SMAD3

UniProt ID: P84022
Organism: Homo sapiens
Review Status: IN PROGRESS
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Gene Description

SMAD3 (Mothers against decapentaplegic homolog 3) is a receptor-regulated SMAD (R-SMAD) that serves as the principal intracellular signal transducer and transcriptional modulator in the TGF-beta/activin signaling pathway. Upon TGF-beta receptor activation, SMAD3 is phosphorylated at its C-terminal SSXS motif by TGF-beta type I receptor (ALK5/TGFBR1), forms heteromeric complexes with SMAD4 (co-SMAD), and translocates to the nucleus where it directly binds DNA at SMAD binding elements (SBE: GTCT/AGAC) and CAGA-box variants through its MH1 domain. SMAD3 cooperates with transcription factors such as c-Jun/c-Fos at composite AP-1/SMAD elements. It also participates in activin and nodal signaling. Key biological outcomes include regulation of cell proliferation, EMT, fibrosis, immune regulation, and cell differentiation. Germline mutations cause Loeys-Dietz syndrome type 3.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 directly binds DNA at SMAD binding elements (SBEs) in cis-regulatory regions of target genes, demonstrated by crystal structure (PMID:9741623), CASTing assays (Itoh et al. 2024), and ChIP-seq studies. IBA is well-supported across SMAD family.
Reason: Core molecular function of SMAD3. The MH1 domain directly binds SBE sequences (GTCT/AGAC) in promoter/enhancer regions, extensively validated by structural and functional studies.
Supporting Evidence:
PMID:9311995
Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was phosphorylated by TbetaR-II kinase.
PMID:9732876
Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription from the TRE
GO:0009653 anatomical structure morphogenesis
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: SMAD3 is involved in anatomical structure morphogenesis via TGF-beta signaling. IBA supported across species.
Reason: This is a broad downstream biological process. While SMAD3 contributes to morphogenesis through TGF-beta signaling, this is a pleiotropic outcome rather than a core function. IBA is valid at this level for SMAD family.
GO:0030154 cell differentiation
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: SMAD3 is involved in cell differentiation through TGF-beta signaling. IBA supported.
Reason: Broad downstream process. TGF-beta/SMAD3 signaling regulates differentiation in many contexts (chondrocyte, osteoblast, T cell, etc.), but this is a pleiotropic outcome rather than core function.
GO:0032924 activin receptor signaling pathway
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 also mediates activin receptor signaling, being phosphorylated by ACVR1B. IBA well-supported.
Reason: Core biological process. SMAD3 is activated by activin type I receptor (ACVR1B/ALK4) in addition to TGF-beta receptors. UniProt confirms interaction with ACVR1B (PMID:9892009).
Supporting Evidence:
PMID:9892009
Roles of pathway-specific and inhibitory Smads in activin receptor signaling
GO:0000981 DNA-binding transcription factor activity, RNA polymerase II-specific
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 is a sequence-specific DNA-binding transcription factor that activates and represses RNA polymerase II target genes in response to TGF-beta signaling. IBA supported across the SMAD family.
Reason: Core molecular function. SMAD3 directly binds DNA through its MH1 domain and recruits transcriptional coactivators (p300/CBP) or corepressors (Ski/SnoN, TGIF) to regulate Pol II transcription.
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription
PMID:21947082
USP15 is required for TGFbeta and BMP responses in mammalian cells
GO:0060395 SMAD protein signal transduction
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 is a core component of the SMAD protein signal transduction pathway. Upon TGF-beta receptor activation, SMAD3 is phosphorylated, complexes with SMAD4, and translocates to the nucleus. IBA well-supported.
Reason: Core biological process. SMAD3 is one of the principal R-SMADs mediating TGF-beta/activin signal transduction through the canonical SMAD pathway.
Supporting Evidence:
PMID:9311995
TGF-beta induces heteromeric complexes of Smads 2, 3 and 4, and their concomitant translocation to the nucleus, which is required for efficient TGF-beta signal transduction
PMID:8774881
hMAD-3 but not hMAD-4 was phosphorylated and associated with the ligand-bound receptor complex
GO:0007179 transforming growth factor beta receptor signaling pathway
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 is a key intracellular mediator of TGF-beta receptor signaling. Activated by TGFBR1/ALK5 phosphorylation. IBA well-supported.
Reason: Core biological process. SMAD3 is phosphorylated directly by the TGF-beta type I receptor and is essential for canonical TGF-beta signal transduction.
Supporting Evidence:
PMID:8774881
The activity of hMAD-3 and -4 was regulated by the TGF-beta receptors, and hMAD-3 but not hMAD-4 was phosphorylated and associated with the ligand-bound receptor complex
PMID:9311995
Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor
GO:0071144 heteromeric SMAD protein complex
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 forms heteromeric SMAD complexes with SMAD4 upon TGF-beta stimulation. IBA well-supported across species.
Reason: Core cellular component. The SMAD3/SMAD4 heteromeric complex is the functional unit of TGF-beta signaling. Crystal structure solved (PMID:15350224).
Supporting Evidence:
PMID:9311995
TGF-beta induces heteromeric complexes of Smads 2, 3 and 4
GO:0045944 positive regulation of transcription by RNA polymerase II
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 positively regulates transcription by RNA polymerase II at TGF-beta target gene promoters. IBA supported.
Reason: Core biological process. SMAD3/SMAD4 complexes recruit p300/CBP coactivators to activate transcription of TGF-beta responsive genes.
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta
GO:0070411 I-SMAD binding
IBA
GO_REF:0000033
ACCEPT
Summary: SMAD3 binds inhibitory SMADs (SMAD6/SMAD7). IBA supported across the SMAD family.
Reason: Well-established interaction. Inhibitory SMADs compete with R-SMADs for receptor binding and recruit phosphatases/ubiquitin ligases. SMAD3 binds SMAD7 as part of pathway regulation.
GO:0003677 DNA binding
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for DNA binding. SMAD3 directly binds DNA through MH1 domain at SBE sequences.
Reason: Correct but overly broad. More specific DNA binding terms (GO:0000978, GO:0000987) are already annotated with experimental evidence. This IEA is acceptable as a general parent term.
GO:0005634 nucleus
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for nucleus localization. SMAD3 translocates to nucleus upon TGF-beta stimulation.
Reason: Well-supported. SMAD3 is found in nucleus upon activation. Multiple IDA annotations also confirm this.
GO:0005667 transcription regulator complex
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for transcription regulator complex. SMAD3 forms complexes with SMAD4 and other transcription factors.
Reason: Correct. SMAD3/SMAD4 complexes function as transcription regulator complexes. Also supported by IDA (PMID:21947082).
GO:0005737 cytoplasm
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for cytoplasm localization. SMAD3 resides in cytoplasm when inactive.
Reason: Well-supported. SMAD3 shuttles between cytoplasm and nucleus, residing primarily in cytoplasm before TGF-beta stimulation.
GO:0006355 regulation of DNA-templated transcription
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for regulation of DNA-templated transcription based on InterPro domain mapping.
Reason: Correct. SMAD3 is a transcription factor that regulates DNA-templated transcription. Consistent with experimental annotations.
GO:0032502 developmental process
IEA
GO_REF:0000117
MARK AS OVER ANNOTATED
Summary: IEA for developmental process. Very broad term from ARBA prediction.
Reason: Excessively broad. While TGF-beta/SMAD3 signaling contributes to development, GO:0032502 is too general to be informative. More specific developmental processes are annotated elsewhere.
GO:0046872 metal ion binding
IEA
GO_REF:0000043
MODIFY
Summary: IEA for metal ion binding from UniProt keyword mapping. SMAD3 MH1 domain coordinates a zinc ion essential for DNA binding.
Reason: The annotation is correct in that SMAD3 binds metal ions, but the term is too broad. The MH1 domain contains a zinc-binding site essential for structural integrity and DNA binding (PMID:12686552). Should be zinc ion binding (GO:0008270) which is already annotated with IDA.
Proposed replacements: zinc ion binding
GO:0071363 cellular response to growth factor stimulus
IEA
GO_REF:0000117
ACCEPT
Summary: IEA for cellular response to growth factor stimulus. SMAD3 mediates TGF-beta responses.
Reason: Correct but broad. SMAD3 mediates cellular responses to TGF-beta and activin, which are growth factors. More specific pathway terms are also annotated.
GO:0141091 transforming growth factor beta receptor superfamily signaling pathway
IEA
GO_REF:0000117
ACCEPT
Summary: IEA for TGF-beta receptor superfamily signaling pathway from ARBA.
Reason: Correct. SMAD3 is a core mediator of TGF-beta receptor superfamily signaling. More specific TGF-beta pathway terms are also present.
GO:0005515 protein binding
IPI
PMID:11278756
Ski-interacting protein interacts with Smad proteins to augm...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11278756). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:11387212
The adaptor molecule Disabled-2 links the transforming growt...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11387212). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:12154125
Smad3 allostery links TGF-beta receptor kinase activation to...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12154125). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:12650946
Identification of three novel Smad binding proteins involved...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12650946). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:12857746
Direct interaction of Ski with either Smad3 or Smad4 is nece...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12857746). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:14525983
DACH1 inhibits transforming growth factor-beta signaling thr...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:14525983). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:15084259
Integration of Smad and forkhead pathways in the control of ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15084259). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:15231748
Functional proteomics mapping of a human signaling pathway.
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15231748). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:15350224
Structural basis of heteromeric smad protein assembly in TGF...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15350224). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:15527767
Proteomics-based identification of proteins interacting with...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15527767). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:16007207
Hepatitis C virus core variants isolated from liver tumor bu...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:16007207). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:17785517
Msk is required for nuclear import of TGF-{beta}/BMP-activat...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17785517). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:18548003
SMAD proteins control DROSHA-mediated microRNA maturation.
