interacts with the homeodomain transcription factor Hoxc-8 (identified by yeast two-hybrid, confirmed by pull-down and co-IP in COS-1 cells); purified SMAD1 inhibits Hoxc-8 binding to its DNA element (osteopontin promoter) in a concentration-dependent manner, thereby de-repressing BMP-induced gene transcription. Yeast two-hybrid screen, GST pull-down, co-immunoprecipitation, EMSA/gel-shift assay, luciferase reporter transfection The Journal of Biological Chemistry High 10224145
1999 SMAD1 interacts with the transcriptional coactivators p300 and CBP both in vitro and in vivo; the C-terminal half of SMAD1 contains two interaction domains that bind the C-terminal region of p300/CBP; phosphorylation of SMAD1 enhances binding to CBP and further stimulates SMAD1-dependent transcription. In vitro binding assay, co-immunoprecipitation, transcription reporter assays, phosphorylation-binding correlation Biochimica et Biophysica Acta High 10673036
1999 The Ras/MEK pathway is partially required for TGF-β- and BMP-induced SMAD1 phosphorylation and transcriptional activity in intestinal epithelial cells; either dominant-negative Ras (RasN17) or the MEK inhibitor PD98059 significantly decreased BMP/TGF-β-induced SMAD1 phosphorylation. Stable inducible dominant-negative Ras expression, MEK inhibitor (PD98059), endogenous SMAD1 phosphorylation assay, luciferase reporter Oncogene Medium 10208426
2000 Calmodulin directly binds to two conserved regions in SMAD1 and increases SMAD1 activity in Xenopus embryos; calmodulin binding inhibits subsequent ERK2-dependent linker phosphorylation of SMAD1, revealing cross-talk between Ca2+/calmodulin, RTK/ERK, and BMP-SMAD1 pathways. Binding assays (structure-function), Xenopus embryo gain/loss-of-function, ERK2 phosphorylation assay The Journal of Biological Chemistry Medium 11007779
2002 SANE (Smad1 Antagonistic Effector), a LEM-domain protein, binds to SMAD1/5 and to BMP type I receptors, inhibiting BMP-induced SMAD1 phosphorylation and blocking ligand-dependent nuclear translocation of SMAD1; a SANE mutant that cannot bind SMAD1 fails to inhibit BMP signaling. Co-immunoprecipitation, Xenopus embryo functional assays, mammalian bone-formation model, nuclear translocation assay, SANE-binding mutant analysis The Journal of Biological Chemistry High 12393873
2003 SMAD1 and GATA factors (GATA-1, -5, or -6) assemble together on the intronic BMP-response element (I-BRE) of the Smad7 gene in living cells; GATA-dependent co-occupancy renders I-BRE more responsive to low BMP concentrations than the promoter BRE that relies solely on SMAD1/SMAD4 binding. Chromatin immunoprecipitation (ChIP), luciferase reporter assays, site-directed mutagenesis of Smad1/Smad4 binding sites Molecular and Cellular Biology High 12944489
2004 SMAD1 interacts with the cytoplasmic domain of CD44 (identified by yeast two-hybrid, confirmed by co-IP); full-length CD44 is required for BMP-7-induced SMAD1 phosphorylation and nuclear translocation—overexpression of a truncated CD44 abolished nuclear SMAD1 translocation—and disruption of extracellular hyaluronan-CD44 interactions (by hyaluronidase) inhibited BMP-7-mediated SMAD1 phosphorylation and SMAD4 nuclear translocation. Yeast two-hybrid, co-immunoprecipitation, nuclear translocation assay, luciferase reporter (SBE4-luc), hyaluronidase treatment The Journal of Cell Biology High 15452148
2005 Interaction between SMAD1 and CHIP (an E3 ubiquitin ligase) was demonstrated by surface plasmon resonance, suggesting CHIP may mediate SMAD1 degradation. Surface plasmon resonance (SPR) binding assay Colloids and Surfaces B: Biointerfaces Low 15708501