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18548003). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:18729074
Identification of novel Smad2 and Smad3 associated proteins ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18729074). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:19032343
Ski co-repressor complexes maintain the basal repressed stat...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19032343). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:19135894
FAM/USP9x, a deubiquitinating enzyme essential for TGFbeta s...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19135894). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:20061380
MTMR4 attenuates transforming growth factor beta (TGFbeta) s...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20061380). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:20211142
An atlas of combinatorial transcriptional regulation in mous...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20211142). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21258410
Selective targeting of activating and inhibitory Smads by di...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21258410). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21297662
Homeodomain protein DLX4 counteracts key transcriptional con...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21297662). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21532621
Nodal enhances the activity of FoxO3a and its synergistic in...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21532621). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21597466
Antagonistic regulation of EMT by TIF1Ξ³ and Smad4 in mammary...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21597466). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21828274). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:21988832
Toward an understanding of the protein interaction network o...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21988832). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:22045334
Ablation of Smurf2 reveals an inhibition in TGF-Ξ² signalling...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22045334). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:22442258
Intercellular variation in signaling through the TGF-Ξ² pathw...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22442258). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:22538441
CSMD1 exhibits antitumor activity in A375 melanoma cells thr...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22538441). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:25416956
A proteome-scale map of the human interactome network.
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25416956). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:25502805
A massively parallel pipeline to clone DNA variants and exam...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25502805). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:25609649
Proteomic analyses reveal distinct chromatin-associated and ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25609649). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:25670079
14-3-3ΞΆ turns TGF-Ξ²'s function from tumor suppressor to meta...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25670079). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:25910212
Widespread macromolecular interaction perturbations in human...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25910212). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:26680585
ShcA Protects against Epithelial-Mesenchymal Transition thro...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:26680585). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:27107012
Pooled-matrix protein interaction screens using Barcode Fusi...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:27107012). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:28471448
MiR-1 suppresses tumor cell proliferation in colorectal canc...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:28471448). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:29892012
An interactome perturbation framework prioritizes damaging m...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29892012). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:29997244
LuTHy: a double-readout bioluminescence-based two-hybrid tec...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29997244). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:31515488
Extensive disruption of protein interactions by genetic vari...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:31515488). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:32296183
A reference map of the human binary protein interactome.
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:32296183). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:32814053
Interactome Mapping Provides a Network of Neurodegenerative ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:32814053). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:33961781). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:35512704
Systematic discovery of mutation-directed neo-protein-protei...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:35512704). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:39243984
Gain-of-function variants in SMAD4 compromise respiratory ep...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:39243984). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0042802 identical protein binding
IPI
PMID:18729074
Identification of novel Smad2 and Smad3 associated proteins ...
ACCEPT
Summary: SMAD3 forms homo-oligomers. Confirmed by co-immunoprecipitation and structural studies.
Reason: SMAD3 homo-oligomerization is well-documented. Crystal structure shows SMAD3 trimeric assembly (PMID:11224571).
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
GO:0042802 identical protein binding
IPI
PMID:22045334
Ablation of Smurf2 reveals an inhibition in TGF-Ξ² signalling...
ACCEPT
Summary: SMAD3 forms homo-oligomers. Confirmed by co-immunoprecipitation and structural studies.
Reason: SMAD3 homo-oligomerization is well-documented. Crystal structure shows SMAD3 trimeric assembly (PMID:11224571).
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
GO:0000122 negative regulation of transcription by RNA polymerase II
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for negative regulation of Pol II transcription. SMAD3 can also repress transcription at certain loci.
Reason: SMAD3 acts as both transcriptional activator and repressor depending on context and cofactors (Ski/SnoN, TGIF). Also supported by IDA (PMID:28467929).
GO:0000165 MAPK cascade
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for MAPK cascade.
Reason: SMAD3 is not a core component of the MAPK cascade. While MAPK phosphorylates SMAD3 linker region, and SMAD3/AP-1 cooperate at promoters, SMAD3 does not directly participate in the MAPK cascade itself. Cross-talk annotation is misleading.
GO:0000977 RNA polymerase II transcription regulatory region sequence-specific DNA binding
IEA
GO_REF:0000120
ACCEPT
Summary: IEA (Ensembl/UniProt) for RNA polymerase II transcription regulatory region sequence-specific DNA binding.
Reason: Correct. SMAD3 binds SBE sequences in Pol II regulatory regions. Also supported by IDA annotations.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for DNA-binding transcription activator activity, Pol II-specific.
Reason: Correct. SMAD3 activates Pol II transcription at TGF-beta target genes. Also supported by multiple IDA annotations.
GO:0001649 osteoblast differentiation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for osteoblast differentiation from mouse data.
Reason: Valid downstream effect. SMAD3 regulates osteoblast differentiation and chondrogenesis (UniProt confirms SMAD3 is a regulator of osteogenesis). Non-core pleiotropic process.
GO:0001657 ureteric bud development
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for ureteric bud development from mouse data.
Reason: Very specific developmental process likely reflecting pleiotropic TGF-beta pathway effects in kidney development. No direct evidence for SMAD3-specific role in ureteric bud development in humans.
GO:0001836 release of cytochrome c from mitochondria
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for release of cytochrome c from mitochondria from rat data.
Reason: Indirect downstream effect. TGF-beta/SMAD3 signaling can trigger apoptosis in some contexts, which may involve cytochrome c release, but this is not a direct function of SMAD3.
GO:0003682 chromatin binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for chromatin binding from mouse data.
Reason: Correct. SMAD3 binds chromatin at enhancer/promoter regions. Also supported by IDA chromatin localization (PMID:21828274).
GO:0003690 double-stranded DNA binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for double-stranded DNA binding from mouse data.
Reason: Correct. Crystal structure confirms SMAD3 MH1 domain binds dsDNA at SBE sequences (PMID:9741623, PMID:12686552).
GO:0003700 DNA-binding transcription factor activity
IEA
GO_REF:0000120
ACCEPT
Summary: IEA for DNA-binding transcription factor activity.
Reason: Correct. SMAD3 is a bona fide DNA-binding transcription factor. Also supported by multiple IDA annotations.
GO:0005518 collagen binding
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for collagen binding from mouse data.
Reason: SMAD3 regulates collagen gene expression as a transcription factor, but there is no evidence it directly binds collagen protein. This appears to be a misannotation or conflation of transcriptional regulation with physical binding.
GO:0005886 plasma membrane
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for plasma membrane localization from mouse data.
Reason: SMAD3 can associate with TGF-beta receptors at the plasma membrane during initial activation, but its primary functional locations are cytoplasm and nucleus. Plasma membrane localization is transient.
GO:0007179 transforming growth factor beta receptor signaling pathway
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for TGF-beta receptor signaling pathway from rat orthologs.
Reason: Correct. Core biological process for SMAD3. Also supported by multiple IDA and IMP annotations.
GO:0007254 JNK cascade
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for JNK cascade from mouse data.
Reason: SMAD3 is not a component of the JNK cascade. While TGF-beta can activate JNK through non-canonical signaling and SMAD3/JUN cooperate at AP-1 sites, SMAD3 does not participate directly in JNK signal transduction.
GO:0008013 beta-catenin binding
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA annotation for beta-catenin binding via Ensembl Compara ortholog transfer. SMAD3 has been reported to interact with beta-catenin in the context of Wnt/TGF-beta crosstalk, but this is not a primary binding activity.
Reason: While SMAD3 can interact with beta-catenin in certain cellular contexts as part of Wnt/TGF-beta signaling crosstalk, beta-catenin binding is not a core molecular function of SMAD3. This IEA annotation likely represents a context-dependent interaction rather than an intrinsic binding activity.
GO:0008285 negative regulation of cell population proliferation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for negative regulation of cell population proliferation from rat data.
Reason: Valid downstream effect. TGF-beta/SMAD3 signaling inhibits cell proliferation through transcriptional regulation of CDK inhibitors (p15, p21). Also supported by IMP (PMID:14555988). Non-core pleiotropic process.
GO:0010332 response to gamma radiation
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for response to gamma radiation from rat data.
Reason: Indirect response. While SMAD3 may be activated or its expression altered by gamma radiation in some cell types, this is not a direct or core function of SMAD3.
GO:0010467 gene expression
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for gene expression from mouse data.
Reason: Excessively broad. 'Gene expression' is too general. SMAD3 regulates transcription, which is a component of gene expression, but more specific terms are already annotated.
GO:0019899 enzyme binding
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for enzyme binding from rat data.
Reason: While SMAD3 binds kinases (TGFBR1, CDKs, PDPK1), phosphatases (PPM1A), and E3 ligases, 'enzyme binding' is too broad. More specific binding terms are already present.
GO:0023019 signal transduction involved in regulation of gene expression
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for signal transduction involved in regulation of gene expression from rat data.
Reason: Correct description of SMAD3 core function: signal transduction (TGF-beta pathway) that directly regulates gene expression.
GO:0030325 adrenal gland development
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for adrenal gland development from rat data.
Reason: Very specific developmental process likely reflecting broad TGF-beta pathway pleiotropic effects. No direct evidence for SMAD3-specific role in human adrenal development.
GO:0030335 positive regulation of cell migration
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for positive regulation of cell migration from rat data.
Reason: Valid downstream effect. TGF-beta/SMAD3 signaling promotes cell migration, particularly in EMT context. UniProt notes SMAD3 role in EMT. Non-core pleiotropic process.
GO:0030501 positive regulation of bone mineralization
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for positive regulation of bone mineralization from rat data.
Reason: Valid downstream effect. UniProt notes SMAD3 is a regulator of osteogenesis. Non-core tissue-specific process.
GO:0031490 chromatin DNA binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for chromatin DNA binding from mouse data.
Reason: Correct. SMAD3 binds DNA within chromatin context at enhancers and promoters. Supported by ChIP studies.
GO:0032332 positive regulation of chondrocyte differentiation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for positive regulation of chondrocyte differentiation from mouse data.
Reason: Valid downstream effect. UniProt notes SMAD3 is a regulator of chondrogenesis. Non-core developmental process.
GO:0032731 positive regulation of interleukin-1 beta production
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of IL-1 beta production from rat data.
Reason: Indirect downstream effect of TGF-beta signaling in immune regulation. Not a direct function of SMAD3 as a transcription factor.
GO:0032916 positive regulation of transforming growth factor beta3 production
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of TGF-beta3 production from rat data.
Reason: While SMAD3 could transcriptionally regulate TGF-beta3 expression as a feedback mechanism, this is a very specific downstream effect. Limited direct human evidence.
GO:0032993 protein-DNA complex
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for protein-DNA complex from mouse data.
Reason: Correct. SMAD3 forms protein-DNA complexes at SBE-containing regulatory regions. Crystal structure of SMAD3 MH1-DNA complex solved (PMID:12686552).
GO:0033689 negative regulation of osteoblast proliferation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for negative regulation of osteoblast proliferation from mouse data.
Reason: Valid downstream effect consistent with SMAD3 role in osteogenesis regulation. Non-core tissue-specific process.
GO:0036120 cellular response to platelet-derived growth factor stimulus
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for cellular response to PDGF stimulus from rat data.
Reason: Indirect. PDGF may modulate TGF-beta/SMAD signaling through MAPK crosstalk, but SMAD3 is not a direct mediator of PDGF responses.
GO:0042110 T cell activation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for T cell activation from mouse data.
Reason: Valid downstream effect. TGF-beta/SMAD3 signaling is critical for T cell differentiation and immune regulation. SMAD3 knockout mice have T cell activation defects. Non-core immune process.
GO:0042177 negative regulation of protein catabolic process
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for negative regulation of protein catabolic process from rat data.
Reason: Indirect downstream effect. While TGF-beta signaling can affect protein stability, this is not a direct SMAD3 function.
GO:0042220 response to cocaine
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for response to cocaine from rat data.
Reason: Highly specific environmental response likely from rat neurological studies. No evidence SMAD3 directly mediates cocaine responses. Over-annotation from ortholog transfer.
GO:0042802 identical protein binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for identical protein binding from mouse data.
Reason: Correct. SMAD3 forms homomeric complexes. SMAD3 homo-oligomerization is well-documented (PMID:9670020).
GO:0043066 negative regulation of apoptotic process
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for negative regulation of apoptotic process from rat data.
Reason: Context-dependent downstream effect. TGF-beta/SMAD3 can both promote and inhibit apoptosis depending on cell type. Non-core pleiotropic process.
GO:0043565 sequence-specific DNA binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for sequence-specific DNA binding from rat data.
Reason: Correct. SMAD3 MH1 domain binds specific DNA sequences (SBE: GTCT/AGAC). Crystal structure confirms sequence-specific binding (PMID:9741623).
GO:0045429 positive regulation of nitric oxide biosynthetic process
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for positive regulation of nitric oxide biosynthesis from rat data.
Reason: Context-dependent downstream effect. TGF-beta/SMAD3 can regulate iNOS expression in certain cell types. Also supported by IDA (PMID:27038547). Non-core process.
GO:0045944 positive regulation of transcription by RNA polymerase II
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for positive regulation of Pol II transcription from rat data.
Reason: Correct. SMAD3 activates Pol II transcription. Also supported by IBA and IMP annotations.
GO:0050728 negative regulation of inflammatory response
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for negative regulation of inflammatory response from mouse data.
Reason: Valid downstream effect. TGF-beta/SMAD3 is a key anti-inflammatory pathway. Also supported by NAS (ComplexPortal). Non-core immune process.
GO:0050776 regulation of immune response
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for regulation of immune response from mouse data.
Reason: Valid broad downstream effect. TGF-beta/SMAD3 is critical for immune regulation. Non-core pleiotropic process.
GO:0050821 protein stabilization
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for protein stabilization from rat data.
Reason: Indirect effect. While SMAD3 can affect protein stability through transcriptional regulation of ubiquitin pathway components, protein stabilization is not a direct function.
GO:0050927 positive regulation of positive chemotaxis
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of positive chemotaxis from rat data.
Reason: Indirect downstream effect of TGF-beta signaling. Not a direct SMAD3 function.
GO:0051496 positive regulation of stress fiber assembly
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of stress fiber assembly from rat data.
Reason: Indirect downstream effect. TGF-beta can promote stress fiber formation through EMT-related cytoskeletal remodeling, but this is not a direct SMAD3 transcription factor function.
GO:0051881 regulation of mitochondrial membrane potential
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for regulation of mitochondrial membrane potential from rat data.
Reason: Indirect downstream effect. Not a direct function of SMAD3 as a transcription factor.
GO:0051894 positive regulation of focal adhesion assembly
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of focal adhesion assembly from rat data.
Reason: Indirect downstream effect of TGF-beta-mediated EMT or cytoskeletal changes. Not a direct SMAD3 function.
GO:0060290 transdifferentiation
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for transdifferentiation from rat data.
Reason: Valid downstream effect. TGF-beta/SMAD3 drives EMT (a form of transdifferentiation). Non-core pleiotropic process.
GO:0060395 SMAD protein signal transduction
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for SMAD protein signal transduction from rat data.
Reason: Correct. Core biological process. Also supported by multiple IBA and IDA annotations.
GO:0061001 regulation of dendritic spine morphogenesis
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for regulation of dendritic spine morphogenesis from rat data.
Reason: Very specific neuronal process. Likely indirect TGF-beta pathway effect. No direct evidence for SMAD3-specific role in dendritic spine morphogenesis in humans.
GO:0061045 negative regulation of wound healing
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for negative regulation of wound healing from mouse data.
Reason: Valid downstream effect. UniProt notes SMAD3 has inhibitory effect on wound healing. Non-core tissue-specific process.
GO:0061629 RNA polymerase II-specific DNA-binding transcription factor binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for RNA polymerase II-specific DNA-binding TF binding from mouse data.
Reason: Correct. SMAD3 binds other Pol II TFs (c-Jun, c-Fos, FOXH1, etc.). Also supported by IPI annotations.
GO:0061767 negative regulation of lung blood pressure
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for negative regulation of lung blood pressure from rat data.
Reason: Very specific physiological process likely from rat cardiovascular studies. Indirect downstream effect of TGF-beta signaling.
GO:0071333 cellular response to glucose stimulus
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for cellular response to glucose stimulus from rat data.
Reason: Indirect. TGF-beta/SMAD signaling may be modulated by metabolic conditions, but SMAD3 is not a direct glucose sensor.
GO:0071560 cellular response to transforming growth factor beta stimulus
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for cellular response to TGF-beta stimulus from rat data.
Reason: Correct. Core biological process. SMAD3 is a primary mediator of cellular responses to TGF-beta. Also supported by IDA (PMID:12902338).
GO:0090263 positive regulation of canonical Wnt signaling pathway
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for positive regulation of canonical Wnt signaling pathway from rat data.
Reason: SMAD3 is not a direct component of the Wnt pathway. While there is TGF-beta/Wnt pathway crosstalk, annotating SMAD3 to Wnt pathway regulation is misleading.
GO:0097191 extrinsic apoptotic signaling pathway
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for extrinsic apoptotic signaling pathway from rat data.
Reason: Valid downstream effect. TGF-beta can trigger apoptosis through SMAD3-dependent transcriptional programs. Also supported by IMP (PMID:15334054). Non-core process.
GO:0097305 response to alcohol
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for response to alcohol from rat data.
Reason: Likely from rat liver/fibrosis studies where alcohol induces TGF-beta signaling. Not a direct SMAD3 function.
GO:0098586 cellular response to virus
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA (Ensembl orthologs) for cellular response to virus from mouse data.
Reason: TGF-beta/SMAD3 signaling is involved in antiviral responses and immune regulation. SARS-CoV nucleoprotein interacts with SMAD3 (PMID:18055455). Non-core immune process.
GO:1903243 negative regulation of cardiac muscle hypertrophy in response to stress
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA (Ensembl orthologs) for negative regulation of cardiac muscle hypertrophy from rat data.
Reason: Very specific cardiac process. Indirect TGF-beta pathway effect. Not a direct SMAD3 function.
GO:1990841 promoter-specific chromatin binding
IEA
GO_REF:0000107
ACCEPT
Summary: IEA (Ensembl orthologs) for promoter-specific chromatin binding from mouse data.
Reason: Correct. SMAD3 binds chromatin at specific promoters of TGF-beta target genes.
GO:0005654 nucleoplasm
IDA
GO_REF:0000052
ACCEPT
Summary: Nucleoplasm localization (GO_REF:0000052).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0006355 regulation of DNA-templated transcription
NAS
PMID:35359452
The Interplay Between TGF-Ξ² Signaling and Cell Metabolism.
ACCEPT
Summary: Regulation of DNA-templated transcription confirmed (PMID:35359452).
Reason: Core biological process. SMAD3 is a transcription factor that regulates gene expression.
GO:0007179 transforming growth factor beta receptor signaling pathway
NAS
PMID:35359452
The Interplay Between TGF-Ξ² Signaling and Cell Metabolism.
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:35359452).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0032924 activin receptor signaling pathway
NAS
PMID:15150278
Nodal and ALK7 inhibit proliferation and induce apoptosis in...
ACCEPT
Summary: Activin receptor signaling pathway confirmed (PMID:15150278).
Reason: Core biological process. SMAD3 mediates activin signaling through ACVR1B.
GO:0005634 nucleus
IPI
PMID:20935647
IL-37 is a fundamental inhibitor of innate immunity.
ACCEPT
Summary: Nucleus localization confirmed (PMID:20935647). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005737 cytoplasm
IPI
PMID:20935647
IL-37 is a fundamental inhibitor of innate immunity.
ACCEPT
Summary: Cytoplasm localization confirmed (PMID:20935647). SMAD3 resides in cytoplasm before activation.
Reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
GO:0006355 regulation of DNA-templated transcription
IDA
PMID:25060702
A Smad3 and TTF-1/NKX2-1 complex regulates Smad4-independent...
ACCEPT
Summary: Regulation of DNA-templated transcription confirmed (PMID:25060702).
Reason: Core biological process. SMAD3 is a transcription factor that regulates gene expression.
GO:0031665 negative regulation of lipopolysaccharide-mediated signaling pathway
NAS
PMID:29789615
IL-37 isoform D downregulates pro-inflammatory cytokines exp...
KEEP AS NON CORE
Summary: Negative regulation of LPS-mediated signaling pathway (PMID:29789615). NAS from ComplexPortal.
Reason: Valid downstream effect reflecting anti-inflammatory role of TGF-beta/SMAD3. Non-core immune process.
GO:0050728 negative regulation of inflammatory response
NAS
PMID:27060871
Suppression of innate inflammation and immunity by interleuk...
KEEP AS NON CORE
Summary: Negative regulation of inflammatory response (PMID:27060871). NAS from ComplexPortal.
Reason: Valid downstream effect. TGF-beta/SMAD3 is a key anti-inflammatory pathway. Non-core immune process.
GO:0071635 negative regulation of transforming growth factor beta production
NAS
PMID:20935647
IL-37 is a fundamental inhibitor of innate immunity.
KEEP AS NON CORE
Summary: Negative regulation of TGF-beta production (PMID:20935647). NAS from ComplexPortal.
Reason: Valid feedback regulation. SMAD3 can modulate TGF-beta expression as part of signaling feedback. Non-core regulatory process.
GO:0061450 trophoblast cell migration
IDA
PMID:34432647
GDF-8 stimulates trophoblast cell invasion by inducing ALK5-...
KEEP AS NON CORE
Summary: Trophoblast cell migration (PMID:34432647).
Reason: Specific developmental process. TGF-beta/SMAD3 promotes trophoblast migration. Non-core tissue-specific process.
GO:0045596 negative regulation of cell differentiation
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Negative regulation of cell differentiation (GO_REF:0000024). ISS from mouse ortholog.
Reason: Valid downstream effect. TGF-beta/SMAD3 can inhibit differentiation in certain contexts. Non-core pleiotropic process.
GO:0060391 positive regulation of SMAD protein signal transduction
IMP
PMID:15107418
Activin receptor-like kinase-7 induces apoptosis through act...
ACCEPT
Summary: Positive regulation of SMAD protein signal transduction (PMID:15107418).
Reason: SMAD3 positively promotes SMAD signal transduction as an R-SMAD that forms complexes with SMAD4.
GO:0097190 apoptotic signaling pathway
IMP
PMID:15107418
Activin receptor-like kinase-7 induces apoptosis through act...
KEEP AS NON CORE
Summary: Apoptotic signaling pathway (PMID:15107418).
Reason: Valid downstream effect. TGF-beta/SMAD3 can trigger apoptosis in certain contexts. Non-core pleiotropic process.
GO:0030279 negative regulation of ossification
IDA
PMID:22155034
Bone morphogenetic protein-3b (BMP-3b) inhibits osteoblast d...
KEEP AS NON CORE
Summary: Negative regulation of ossification (PMID:22155034).
Reason: Valid downstream effect. UniProt notes SMAD3 inhibits early healing of bone fractures. Non-core tissue-specific process.
GO:0007179 transforming growth factor beta receptor signaling pathway
IMP
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:8774881).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0060395 SMAD protein signal transduction
IMP
PMID:30729664
Novel role of the clustered miR-23b-3p and miR-27b-3p in enh...
ACCEPT
Summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:30729664).
Reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
GO:1901203 positive regulation of extracellular matrix assembly
IMP
PMID:30729664
Novel role of the clustered miR-23b-3p and miR-27b-3p in enh...
KEEP AS NON CORE
Summary: Positive regulation of extracellular matrix assembly (PMID:30729664).
Reason: Valid downstream effect. TGF-beta/SMAD3 transcriptionally activates ECM genes (collagen, fibronectin). Non-core but well-documented in fibrosis context.
GO:1902895 positive regulation of miRNA transcription
IMP
PMID:30729664
Novel role of the clustered miR-23b-3p and miR-27b-3p in enh...
KEEP AS NON CORE
Summary: Positive regulation of miRNA transcription (PMID:30729664).
Reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
GO:1990776 response to angiotensin
IMP
PMID:30729664
Novel role of the clustered miR-23b-3p and miR-27b-3p in enh...
KEEP AS NON CORE
Summary: Response to angiotensin (PMID:30729664). Acts upstream of.
Reason: TGF-beta/SMAD3 signaling cross-talks with angiotensin pathways in cardiovascular/fibrotic contexts. Non-core.
GO:0060395 SMAD protein signal transduction
IDA
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9732876).
Reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
GO:0031625 ubiquitin protein ligase binding
IPI
PMID:15781469
CHIP controls the sensitivity of transforming growth factor-...
ACCEPT
Summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP (PMID:15781469).
Reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:9311995).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0060395 SMAD protein signal transduction
IDA
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9311995).
Reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
GO:0001222 transcription corepressor binding
IPI
PMID:14612439
A novel E1A-like inhibitor of differentiation (EID) family m...
ACCEPT
Summary: Transcription corepressor binding confirmed. SMAD3 interacts with corepressors including Ski, SnoN, TGIF (PMID:14612439).
Reason: Well-supported. SMAD3 recruits transcriptional corepressors at certain target genes.
GO:0003700 DNA-binding transcription factor activity
IDA
PMID:10823886
Transforming growth factor beta -inducible independent bindi...
ACCEPT
Summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:10823886).
Reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:10823886
Transforming growth factor beta -inducible independent bindi...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:10823886).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0060395 SMAD protein signal transduction
IDA
PMID:10823886
Transforming growth factor beta -inducible independent bindi...
ACCEPT
Summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:10823886).
Reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
GO:0060395 SMAD protein signal transduction
IDA
PMID:9111321
Heteromeric and homomeric interactions correlate with signal...
ACCEPT
Summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9111321).
Reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
GO:0000987 cis-regulatory region sequence-specific DNA binding
IMP
PMID:32141990
miR-4286/TGF-Ξ²1/Smad3-Negative Feedback Loop Ameliorated Vas...
ACCEPT
Summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:32141990).
Reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
GO:0003700 DNA-binding transcription factor activity
IMP
PMID:32141990
miR-4286/TGF-Ξ²1/Smad3-Negative Feedback Loop Ameliorated Vas...
ACCEPT
Summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:32141990).
Reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
GO:1902894 negative regulation of miRNA transcription
IMP
PMID:32141990
miR-4286/TGF-Ξ²1/Smad3-Negative Feedback Loop Ameliorated Vas...
KEEP AS NON CORE
Summary: Negative regulation of miRNA transcription (PMID:32141990).
Reason: Valid specific function. SMAD3 can also repress miRNA transcription at certain loci. Non-core regulatory process.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IDA
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
ACCEPT
Summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:24378993).
Reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
IDA
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
ACCEPT
Summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24378993).
Reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
IMP
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
ACCEPT
Summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24378993).
Reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
GO:0010629 negative regulation of gene expression
IMP
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
ACCEPT
Summary: Negative regulation of gene expression (PMID:24378993).
Reason: Valid. SMAD3 can repress gene expression at certain loci through corepressor recruitment.
GO:1902895 positive regulation of miRNA transcription
IDA
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
KEEP AS NON CORE
Summary: Positive regulation of miRNA transcription (PMID:24378993).
Reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
GO:1902895 positive regulation of miRNA transcription
IMP
PMID:24378993
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathwa...
KEEP AS NON CORE
Summary: Positive regulation of miRNA transcription (PMID:24378993).
Reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2179276
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2179276).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005515 protein binding
IPI
PMID:15051726
Atrophin-1-interacting protein 4/human Itch is a ubiquitin E...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15051726). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:29899023
NEDD9 targets COL3A1 to promote endothelial fibrosis and pul...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29899023). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
ISS
PMID:24964035
Pin1 promotes degradation of Smad proteins and their interac...
ACCEPT
Summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24964035).
Reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
IDA
PMID:25605017
Reduction of miR-29c enhances pancreatic cancer cell migrati...
ACCEPT
Summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:25605017).
Reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
GO:0032810 sterol response element binding
IGI
PMID:25605017
Reduction of miR-29c enhances pancreatic cancer cell migrati...
KEEP AS NON CORE
Summary: Sterol response element binding. SMAD3 binds SRE in IGI with TGF-beta1 (PMID:25605017).
Reason: Specialized binding activity at specific genomic loci. Non-core but documented.
GO:0061629 RNA polymerase II-specific DNA-binding transcription factor binding
IPI
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: SMAD3 binds Pol II-specific DNA-binding TFs including c-Jun, c-Fos, PAX6 (PMID:9732876).
Reason: Core molecular function. SMAD3 cooperates with other TFs at composite promoter elements.
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta, through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos
GO:0140297 DNA-binding transcription factor binding
IPI
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
ACCEPT
Summary: DNA-binding transcription factor binding. SMAD3 binds FOXH1 (PMID:21828274).
Reason: Well-supported. SMAD3 cooperates with multiple DNA-binding TFs.
GO:0016922 nuclear receptor binding
IPI
PMID:31023188
LRP1 Deficiency in Vascular SMC Leads to Pulmonary Arterial ...
ACCEPT
Summary: Nuclear receptor binding. SMAD3 binds PPARgamma (PMID:31023188).
Reason: Well-documented interaction. SMAD3 interacts with nuclear receptors as part of transcriptional cross-talk.
GO:0008285 negative regulation of cell population proliferation
IMP
PMID:14555988
Differential regulation of TGF-beta signaling through Smad2,...