2005 In cardiomyocytes, SMAD1 signaling mediates BMP2-induced protection against apoptosis during hypoxia-reoxygenation via upregulation of Bcl-xL and β-catenin; transgenic mice with cardiac-specific Smad1 overexpression showed significantly smaller myocardial infarctions and fewer apoptotic cardiomyocyte deaths after ischemia-reperfusion injury. Cardiac-specific transgenic mouse model, adenoviral gene transfer, TUNEL/DNA ladder apoptosis assay, western blot for Bcl-xL, β-catenin, and caspase-3 Circulation High 15911698
2006 Angiotensin II activates SMAD1 through a Src kinase-dependent mechanism in mesangial cells: Ang II induces phosphorylation of Src and SMAD1; Src inhibitor PP2, dominant-negative Src, or Src siRNA all reduce SMAD1 phosphorylation and type IV collagen synthesis. Pharmacological Src inhibitor (PP2), dominant-negative Src overexpression, Src siRNA knockdown, western blot for p-Src and p-Smad1, collagen IV assay Laboratory Investigation High 16767106
2006 Nanog physically interacts with SMAD1 and blocks BMP-induced mesoderm differentiation of mouse ES cells by interfering with the recruitment of coactivators to active Smad transcriptional complexes. Co-immunoprecipitation, ES cell differentiation assays, transcriptional reporter assays Proceedings of the National Academy of Sciences USA Medium 16801560
2007 In SSc fibroblasts, SMAD1 directly binds and activates the CCN2 (CTGF) gene promoter; siRNA-mediated depletion of SMAD1 normalizes CCN2 and collagen production; imatinib mesylate reverses Smad1 pathway activation via blockade of c-Abl. In vitro and in vivo DNA binding assays, siRNA knockdown, luciferase reporter assay, western blot, immunohistochemistry Arthritis and Rheumatism High 18668566
2008 Linker-region sequential phosphorylation of SMAD1 by MAPK and then GSK3 negatively regulates BMP-SMAD1 signaling by directing SMAD1 to the centrosomal region for polyubiquitination and proteasomal degradation; Wnt signaling, via GSK3 inhibition, prolongs the BMP-SMAD1 signal. Xenopus embryo epistasis experiments, phosphorylation-resistant SMAD1 mutants, proteasome inhibitor assays, centrosome fractionation (reviewed/synthesized from multiple primary papers) Current Opinion in Genetics & Development Medium 18590818
2010 Crystal structure of the SMAD1 MH1 domain bound to a palindromic Smad binding element (SBE) reveals that the N-terminal helix 1 of SMAD1 is dislodged from its intramolecular binding site and adopts a domain-swapped arrangement compared to SMAD3, causing helix 2 to kink away and disabling key DNA backbone contacts; this confers lower overall DNA affinity but increased cooperativity on palindromic elements. X-ray crystallography, thermal melting analysis, DNA binding assays Nucleic Acids Research High 20147459
2011 In endothelial cells, SnoN directly binds ALK1 at the plasma membrane and facilitates the interaction between ALK1 and SMAD1/5, enhancing SMAD1/5 phosphorylation; disruption of SnoN-SMAD interaction impairs SMAD1/5 activation, upregulates SMAD2/3 activity, and causes defective angiogenesis and arteriovenous malformations. Co-immunoprecipitation, direct binding assay (SnoN-ALK1 interaction), in vivo embryonic angiogenesis phenotype, SMAD1/5 phosphorylation assay The Journal of Cell Biology High 24019535
2011 Endoglin promotes TGF-β/SMAD1 signaling in SSc fibroblasts: endoglin is constitutively found in complexes with ALK1, and siRNA depletion of endoglin inhibits SMAD1 phosphorylation without affecting pSMAD2/3, reducing collagen and CCN2 levels. Co-immunoprecipitation, siRNA knockdown, western blot for pSMAD1 Journal of Cellular Physiology Medium 21344387
2011 In the developing lung, SMAD1 (but not SMAD5) in epithelial cells transcriptionally activates Wnt inhibitory factor 1 (Wif1), identified by combined cDNA microarray and ChIP-chip; loss of SMAD1-dependent Wif1 activation results in increased Wnt/β-catenin signaling and specific fetal lung abnormalities. Conditional knockout mice (lung epithelial-specific Smad1 or Smad5 deletion), cDNA microarray, ChIP-chip, Wnt/β-catenin reporter assays Development High 21270055