KEEP AS NON CORE
Summary: Negative regulation of cell population proliferation confirmed (PMID:14555988).
Reason: Valid downstream effect. TGF-beta/SMAD3 mediates growth arrest through CDK inhibitor induction. Non-core but important pleiotropic process.
GO:1902893 regulation of miRNA transcription
IC
PMID:25605017
Reduction of miR-29c enhances pancreatic cancer cell migrati...
KEEP AS NON CORE
Summary: Regulation of miRNA transcription (PMID:25605017).
Reason: Valid. SMAD3 regulates miRNA transcription. Parent term of both positive and negative regulation annotations. Non-core.
GO:0000122 negative regulation of transcription by RNA polymerase II
IDA
PMID:28467929
PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscl...
ACCEPT
Summary: Negative regulation of transcription by Pol II confirmed (PMID:28467929). SMAD3 can repress transcription.
Reason: Valid. SMAD3 acts as transcriptional repressor at certain loci with Ski/SnoN/TGIF cofactors.
GO:0001217 DNA-binding transcription repressor activity
IDA
PMID:28467929
PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscl...
ACCEPT
Summary: DNA-binding transcription repressor activity confirmed (PMID:28467929). SMAD3 can also repress transcription at certain loci.
Reason: Valid. SMAD3 acts as both activator and repressor depending on cofactor context (Ski/SnoN, TGIF for repression).
GO:0003677 DNA binding
IDA
PMID:28467929
PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscl...
ACCEPT
Summary: DNA binding confirmed by direct assay (PMID:28467929).
Reason: Core molecular function. SMAD3 MH1 domain directly binds DNA.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:28467929
PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscl...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:28467929).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9731111
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-9731111).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0001228 DNA-binding transcription activator activity, RNA polymerase II-specific
IMP
PMID:26311719
MicroRNA-143 Activation Regulates Smooth Muscle and Endothel...
ACCEPT
Summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:26311719).
Reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
GO:1902895 positive regulation of miRNA transcription
IMP
PMID:26311719
MicroRNA-143 Activation Regulates Smooth Muscle and Endothel...
KEEP AS NON CORE
Summary: Positive regulation of miRNA transcription (PMID:26311719).
Reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
GO:0005515 protein binding
IPI
PMID:23723426
Kindlin-2 mediates activation of TGF-Ξ²/Smad signaling and re...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:23723426). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005634 nucleus
IGI
PMID:23723426
Kindlin-2 mediates activation of TGF-Ξ²/Smad signaling and re...
ACCEPT
Summary: Nucleus localization confirmed (PMID:23723426). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005829 cytosol
IGI
PMID:23723426
Kindlin-2 mediates activation of TGF-Ξ²/Smad signaling and re...
ACCEPT
Summary: Cytosol localization (PMID:23723426).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IDA
PMID:18832382
HMGA2 and Smads co-regulate SNAIL1 expression during inducti...
ACCEPT
Summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:18832382).
Reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
GO:0005634 nucleus
IDA
PMID:21145499
The Crumbs complex couples cell density sensing to Hippo-dep...
ACCEPT
Summary: Nucleus localization confirmed (PMID:21145499). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005737 cytoplasm
IDA
PMID:21145499
The Crumbs complex couples cell density sensing to Hippo-dep...
ACCEPT
Summary: Cytoplasm localization confirmed (PMID:21145499). SMAD3 resides in cytoplasm before activation.
Reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IDA
PMID:31582430
Structural basis for distinct roles of SMAD2 and SMAD3 in FO...
ACCEPT
Summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:31582430).
Reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
GO:0001223 transcription coactivator binding
IPI
PMID:16777850
The novel PIAS-like protein hZimp10 enhances Smad transcript...
ACCEPT
Summary: Transcription coactivator binding confirmed. SMAD3 interacts with coactivators including p300/CBP, ZMIZ1 (PMID:16777850).
Reason: Well-supported. SMAD3 linker region and MH2 domain recruit p300/CBP coactivators.
GO:0000987 cis-regulatory region sequence-specific DNA binding
IC
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
ACCEPT
Summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:21828274).
Reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
GO:0000785 chromatin
ISA
GO_REF:0000113
ACCEPT
Summary: Chromatin localization (GO_REF:0000113). SMAD3 is found at chromatin at TGF-beta target gene loci.
Reason: Correct. ChIP studies confirm SMAD3 binding to chromatin at enhancer/promoter regions.
GO:0000981 DNA-binding transcription factor activity, RNA polymerase II-specific
ISA
GO_REF:0000113
ACCEPT
Summary: ISA from TFClass database for DNA-binding TF activity Pol II-specific.
Reason: Correct. SMAD3 is classified as a TF in TFClass. Also supported by IDA and IBA.
GO:0000981 DNA-binding transcription factor activity, RNA polymerase II-specific
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: SMAD3 functions as a DNA-binding transcription factor that directly binds SMAD binding elements (SBE: GTCT/AGAC) and CAGA-box motifs through its MH1 domain, and regulates RNA polymerase II-dependent transcription of TGF-beta responsive genes.
Reason: Core molecular function. SMAD3 is a bona fide sequence-specific DNA-binding transcription factor. Its MH1 domain directly contacts DNA at SBE sequences, and it recruits transcriptional coactivators (p300/CBP) or corepressors (Ski/SnoN) to regulate Pol II transcription. Consistent with IBA and ISA evidence already accepted for this term.
GO:0042307 positive regulation of protein import into nucleus
NAS
PMID:15799969
Nuclear targeting of transforming growth factor-beta-activat...
ACCEPT
Summary: Positive regulation of protein import into nucleus (PMID:15799969).
Reason: SMAD3 promotes nuclear import of the SMAD2/3-SMAD4 complex. Part of core signaling mechanism.
GO:0005515 protein binding
IPI
PMID:25556234
New host factors important for respiratory syncytial virus (...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25556234). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:18548003
SMAD proteins control DROSHA-mediated microRNA maturation.
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:18548003).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0017151 DEAD/H-box RNA helicase binding
IPI
PMID:18548003
SMAD proteins control DROSHA-mediated microRNA maturation.
ACCEPT
Summary: DEAD/H-box RNA helicase binding. SMAD3 interacts with DDX5 (p68) (PMID:18548003).
Reason: Documented interaction. DDX5/p68 modulates SMAD3-dependent transcription.
GO:0005515 protein binding
IPI
PMID:24613385
Hsp70 and Hsp90 oppositely regulate TGF-Ξ² signaling through ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:24613385). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0031962 nuclear mineralocorticoid receptor binding
IPI
PMID:12902338
Identification of glucocorticoid receptor domains involved i...
KEEP AS NON CORE
Summary: Nuclear mineralocorticoid receptor binding. SMAD3 interacts with NR3C2 (PMID:12902338).
Reason: Documented interaction representing transcriptional cross-talk between TGF-beta and mineralocorticoid signaling.
GO:0035259 nuclear glucocorticoid receptor binding
IPI
PMID:12902338
Identification of glucocorticoid receptor domains involved i...
KEEP AS NON CORE
Summary: Nuclear glucocorticoid receptor binding. SMAD3 interacts with NR3C1 (PMID:12902338).
Reason: Documented interaction representing transcriptional cross-talk between TGF-beta and glucocorticoid signaling.
GO:0071560 cellular response to transforming growth factor beta stimulus
IDA
PMID:12902338
Identification of glucocorticoid receptor domains involved i...
ACCEPT
Summary: Cellular response to TGF-beta stimulus confirmed (PMID:12902338).
Reason: Core biological process. SMAD3 is a primary mediator of cellular TGF-beta responses.
GO:0045429 positive regulation of nitric oxide biosynthetic process
IDA
PMID:27038547
MiRNA-199a-5p influences pulmonary artery hypertension via d...
KEEP AS NON CORE
Summary: Positive regulation of nitric oxide biosynthetic process (PMID:27038547).
Reason: Specific downstream effect in certain cell types. Non-core pleiotropic process.
GO:0051481 negative regulation of cytosolic calcium ion concentration
IDA
PMID:27038547
MiRNA-199a-5p influences pulmonary artery hypertension via d...
MARK AS OVER ANNOTATED
Summary: Negative regulation of cytosolic calcium ion concentration (PMID:27038547).
Reason: Indirect downstream effect. SMAD3 as a transcription factor does not directly regulate calcium concentrations.
GO:0005829 cytosol
TAS
Reactome:R-HSA-6781764
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-6781764).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005634 nucleus
IDA
PMID:25893292
Syntenin regulates TGF-Ξ²1-induced Smad activation and the ep...
ACCEPT
Summary: Nucleus localization confirmed (PMID:25893292). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0045599 negative regulation of fat cell differentiation
IDA
PMID:19816956
MiR-21 regulates adipogenic differentiation through the modu...
KEEP AS NON CORE
Summary: Negative regulation of fat cell differentiation (PMID:19816956).
Reason: Valid downstream effect. TGF-beta/SMAD3 inhibits adipogenesis. Non-core tissue-specific process.
GO:1901203 positive regulation of extracellular matrix assembly
IDA
PMID:21307346
Response gene to complement 32 interacts with Smad3 to promo...
KEEP AS NON CORE
Summary: Positive regulation of extracellular matrix assembly (PMID:21307346).
Reason: Valid downstream effect. TGF-beta/SMAD3 transcriptionally activates ECM genes (collagen, fibronectin). Non-core but well-documented in fibrosis context.
GO:0005634 nucleus
IDA
PMID:16007207
Hepatitis C virus core variants isolated from liver tumor bu...
ACCEPT
Summary: Nucleus localization confirmed (PMID:16007207). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:21947082).
Reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
GO:0006357 regulation of transcription by RNA polymerase II
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: Regulation of transcription by RNA polymerase II confirmed (PMID:21947082).
Reason: Core biological process. SMAD3 regulates Pol II transcription at target genes.