2011 Smad1 plays a biphasic role in hematopoiesis: early Smad1 expression is required for hemangioblast commitment from mesoderm, while later Smad1 expression in FLK1+ mesoderm restricts expansion of hematopoietic progenitors; depletion of Smad1 after hemangioblast commitment expands hematopoietic progenitors and correlates with increased nuclear SMAD2 activity, indicating cross-regulation between BMP and TGF-β SMAD pathways. Conditional/temporal siRNA knockdown in embryoid body cultures, embryonic stem cell differentiation, SMAD2 nuclear activity assay Blood Medium 21515822
2012 On genotoxic stress, ATM kinase phosphorylates BMP-activated SMAD1 in the nucleus at Ser239; this disrupts SMAD1 interaction with phosphatase PPM1A (leading to enhanced SMAD1 activation), and activated SMAD1 then interacts with p53 to inhibit MDM2-mediated p53 ubiquitination and degradation. ATM kinase assay, S239 phosphorylation site mutagenesis, Co-IP (SMAD1-PPM1A, SMAD1-p53), ubiquitination assay, cell proliferation and survival assays, clinical sample analysis Nature Communications High 22588298
2012 In macrophages, BMP-6-induced IL-1β expression requires a physical interaction between SMAD1 and the transcription factor PU.1; both the canonical SMAD pathway and non-SMAD ERK/JNK pathways are required, and cross-talk between them is mediated through PU.1-SMAD1 interaction. Co-immunoprecipitation (PU.1-SMAD1), siRNA knockdown, ERK/JNK pathway inhibitors, luciferase reporter Molecular Immunology Medium 21571370
2012 In mesangial cells, scleraxis physically associates with E12 to bind the SMA promoter E-box; scleraxis also induces BMP4 secretion, which activates SMAD1 to induce smooth muscle α-actin (SMA); Id1, induced by extended AGE treatment, dislodges scleraxis from the SMA promoter, thereby modulating the BMP4-SMAD1-SMA pathway. Co-immunoprecipitation, chromatin immunoprecipitation, promoter binding assays, overexpression/knockdown, ELISA for BMP4 The Journal of Biological Chemistry Medium 22474292
2013 In endothelial cells, oscillatory shear stress (OSS) induces sustained BMPRIB-αvβ3 integrin association; the intracytoplasmic kinase domain of BMPRII mediates this interaction and subsequently activates the Shc/FAK/ERK cascade, leading to SMAD1/5 activation; SMAD1/5 then associates with and activates Runx2, leading to mTOR/p70S6K activation and EC proliferation. Co-immunoprecipitation, in situ proximity ligation assay, shRNA knockdown of SMAD5, kinase domain deletion analysis, in vivo intra-arterial lentiviral injection Journal of Thrombosis and Haemostasis Medium 23387849
2013 KSHV LANA protein interacts with SMAD1 (identified as novel binding partner), sustains BMP-activated pSMAD1 in the nucleus, and enhances SMAD1 loading on Id gene promoters, resulting in Id upregulation that drives KSHV-induced oncogenic transformation; genetic/chemical inhibition of BMP-SMAD1-Id pathway blocks the oncogenic phenotype. Co-immunoprecipitation (LANA-SMAD1), ChIP on Id promoters, genetic inhibition (siRNA), chemical inhibitor, in vivo tumor xenograft PLoS Pathogens Medium 25010525
2016 YAP (yes-associated protein) physically interacts with SMAD1 in neural stem cells and is required for BMP2-induced stabilization and nuclear accumulation of SMAD1; in YAP-deficient NSCs, SMAD1 expression is reduced and astrocytic differentiation is impaired; SMAD1 re-expression in YAP-deficient NSCs partially rescues astrocytic differentiation. Conditional knockout mice (Yap-nestin, Yap-GFAP), co-immunoprecipitation, western blot, rescue by SMAD1 re-expression Development High 27381227
2017 Erythropoietin and erythroferrone suppress hepcidin transcription by targeting SMAD1/5 signaling: EPO treatment reduces liver SMAD1/5 phosphorylation in parallel with hepcidin mRNA reduction in control mice, but fails to suppress hepcidin in hepatocyte-specific Smad1/Smad5 double-knockout mice or SMAD1/5 knockdown cells; SMAD1 and SMAD5 have overlapping (redundant) functions in hepcidin regulation. Hepatocyte-specific conditional double-knockout mice, EPO/erythroferrone treatment, siRNA knockdown, primary hepatocyte cultures, SMAD1/5 phosphorylation assay Blood High 28438754
2018 TGF-β induces SMAD1/5 phosphorylation through a novel two-receptor mechanism: TGFBR1 phosphorylates and activates ACVR1 (a second type I receptor), which then phosphorylates SMAD1/5; approximately a quarter of the TGF-β-induced transcriptome depends on SMAD1/5 signaling; TGF-β-induced epithelial-to-mesenchymal transition requires both SMAD3 and SMAD1/5 pathways, with SMAD1/5 being essential for ID1 induction. Receptor knockdown/knockout, phosphorylation assays, transcriptome analysis (RNA-seq), EMT functional assays eLife High 29376829
2019 SMAD1 is phosphorylated at Ser239 by ATM in the DNA damage response, and this disrupts SMAD1 interaction with phosphatase PPM1A; conversely, DUSP5 (a dual-specificity phosphatase) promotes osteogenic differentiation by competitively binding SCP1/2 (SMAD1 phosphatases) via its linker region, thereby preventing SCP1/2-mediated SMAD1 dephosphorylation and maintaining SMAD1 activation. Co-immunoprecipitation (DUSP5-SCP1/2-SMAD1), domain mapping, in vitro dephosphorylation assays, osteogenic differentiation assays, OVX mouse model rescue by Dusp5 overexpression Stem Cells Medium 34169608
2019 Sema4C (semaphorin 4C) reverse signaling co-immunoprecipitates with TGFBRII and BMPR1, eliciting non-conventional TGF-β/BMP receptor activation and selective SMAD1/5 phosphorylation leading to increased ID1/3 expression and invasive gene reprogramming in cancer cells. Co-immunoprecipitation (Sema4C-TGFBRII-BMPR1), SMAD1/5 phosphorylation assay, gene expression profiling, in vivo metastasis assay Science Signaling Medium 31431542
2022 METTL14-mediated m6A methylation of SMAD1 mRNA promotes its stability; IGF2BP1 is the m6A reader for SMAD1 transcripts; knockdown of METTL14 reduces m6A modification of SMAD1, decreasing SMAD1 protein levels and suppressing osteogenic differentiation. RIP (RNA immunoprecipitation), MeRIP (m6A-seq), luciferase reporter assay, siRNA knockdown, OVX mouse model Cell Death & Disease Medium 36319624
2022 Hypoxia-induced ALKBH5 demethylase translocates from nucleus to cytoplasm and demethylates m6A-modified SMAD1 (and SMAD5) mRNAs, enhancing their translation and promoting trophoblast invasion via upregulation of MMP9 and ITGA1. m6A-seq (m6A-RIP), ALKBH5 subcellular fractionation, knockdown in mouse placenta (trophoblast-specific), western blot for SMAD1/5 protein Biochimica et Biophysica Acta – Molecular Cell Research Medium 35724807
2022 SMAD1 mediates BMP-dependent repression of SOX2 by recruiting histone H3K27me3 and DNA methylation at the SOX2 promoter; conversely, TGF-β/activin activates SOX2 through SMAD3-dependent H3K4me3 recruitment, defining a reciprocal epigenetic regulatory axis controlling anchorage-independent survival. Gene expression profiling, ChIP (H3K27me3, H3K4me3), DNA methylation assay, in vivo intraperitoneal tumor model Cell Reports Medium 35905726
2024 In adult mouse neocortex, BMP2 acts on parvalbumin (PV) interneurons via SMAD1 to control glutamatergic synapse proteins and perineuronal net components; PV-interneuron-specific disruption of BMP2-SMAD1 signaling reduces glutamatergic innervation of PV cells, impairs perineuronal net development and PV cell excitability, disrupts cortical excitation-inhibition balance, and causes spontaneous epileptic seizures. PV interneuron-specific conditional knockout of SMAD1, electrophysiology, immunohistochemistry, proteomics of synapse proteins, in vivo EEG for seizure detection Nature High 38632412