GO:0045944 positive regulation of transcription by RNA polymerase II
IMP
PMID:14555988
Differential regulation of TGF-beta signaling through Smad2,...
ACCEPT
Summary: Positive regulation of Pol II transcription confirmed (PMID:14555988).
Reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
GO:0010628 positive regulation of gene expression
IDA
PMID:21307346
Response gene to complement 32 interacts with Smad3 to promo...
ACCEPT
Summary: Positive regulation of gene expression (PMID:21307346).
Reason: Core biological process. SMAD3 activates expression of TGF-beta target genes.
GO:0005515 protein binding
IPI
PMID:20129061
TMEPAI, a transmembrane TGF-beta-inducible protein, sequeste...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20129061). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:24627487
C18 ORF1, a novel negative regulator of transforming growth ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:24627487). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0070412 R-SMAD binding
IPI
PMID:20129061
TMEPAI, a transmembrane TGF-beta-inducible protein, sequeste...
ACCEPT
Summary: R-SMAD binding confirmed. SMAD3 binds other R-SMADs (SMAD2) and itself (PMID:20129061).
Reason: Well-supported interaction. SMAD3 interacts with SMAD2 as part of the TGF-beta signaling pathway.
Supporting Evidence:
PMID:9311995
we observed TbetaR-activation-dependent interaction between Smad2 and Smad3
GO:0000785 chromatin
IDA
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
ACCEPT
Summary: Chromatin localization (PMID:21828274). SMAD3 is found at chromatin at TGF-beta target gene loci.
Reason: Correct. ChIP studies confirm SMAD3 binding to chromatin at enhancer/promoter regions.
GO:0043425 bHLH transcription factor binding
IPI
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
ACCEPT
Summary: bHLH transcription factor binding. SMAD3 binds HEB/TCF12 (PMID:21828274).
Reason: Documented interaction between SMAD3 and bHLH TFs at target gene promoters.
GO:0071144 heteromeric SMAD protein complex
IDA
PMID:21828274
HEB and E2A function as SMAD/FOXH1 cofactors.
ACCEPT
Summary: Heteromeric SMAD protein complex confirmed (PMID:21828274).
Reason: Core cellular component. SMAD3/SMAD4 heterotrimer is the functional transcriptional complex.
GO:0097191 extrinsic apoptotic signaling pathway
IMP
PMID:15334054
Hepatitis C viral proteins interact with Smad3 and different...
KEEP AS NON CORE
Summary: Extrinsic apoptotic signaling pathway (PMID:15334054).
Reason: Valid downstream effect. Non-core pleiotropic process.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2176475
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2176475).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2176491
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2176491).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2176502
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2176502).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2176503
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2176503).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2179274
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2179274).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-1535903
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-1535903).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-173481
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-173481).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-173488
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-173488).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-173545
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-173545).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-209055
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-209055).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2106579
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2106579).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2127257
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2127257).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2186607
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2186607).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2186643
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2186643).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2187309
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2187309).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2187325
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2187325).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2187330
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2187330).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2187388
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2187388).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-870449
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-870449).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-870538
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-870538).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-8878143
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-8878143).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-8878178
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-8878178).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-8952226
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-8952226).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9617996
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9617996).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9618004
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9618004).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9618021
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9618021).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9733207
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9733207).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9733247
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9733247).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9736959
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9736959).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9736970
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9736970).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9736979
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9736979).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9736984
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9736984).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9736992
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9736992).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9737710
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9737710).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9823952
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9823952).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-9823959
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-9823959).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-NUL-2186736
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-NUL-2186736).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-NUL-2186755
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-NUL-2186755).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-NUL-9625758
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-NUL-9625758).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-3315483
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-3315483).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005654 nucleoplasm
TAS
Reactome:R-HSA-2187395
ACCEPT
Summary: Nucleoplasm localization (Reactome:R-HSA-2187395).
Reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-1549526
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-1549526).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-170847
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-170847).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-170850
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-170850).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-170868
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-170868).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-173488
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-173488).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2031355
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2031355).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2106579
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2106579).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187355
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187355).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187401
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187401).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187405
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187405).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-3311014
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-3311014).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9008928
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-9008928).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9009910
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-9009910).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0000987 cis-regulatory region sequence-specific DNA binding
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:21947082).
Reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
GO:0003700 DNA-binding transcription factor activity
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:21947082).
Reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
GO:0005515 protein binding
IPI
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21947082). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005634 nucleus
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: Nucleus localization confirmed (PMID:21947082). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005667 transcription regulator complex
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: Transcription regulator complex confirmed (PMID:21947082). SMAD3 forms complexes with SMAD4 and cofactors.
Reason: Correct. SMAD3/SMAD4 heteromeric complexes function as transcription regulator complexes.
GO:0005737 cytoplasm
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: Cytoplasm localization confirmed (PMID:21947082). SMAD3 resides in cytoplasm before activation.
Reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:21947082).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0071141 SMAD protein complex
IDA
PMID:21947082
USP15 is a deubiquitylating enzyme for receptor-activated SM...
ACCEPT
Summary: SMAD protein complex confirmed (PMID:21947082).
Reason: Correct. SMAD3 forms SMAD protein complexes including homo-oligomers and hetero-complexes with SMAD4.
GO:0005515 protein binding
IPI
PMID:21307346
Response gene to complement 32 interacts with Smad3 to promo...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21307346). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0010718 positive regulation of epithelial to mesenchymal transition
IDA
PMID:21307346
Response gene to complement 32 interacts with Smad3 to promo...
KEEP AS NON CORE
Summary: Positive regulation of EMT (PMID:21307346).
Reason: Valid downstream effect. TGF-beta/SMAD3 is a major driver of EMT. Non-core but well-documented process.
GO:0005515 protein binding
IPI
PMID:19289081
Nuclear export of Smad2 and Smad3 by RanBP3 facilitates term...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19289081). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:9865696
SARA, a FYVE domain protein that recruits Smad2 to the TGFbe...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:9865696). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:9892009
Roles of pathway-specific and inhibitory Smads in activin re...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:9892009). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0032924 activin receptor signaling pathway
IMP
PMID:15150278
Nodal and ALK7 inhibit proliferation and induce apoptosis in...
ACCEPT
Summary: Activin receptor signaling pathway confirmed (PMID:15150278).
Reason: Core biological process. SMAD3 mediates activin signaling through ACVR1B.
GO:0038092 nodal signaling pathway
IMP
PMID:15150278
Nodal and ALK7 inhibit proliferation and induce apoptosis in...
ACCEPT
Summary: Nodal signaling pathway (PMID:15150278).
Reason: SMAD3 also mediates Nodal signaling, which uses the same R-SMAD pathway as activin/TGF-beta.
GO:0005829 cytosol
TAS
Reactome:R-HSA-170835
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-170835).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187358
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187358).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187368
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187368).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187375
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187375).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187382
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187382).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-2187395
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-2187395).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-3656523
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-3656523).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9008692
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-9008692).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005829 cytosol
TAS
Reactome:R-HSA-9727922
ACCEPT
Summary: Cytosol localization (Reactome:R-HSA-9727922).
Reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
GO:0005515 protein binding
IPI
PMID:10681527
Identification and characterization of a PDZ protein that in...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:10681527). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0043130 ubiquitin binding
IDA
PMID:18794808
Ligand-dependent ubiquitination of Smad3 is regulated by cas...
ACCEPT
Summary: Ubiquitin binding confirmed (PMID:18794808). SMAD3 is subject to monoubiquitination that regulates DNA binding.
Reason: Well-supported by PMID:18794808 and PMID:21947082. Monoubiquitination of SMAD3 prevents DNA binding; USP15 deubiquitination restores it.
GO:0005515 protein binding
IPI
PMID:17469184
Involvement of the constitutive complex formation of c-Ski/S...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17469184). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:18832382
HMGA2 and Smads co-regulate SNAIL1 expression during inducti...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18832382). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0045944 positive regulation of transcription by RNA polymerase II
IDA
PMID:18832382
HMGA2 and Smads co-regulate SNAIL1 expression during inducti...
ACCEPT
Summary: Positive regulation of Pol II transcription confirmed (PMID:18832382).
Reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
GO:0071141 SMAD protein complex
IDA
PMID:18832382
HMGA2 and Smads co-regulate SNAIL1 expression during inducti...
ACCEPT
Summary: SMAD protein complex confirmed (PMID:18832382).
Reason: Correct. SMAD3 forms SMAD protein complexes including homo-oligomers and hetero-complexes with SMAD4.
GO:0005515 protein binding
IPI
PMID:15588252
The Smad3 linker region contains a transcriptional activatio...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15588252). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0008270 zinc ion binding
IDA
PMID:12686552
Features of a Smad3 MH1-DNA complex. Roles of water and zinc...
ACCEPT
Summary: Zinc ion binding confirmed (PMID:12686552). SMAD3 MH1 domain coordinates zinc via Cys and His residues.
Reason: Crystal structure confirms zinc coordination in MH1 domain essential for DNA binding (PMID:12686552).
GO:0019902 phosphatase binding
IPI
PMID:16751101
PPM1A functions as a Smad phosphatase to terminate TGFbeta s...
ACCEPT
Summary: Phosphatase binding. SMAD3 binds PPM1A (PMID:16751101).
Reason: Well-documented. PPM1A dephosphorylates SMAD3 C-terminal SSXS motif to terminate TGF-beta signaling (PMID:16751101).
GO:0061629 RNA polymerase II-specific DNA-binding transcription factor binding
IPI
PMID:17251190
The MH1 domain of Smad3 interacts with Pax6 and represses au...
ACCEPT
Summary: SMAD3 binds Pol II-specific DNA-binding TFs including c-Jun, c-Fos, PAX6 (PMID:17251190).
Reason: Core molecular function. SMAD3 cooperates with other TFs at composite promoter elements.
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta, through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos
GO:0005515 protein binding
IPI
PMID:18568018
TAZ controls Smad nucleocytoplasmic shuttling and regulates ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18568018). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0031625 ubiquitin protein ligase binding
IPI
PMID:11278251
Smurf1 interacts with transforming growth factor-beta type I...
ACCEPT
Summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP (PMID:11278251).
Reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
GO:0005515 protein binding
IPI
PMID:15897867
Oligomerization of Evi-1 regulated by the PR domain contribu...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15897867). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:17099224
Potentiation of Smad-mediated transcriptional activation by ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17099224). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:15647271
The integral inner nuclear membrane protein MAN1 physically ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15647271). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005637 nuclear inner membrane
IDA
PMID:15647271
The integral inner nuclear membrane protein MAN1 physically ...
KEEP AS NON CORE
Summary: Nuclear inner membrane localization (PMID:15647271). SMAD3 co-localizes with LEMD3/MAN1.
Reason: SMAD3 interacts with LEMD3/MAN1 at the nuclear inner membrane, which sequesters SMAD3 to antagonize signaling. Specialized localization.
GO:0005515 protein binding
IPI
PMID:19049980
SKI and MEL1 cooperate to inhibit transforming growth factor...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19049980). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0019901 protein kinase binding
IPI
PMID:12874272
Requirement of the co-repressor homeodomain-interacting prot...
ACCEPT
Summary: Protein kinase binding. SMAD3 binds CDK8/CDK9 (PMID:12874272).
Reason: SMAD3 interacts with kinases including CDK8/9 that phosphorylate its linker region.
GO:0010718 positive regulation of epithelial to mesenchymal transition
IMP
PMID:18505915
Transforming growth factor-beta1, transforming growth factor...
KEEP AS NON CORE
Summary: Positive regulation of EMT (PMID:18505915).
Reason: Valid downstream effect. TGF-beta/SMAD3 is a major driver of EMT. Non-core but well-documented process.
GO:0045216 cell-cell junction organization
IMP
PMID:18505915
Transforming growth factor-beta1, transforming growth factor...
KEEP AS NON CORE
Summary: Cell-cell junction organization (PMID:18505915). Related to EMT.
Reason: Valid downstream effect related to TGF-beta-induced EMT. Non-core pleiotropic process.
GO:0005515 protein binding
IPI
PMID:16200078
Cloning and functional characterization of a new Ski homolog...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:16200078). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0005515 protein binding
IPI
PMID:17292623
Fussel-15, a novel Ski/Sno homolog protein, antagonizes BMP ...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17292623). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0070410 co-SMAD binding
IPI
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
ACCEPT
Summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric complex (PMID:8774881).
Reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
Supporting Evidence:
PMID:9311995
Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
GO:0070410 co-SMAD binding
IPI
PMID:9111321
Heteromeric and homomeric interactions correlate with signal...
ACCEPT
Summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric complex (PMID:9111321).
Reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
Supporting Evidence:
PMID:9311995
Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
GO:0070410 co-SMAD binding
IPI
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric complex (PMID:9311995).
Reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
Supporting Evidence:
PMID:9311995
Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
GO:0070410 co-SMAD binding
IPI
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric complex (PMID:9732876).
Reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
Supporting Evidence:
PMID:9311995
Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
GO:0070412 R-SMAD binding
IPI
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: R-SMAD binding confirmed. SMAD3 binds other R-SMADs (SMAD2) and itself (PMID:9311995).
Reason: Well-supported interaction. SMAD3 interacts with SMAD2 as part of the TGF-beta signaling pathway.
Supporting Evidence:
PMID:9311995
we observed TbetaR-activation-dependent interaction between Smad2 and Smad3
GO:0000976 transcription cis-regulatory region binding
IDA
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: SMAD3 directly binds to TPA-responsive elements (TREs/AP-1 sites) in promoter cis-regulatory regions. Zhang et al. (1998) showed that Smad3 interacts directly with the TRE and can activate TGF-beta-inducible transcription from the TRE (PMID:9732876).
Reason: Core molecular function. SMAD3 directly binds DNA at cis-regulatory regions including SMAD binding elements (SBEs) and TPA-responsive elements (TREs). This is supported by direct assay evidence from Zhang et al. (1998) in Nature.
Supporting Evidence:
PMID:9732876
Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription from the TRE in the absence of c-Jun and c-Fos.
GO:0003700 DNA-binding transcription factor activity
IDA
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:9732876).
Reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
GO:0007179 transforming growth factor beta receptor signaling pathway
IDA
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: TGF-beta receptor signaling pathway confirmed (PMID:9732876).
Reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
GO:0045893 positive regulation of DNA-templated transcription
IDA
PMID:9732876
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-be...
ACCEPT
Summary: SMAD3 activates DNA-templated transcription. Zhang et al. (1998) showed that Smad3 and Smad4 cooperate with c-Jun/c-Fos to activate transcription in response to TGF-beta through TPA-responsive elements (PMID:9732876).
Reason: Core biological process. SMAD3 is a transcriptional activator of TGF-beta target genes. Direct assay evidence demonstrates SMAD3 activates transcription both independently and in cooperation with AP-1 factors.
Supporting Evidence:
PMID:9732876
Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta, through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos.
GO:0003700 DNA-binding transcription factor activity
IDA
PMID:9111321
Heteromeric and homomeric interactions correlate with signal...
ACCEPT
Summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:9111321).
Reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
GO:0005160 transforming growth factor beta receptor binding
IPI
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: SMAD3 binds the TGF-beta type I receptor. Nakao et al. (1997) showed that Smad3 interacted with the kinase-deficient TGF-beta type I receptor after it was phosphorylated by TbetaR-II kinase (PMID:9311995).
Reason: Core molecular function. SMAD3 physically interacts with the activated TGF-beta type I receptor (ALK5/TGFBR1) as an essential step in signal transduction. This interaction is required for SMAD3 phosphorylation and pathway activation.
Supporting Evidence:
PMID:9311995
Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was phosphorylated by TbetaR-II kinase.
GO:0005634 nucleus
IDA
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: Nucleus localization confirmed (PMID:9311995). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005737 cytoplasm
IDA
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: Cytoplasm localization confirmed (PMID:9311995). SMAD3 resides in cytoplasm before activation.
Reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
GO:0031053 primary miRNA processing
TAS
PMID:19018011
Holding their own: the noncanonical roles of Smad proteins.
KEEP AS NON CORE
Summary: SMAD3 has been reported to participate in primary miRNA processing through noncanonical roles. Davis et al. showed that SMADs directly interact with DROSHA and facilitate maturation of specific miRNAs (PMID:18548003, PMID:19018011).
Reason: Valid but represents a noncanonical function of SMAD3. While SMAD proteins can interact with the DROSHA/DGCR8 microprocessor complex to promote processing of specific pri-miRNAs (e.g., miR-21), this is a secondary function distinct from the core TGF-beta signal transduction and transcriptional regulation roles.
GO:0042060 wound healing
TAS
PMID:19018011
Holding their own: the noncanonical roles of Smad proteins.
KEEP AS NON CORE
Summary: SMAD3 participates in wound healing through TGF-beta signaling. TGF-beta/SMAD3 signaling is a major regulator of wound healing and fibrosis responses (PMID:19018011).
Reason: Valid downstream biological outcome of TGF-beta/SMAD3 signaling. Wound healing is a well-known physiological context in which TGF-beta/SMAD3 signaling plays a role, but represents a tissue-level process rather than a core molecular or signaling function.
GO:0045893 positive regulation of DNA-templated transcription
IDA
PMID:9111321
Heteromeric and homomeric interactions correlate with signal...
ACCEPT
Summary: SMAD3 positively regulates DNA-templated transcription. Shi et al. (1997) demonstrated heteromeric and homomeric interactions of Smad3 and Smad4/DPC4 that correlate with signaling activity and functional cooperativity in transcriptional activation (PMID:9111321).
Reason: Core biological process. Positive regulation of transcription is a primary function of SMAD3. This IDA evidence is consistent with other accepted annotations for this GO term.
GO:0045893 positive regulation of DNA-templated transcription
IDA
PMID:9311995
TGF-beta receptor-mediated signalling through Smad2, Smad3 a...
ACCEPT
Summary: SMAD3 activates transcription in TGF-beta signaling. Nakao et al. (1997) showed that Smads 2, 3 and 4 showed a synergistic effect in a transcriptional reporter assay using the TGF-beta-inducible PAI-1 promoter (PMID:9311995).
Reason: Core biological process. Transcriptional activation is a primary function of SMAD3. Direct assay evidence from Nakao et al. showing synergistic transcriptional activation using a TGF-beta responsive promoter.
Supporting Evidence:
PMID:9311995
Smads 2, 3 and 4 accumulated in the nucleus upon TGF-beta1 treatment in Mv1Lu cells, and showed a synergistic effect in a transcriptional reporter assay using the TGF-beta-inducible plasminogen activator inhibitor-1 promoter.
GO:0005515 protein binding
IPI
PMID:19122240
Pin1 down-regulates transforming growth factor-beta (TGF-bet...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19122240). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0031625 ubiquitin protein ligase binding
IPI
PMID:19122240
Pin1 down-regulates transforming growth factor-beta (TGF-bet...
ACCEPT
Summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP (PMID:19122240).
Reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
GO:0006955 immune response
IMP
PMID:16886151
Association of polymorphisms of IGF1R and genes in the trans...
KEEP AS NON CORE
Summary: SMAD3 participates in immune regulation as a downstream effector of TGF-beta signaling. TGF-beta is a key immunomodulatory cytokine, and SMAD3-dependent signaling regulates T cell differentiation, tolerance, and inflammatory responses (PMID:16886151).
Reason: Valid but represents a downstream pleiotropic effect of TGF-beta/SMAD3 signaling rather than a core molecular function. TGF-beta/SMAD3 signaling is critical for immune homeostasis, particularly T regulatory cell differentiation and suppression of inflammatory responses, but this is a pathway-level outcome.