2024 High fluid shear stress suppresses endothelial SMAD1/5 signaling via a KLF2-BMPER axis: high FSS elevates KLF2, which transcriptionally induces BMPER (a BMP pathway inhibitor), thereby de-inhibiting AKT and facilitating outward vessel remodeling; endothelial BMPER deletion impairs blood flow recovery. Surgically induced high-FSS mouse model, endothelial BMPER conditional knockout, SMAD1/5 phosphorylation assay, KLF2 overexpression/knockdown, luciferase reporter Nature Cardiovascular Research Medium 39196179

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1996 MADR1, a MAD-related protein that functions in BMP2 signaling pathways. Cell 628 8653785
2006 Nanog binds to Smad1 and blocks bone morphogenetic protein-induced differentiation of embryonic stem cells. Proceedings of the National Academy of Sciences of the United States of America 202 16801560
2012 Stalk cell phenotype depends on integration of Notch and Smad1/5 signaling cascades. Developmental cell 189 22364862
2018 TGF-β uses a novel mode of receptor activation to phosphorylate SMAD1/5 and induce epithelial-to-mesenchymal transition. eLife 155 29376829
1999 Smad1 interacts with homeobox DNA-binding proteins in bone morphogenetic protein signaling. The Journal of biological chemistry 145 10224145
2006 Dose-dependent Smad1, Smad5 and Smad8 signaling in the early mouse embryo. Developmental biology 130 16765933
2016 YAP stabilizes SMAD1 and promotes BMP2-induced neocortical astrocytic differentiation. Development (Cambridge, England) 101 27381227
1999 Cross-talk between the Smad1 and Ras/MEK signaling pathways for TGFbeta. Oncogene 85 10208426
2022 Downregulation of METTL14 improves postmenopausal osteoporosis via IGF2BP1 dependent posttranscriptional silencing of SMAD1. Cell death & disease 77 36319624
2020 Correcting Smad1/5/8, mTOR, and VEGFR2 treats pathology in hereditary hemorrhagic telangiectasia models. The Journal of clinical investigation 76 31689244
2017 Smad1/5 is required for erythropoietin-mediated suppression of hepcidin in mice. Blood 73 28438754
2003 GATA- and Smad1-dependent enhancers in the Smad7 gene differentially interpret bone morphogenetic protein concentrations. Molecular and cellular biology 73 12944489
2002 SANE, a novel LEM domain protein, regulates bone morphogenetic protein signaling through interaction with Smad1. The Journal of biological chemistry 73 12393873
2021 Endometrial receptivity and implantation require uterine BMP signaling through an ACVR2A-SMAD1/SMAD5 axis. Nature communications 71 34099644
2011 Endoglin promotes TGF-β/Smad1 signaling in scleroderma fibroblasts. Journal of cellular physiology 69 21344387
2006 Angiotensin II-dependent Src and Smad1 signaling pathway is crucial for the development of diabetic nephropathy. Laboratory investigation; a journal of technical methods and pathology 69 16767106
2008 Smad1 pathway is activated in systemic sclerosis fibroblasts and is targeted by imatinib mesylate. Arthritis and rheumatism 68 18668566
2007 Smad1 and Smad5 differentially regulate embryonic hematopoiesis. Blood 66 17761518
2016 Differential expression of TGF-β superfamily members and role of Smad1/5/9-signalling in chondral versus endochondral chondrocyte differentiation. Scientific reports 65 27848974
2008 Integrating positional information at the level of Smad1/5/8. Current opinion in genetics & development 65 18590818
2013 Bone morphogenetic protein 4 promotes mammalian oogonial stem cell differentiation via Smad1/5/8 signaling. Fertility and sterility 64 23993924
2013 ALK1-Smad1/5 signaling pathway in fibrosis development: friend or foe? Cytokine & growth factor reviews 64 24055043
2013 p53 regulates neural stem cell proliferation and differentiation via BMP-Smad1 signaling and Id1. Stem cells and development 63 23199293
2010 Structure of Smad1 MH1/DNA complex reveals distinctive rearrangements of BMP and TGF-beta effectors. Nucleic acids research 62 20147459
2004 CD44 modulates Smad1 activation in the BMP-7 signaling pathway. The Journal of cell biology 62 15452148
2005 Smad1 protects cardiomyocytes from ischemia-reperfusion injury. Circulation 56 15911698
2013 The ALK-1/Smad1 pathway in cardiovascular physiopathology. A new target for therapy? Biochimica et biophysica acta 54 23707512
2011 Smad1 plays an essential role in bone development and postnatal bone formation. Osteoarthritis and cartilage 54 21420501
2013 BMP receptor-integrin interaction mediates responses of vascular endothelial Smad1/5 and proliferation to disturbed flow. Journal of thrombosis and haemostasis : JTH 53 23387849
2007 Functional characterization of bone morphogenetic protein binding sites and Smad1/5 activation in human vascular cells. Molecular pharmacology 53 17989347
1999 Activation of Smad1-mediated transcription by p300/CBP. Biochimica et biophysica acta 50 10673036
2009 Integration of BMP and Wnt signaling via vertebrate Smad1/5/8 and Drosophila Mad. Cytokine & growth factor reviews 48 19896409
2022 Reciprocal SOX2 regulation by SMAD1-SMAD3 is critical for anoikis resistance and metastasis in cancer. Cell reports 47 35905726
2011 Smad1 and its target gene Wif1 coordinate BMP and Wnt signaling activities to regulate fetal lung development. Development (Cambridge, England) 47 21270055
2000 Calmodulin differentially modulates Smad1 and Smad2 signaling. The Journal of biological chemistry 46 11007779
2019 LRG1 Promotes Apoptosis and Autophagy through the TGFβ-smad1/5 Signaling Pathway to Exacerbate Ischemia/Reperfusion Injury. Neuroscience 45 31220542
2012 BMP-9 induced endothelial cell tubule formation and inhibition of migration involves Smad1 driven endothelin-1 production. PloS one 45 22299030
2010 Endothelial cells are activated during hypoxia via endoglin/ALK-1/SMAD1/5 signaling in vivo and in vitro. Biochemical and biophysical research communications 45 20060813
2012 A crucial role for bone morphogenetic protein-Smad1 signalling in the DNA damage response. Nature communications 44 22588298
2021 Circular RNA AFF4 modulates osteogenic differentiation in BM-MSCs by activating SMAD1/5 pathway through miR-135a-5p/FNDC5/Irisin axis. Cell death & disease 41 34145212
2019 Ablation of Hepatocyte Smad1, Smad5, and Smad8 Causes Severe Tissue Iron Loading and Liver Fibrosis in Mice. Hepatology (Baltimore, Md.) 41 31127639
2024 Control of neuronal excitation-inhibition balance by BMP-SMAD1 signalling. Nature 36 38632412
2016 Thyroid follicle development requires Smad1/5- and endothelial cell-dependent basement membrane assembly. Development (Cambridge, England) 36 27068110
2017 Genomic integration of Wnt/β-catenin and BMP/Smad1 signaling coordinates foregut and hindgut transcriptional programs. Development (Cambridge, England) 35 28219948
2016 Sphingosine-1-phosphate/S1PR2-mediated signaling triggers Smad1/5/8 phosphorylation and thereby induces Runx2 expression in osteoblasts. Bone 35 27612439
2015 Bone Morphogenetic Protein 4 and Smad1 Mediate Extracellular Matrix Production in the Development of Diabetic Nephropathy. Diabetes 35 25995358
2020 Impaired SMAD1/5 Mechanotransduction and Cx37 (Connexin37) Expression Enable Pathological Vessel Enlargement and Shunting. Arteriosclerosis, thrombosis, and vascular biology 34 32078368
2019 Adamts17 is involved in skeletogenesis through modulation of BMP-Smad1/5/8 pathway. Cellular and molecular life sciences : CMLS 34 31201465
2020 BMP-SMAD1/5 Signaling Regulates Retinal Vascular Development. Biomolecules 32 32210087
2017 Fstl1 Promotes Glioma Growth Through the BMP4/Smad1/5/8 Signaling Pathway. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 30 29212066
2014 Transcriptional factors smad1 and smad9 act redundantly to mediate zebrafish ventral specification downstream of smad5. The Journal of biological chemistry 30 24488494
1997 Evidence that Mothers-against-dpp-related 1 (Madr1) plays a role in the initiation and maintenance of spermatogenesis in the mouse. Mechanisms of development 30 9076678
2022 BMP-7 ameliorates partial epithelial-mesenchymal transition by restoring SnoN protein level via Smad1/5 pathway in diabetic kidney disease. Cell death & disease 29 35314669
2022 Hypoxia induced ALKBH5 prevents spontaneous abortion by mediating m6A-demethylation of SMAD1/5 mRNAs. Biochimica et biophysica acta. Molecular cell research 29 35724807
2014 Smad1 and 5 but not Smad8 establish stem cell quiescence which is critical to transform the premature hair follicle during morphogenesis toward the postnatal state. Stem cells (Dayton, Ohio) 28 24023003
2011 Role of Smad1 in diabetic nephropathy: Molecular mechanisms and implications as a diagnostic marker. Histology and histopathology 28 21360446
2023 Apoptotic Vesicles Regulate Bone Metabolism via the miR1324/SNX14/SMAD1/5 Signaling Axis. Small (Weinheim an der Bergstrasse, Germany) 27 36670083
2018 SMAD1/5 signaling in osteoclasts regulates bone formation via coupling factors. PloS one 27 30188920
2014 BMP4 enhances foam cell formation by BMPR-2/Smad1/5/8 signaling. International journal of molecular sciences 27 24690996
2014 Oncogenic herpesvirus KSHV Hijacks BMP-Smad1-Id signaling to promote tumorigenesis. PLoS pathogens 27 25010525
2000 Characterization of the mouse Smad1 gene and its expression pattern in adult mouse tissues. Gene 27 11111041
2019 Activation of the BMP4/Smad1 Pathway Promotes Retinal Ganglion Cell Survival and Axon Regeneration. Investigative ophthalmology & visual science 25 31022296
2018 BMP6 Downregulates GDNF Expression Through SMAD1/5 and ERK1/2 Signaling Pathways in Human Granulosa-Lutein Cells. Endocrinology 25 29750278
2022 BMP2 promotes lung adenocarcinoma metastasis through BMP receptor 2-mediated SMAD1/5 activation. Scientific reports 24 36175474
2016 Icariin regulates systemic iron metabolism by increasing hepatic hepcidin expression through Stat3 and Smad1/5/8 signaling. International journal of molecular medicine 24 27035325
2015 MicroRNA-26b inhibits metastasis of osteosarcoma via targeting CTGF and Smad1. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 24 25761878
2005 Smad1 expression and function during mouse embryonic lung branching morphogenesis. American journal of physiology. Lung cellular and molecular physiology 24 15681399
2017 Smad1 promotes colorectal cancer cell migration through Ajuba transactivation. Oncotarget 23 29299158
2015 Smad1/5 and Smad4 expression are important for osteoclast differentiation. Journal of cellular biochemistry 23 25711193
2020 Celastrol attenuates arterial and valvular calcification via inhibiting BMP2/Smad1/5 signalling. Journal of cellular and molecular medicine 22 32954678
2019 Reverse signaling by semaphorin 4C elicits SMAD1/5- and ID1/3-dependent invasive reprogramming in cancer cells. Science signaling 22 31431542
2018 TGF-β and BMP signals regulate insect diapause through Smad1-POU-TFAM pathway. Biochimica et biophysica acta. Molecular cell research 22 29902488
2012 Scleraxis modulates bone morphogenetic protein 4 (BMP4)-Smad1 protein-smooth muscle α-actin (SMA) signal transduction in diabetic nephropathy. The Journal of biological chemistry 22 22474292
2022 Mechanical Stretch Induced Osteogenesis on Human Annulus Fibrosus Cells through Upregulation of BMP-2/6 Heterodimer and Activation of P38 and SMAD1/5/8 Signaling Pathways. Cells 20 36010676
2019 Pharmacological DNA demethylation restores SMAD1 expression and tumor suppressive signaling in diffuse large B-cell lymphoma. Blood advances 20 31648327
2020 Kaempferol Protects Renal Fibrosis through Activating the BMP-7-Smad1/5 Signaling Pathway. Biological & pharmaceutical bulletin 19 32115512
2017 Negative regulation of Smad1 pathway and collagen IV expression by store-operated Ca2+ entry in glomerular mesangial cells. American journal of physiology. Renal physiology 19 28298362
2013 SnoN facilitates ALK1-Smad1/5 signaling during embryonic angiogenesis. The Journal of cell biology 18 24019535
2011 Smad1 signaling restricts hematopoietic potential after promoting hemangioblast commitment. Blood 18 21515822
2020 BMP9 attenuates occurrence of venous malformation by maintaining endothelial quiescence and strengthening vessel walls via SMAD1/5/ID1/α-SMA pathway. Journal of molecular and cellular cardiology 17 32730768
2011 Bone morphogenetic protein 6-induced interleukin-1β expression in macrophages requires PU.1/Smad1 interaction. Molecular immunology 17 21571370
2007 Induction of Smad1 by MT1-MMP contributes to tumor growth. International journal of cancer 17 17455258
2003 Expression and localization of Smad1, Smad2 and Smad4 proteins in rat testis during postnatal development. Asian journal of andrology 17 12647004
2022 BMP/SMAD1/5 Signaling in the Endometrial Epithelium Is Essential for Receptivity and Early Pregnancy. Endocrinology 16 35383354
2022 Circular RNA CREBBP modulates cartilage degradation by activating the Smad1/5 pathway through the TGFβ2/ALK1 axis. Experimental & molecular medicine 16 36224344
2021 SMAD1 as a biomarker and potential therapeutic target in drug-resistant multiple myeloma. Biomarker research 16 34134766
2021 DUSP5 promotes osteogenic differentiation through SCP1/2-dependent phosphorylation of SMAD1. Stem cells (Dayton, Ohio) 16 34169608
2015 P53 deficiency-induced Smad1 upregulation suppresses tumorigenesis and causes chemoresistance in colorectal cancers. Journal of molecular cell biology 16 25757624
2015 Smad1/5/8 are myogenic regulators of murine and human mesoangioblasts. Journal of molecular cell biology 16 26450990
2012 Smad1/Smad5 signaling in limb ectoderm functions redundantly and is required for interdigital programmed cell death. Developmental biology 16 22240098
2023 BMP4-SMAD1/5/9-RUNX2 pathway activation inhibits neurogenesis and oligodendrogenesis in Alzheimer's patients' iPSCs in senescence-related conditions. Stem cell reports 15 36764297
2020 miR-203a-3p promotes loureirin A-induced hair follicle stem cells differentiation by targeting Smad1. Anatomical record (Hoboken, N.J. : 2007) 15 32589363
2019 BMP4/Smad1 Signalling Promotes Spinal Dorsal Column Axon Regeneration and Functional Recovery After Injury. Molecular neurobiology 15 30924076
2007 Increased Smad1 expression and transcriptional activity enhances trans-differentiation of hepatic stellate cells. Journal of cellular physiology 15 17525996
2024 A KLF2-BMPER-Smad1/5 checkpoint regulates high fluid shear stress-mediated artery remodeling. Nature cardiovascular research 14 39196179
2021 Bone Morphogenetic Protein-2 Promotes Osteoclasts-mediated Osteolysis via Smad1 and p65 Signaling Pathways. Spine 14 33156278
2021 mTOR signaling regulates gastric epithelial progenitor homeostasis and gastric tumorigenesis via MEK1-ERKs and BMP-Smad1 pathways. Cell reports 14 33951440
2019 MicroRNA-145 targets Smad1 in endometrial stromal cells and regulates decidualization in rat. Journal of molecular medicine (Berlin, Germany) 14 30729278
2018 Reversal of Sp1 transactivation and TGFβ1/SMAD1 signaling by H2S prevent nickel-induced fibroblast activation. Toxicology and applied pharmacology 14 30055191
2005 Specific interaction between Smad1 and CHIP: a surface plasmon resonance study. Colloids and surfaces. B, Biointerfaces 14 15708501

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