GO:0005515 protein binding
IPI
PMID:11274402
Inactivation of menin, a Smad3-interacting protein, blocks t...
REMOVE
Summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11274402). SMAD3 interacts with numerous proteins as a signaling hub.
Reason: 'Protein binding' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already present in the annotation set.
GO:0000122 negative regulation of transcription by RNA polymerase II
IMP
PMID:14555988
Differential regulation of TGF-beta signaling through Smad2,...
ACCEPT
Summary: SMAD3 represses transcription of specific target genes in the TGF-beta pathway. Differential regulation through SMAD2, SMAD3 and SMAD4 has been demonstrated, with SMAD3 contributing to transcriptional repression at specific promoters (PMID:14555988).
Reason: Core transcriptional function. SMAD3 acts as both a transcriptional activator and repressor depending on the target gene and cofactor context. At repressive loci, SMAD3 recruits co-repressors such as Ski/SnoN/TGIF to silence gene expression. This is consistent with other annotations for this GO term already accepted with IDA evidence.
GO:0000122 negative regulation of transcription by RNA polymerase II
IDA
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
ACCEPT
Summary: Negative regulation of transcription by Pol II confirmed (PMID:8774881). SMAD3 can repress transcription.
Reason: Valid. SMAD3 acts as transcriptional repressor at certain loci with Ski/SnoN/TGIF cofactors.
GO:0045944 positive regulation of transcription by RNA polymerase II
IDA
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
ACCEPT
Summary: Positive regulation of Pol II transcription confirmed (PMID:8774881).
Reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
GO:0001666 response to hypoxia
IMP
PMID:12411310
Cellular response to hypoxia involves signaling via Smad pro...
KEEP AS NON CORE
Summary: Cellular response to hypoxia involves signaling via Smad proteins including SMAD3 (PMID:12411310). Hypoxia activates TGF-beta/SMAD signaling, with SMAD3 acting as a mediator of hypoxia-induced gene expression changes.
Reason: Valid annotation supported by IMP evidence, but represents a stimulus-response context for SMAD3 signaling rather than a core function. SMAD3 mediates responses to hypoxia through its role in TGF-beta signaling, but hypoxia response is not a primary function of SMAD3.
GO:0032909 regulation of transforming growth factor beta2 production
IMP
PMID:12411310
Cellular response to hypoxia involves signaling via Smad pro...
KEEP AS NON CORE
Summary: SMAD3 participates in regulation of TGF-beta2 production (PMID:12411310). This represents a feedback mechanism within the TGF-beta signaling pathway, where SMAD3-dependent transcription influences production of TGF-beta2 ligand.
Reason: Valid annotation representing a downstream transcriptional outcome of SMAD3 signaling. Regulation of TGF-beta2 production is a specific transcriptional target/feedback mechanism rather than a core function. SMAD3 regulates many target genes; this is one pathway-specific output.
GO:0005634 nucleus
IDA
PMID:12446380
TGFbeta-Smad signalling in postoperative human lens epitheli...
ACCEPT
Summary: Nucleus localization confirmed (PMID:12446380). SMAD3 translocates to nucleus upon TGF-beta activation.
Reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
GO:0005737 cytoplasm
IDA
PMID:12446380
TGFbeta-Smad signalling in postoperative human lens epitheli...
ACCEPT
Summary: Cytoplasm localization confirmed (PMID:12446380). SMAD3 resides in cytoplasm before activation.
Reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
GO:0030308 negative regulation of cell growth
IDA
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
KEEP AS NON CORE
Summary: SMAD3 mediates TGF-beta-induced growth inhibition. Zhang et al. (1996) demonstrated that hMAD-3 (SMAD3) and hMAD-4 (SMAD4) synergize to induce TGF-beta-like responses including growth arrest, and that C-terminally truncated forms act as dominant-negative inhibitors of normal TGF-beta response (PMID:8774881).
Reason: Well-supported by early functional studies. TGF-beta-induced growth inhibition is a major biological outcome of SMAD3 signaling, but is a downstream cellular response rather than a core molecular or signaling function. SMAD3 mediates this through transcriptional regulation of cell cycle inhibitors like p15INK4b and p21.
Supporting Evidence:
PMID:8774881
hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses. When truncated at their carboxy termini, hMAD-3 and -4 act as dominant-negative inhibitors of the normal TGF-beta response.
GO:0042803 protein homodimerization activity
IPI
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
ACCEPT
Summary: SMAD3 forms homomeric complexes. Zhang et al. (1996) showed that hMAD-3 (SMAD3) associates with itself, and Nakao et al. (1997) demonstrated TGF-beta receptor activation-dependent interaction between SMAD2 and SMAD3, as well as SMAD3 self-association (PMID:8774881, PMID:9311995).
Reason: Valid molecular function. SMAD3 forms homotrimers via its MH2 domain, a key step in the signaling pathway. Crystal structures confirm the MH2 domain mediates trimerization. While SMAD3 primarily signals as a heterotrimer with SMAD4, homodimerization/trimerization is a well-documented intrinsic activity.
Supporting Evidence:
PMID:8774881
hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses.
GO:0043235 receptor complex
IMP
PMID:8774881
Receptor-associated Mad homologues synergize as effectors of...
KEEP AS NON CORE
Summary: SMAD3 transiently associates with the TGF-beta receptor complex. Zhang et al. (1996) showed that hMAD-3 (SMAD3) was phosphorylated and associated with the ligand-bound receptor complex (PMID:8774881). Nakao et al. (1997) confirmed that SMAD3 interacts with the activated TGF-beta type I receptor (PMID:9311995).
Reason: Valid but SMAD3 is not a permanent constituent of the receptor complex. SMAD3 transiently associates with the activated TGF-beta type I receptor for phosphorylation and then dissociates. The primary localization of SMAD3 is cytoplasm (basal) and nucleus (activated). This CC annotation is technically correct but could be misleading about stable residency.
Supporting Evidence:
PMID:8774881
hMAD-3 but not hMAD-4 was phosphorylated and associated with the ligand-bound receptor complex.
GO:0043565 sequence-specific DNA binding
IDA
PMID:10823886
Transforming growth factor beta -inducible independent bindi...
ACCEPT
Summary: SMAD3 directly binds DNA in a sequence-specific manner. The MH1 domain of SMAD3 recognizes the SMAD binding element (SBE: GTCT/AGAC) and CAGA-box variants. Structural studies confirm zinc-dependent DNA binding by the MH1 domain (PMID:10823886, PMID:12686552).
Reason: Core molecular function. Sequence-specific DNA binding through the MH1 domain is a fundamental activity of SMAD3. This is well-established by crystal structures, EMSA, and ChIP experiments. Consistent with the IEA annotation for this term already accepted.

Core Functions

SMAD3 functions as a sequence-specific DNA-binding transcription factor for RNA polymerase II. The MH1 domain directly binds SMAD binding elements (SBE: GTCT/AGAC) and CAGA-box motifs, and cooperates with coactivators (p300/CBP) or corepressors (Ski/SnoN/TGIF) to regulate target gene transcription.

Supporting Evidence:
  • PMID:9732876
    Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription from the TRE in the absence of c-Jun and c-Fos.
  • PMID:8774881
    hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses.

SMAD3 binds the activated TGF-beta type I receptor (ALK5/TGFBR1) as the principal intracellular signal transducer. Upon receptor binding and C-terminal SSXS phosphorylation, SMAD3 forms heteromeric complexes with SMAD4 and translocates to the nucleus.

Supporting Evidence:
  • PMID:9311995
    Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was phosphorylated by TbetaR-II kinase. TGF-beta1 induced phosphorylation of Smad2 and Smad3 in Mv1Lu mink lung epithelial cells.

SMAD3 binds to cis-regulatory DNA sequences including SMAD binding elements (SBE) and TPA-responsive elements (TRE/AP-1 sites), acting as a direct DNA-binding component of the TGF-beta transcriptional response.

Supporting Evidence:
  • PMID:9732876
    Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription from the TRE in the absence of c-Jun and c-Fos.

References

Manual transfer of experimentally-verified manual GO annotation data to orthologs by curator judgment of sequence similarity
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot keyword mapping
Gene Ontology annotation based on curation of immunofluorescence data
Automatic transfer of experimentally verified manual GO annotation data to orthologs using Ensembl Compara
Gene Ontology annotation of human sequence-specific DNA binding transcription factors (DbTFs) based on the TFClass database
Electronic Gene Ontology annotations created by ARBA machine learning models
Combined Automated Annotation using Multiple IEA Methods
Identification and characterization of a PDZ protein that interacts with activin type II receptors.
Transforming growth factor beta -inducible independent binding of SMAD to the Smad7 promoter.
Inactivation of menin, a Smad3-interacting protein, blocks transforming growth factor type beta signaling.
Smurf1 interacts with transforming growth factor-beta type I receptor through Smad7 and induces receptor degradation.
Ski-interacting protein interacts with Smad proteins to augment transforming growth factor-beta-dependent transcription.
The adaptor molecule Disabled-2 links the transforming growth factor beta receptors to the Smad pathway.
Smad3 allostery links TGF-beta receptor kinase activation to transcriptional control.
Cellular response to hypoxia involves signaling via Smad proteins.
TGFbeta-Smad signalling in postoperative human lens epithelial cells.
Identification of three novel Smad binding proteins involved in cell polarity.
Features of a Smad3 MH1-DNA complex. Roles of water and zinc in DNA binding.
Direct interaction of Ski with either Smad3 or Smad4 is necessary and sufficient for Ski-mediated repression of transforming growth factor-beta signaling.
Requirement of the co-repressor homeodomain-interacting protein kinase 2 for ski-mediated inhibition of bone morphogenetic protein-induced transcriptional activation.
Identification of glucocorticoid receptor domains involved in transrepression of transforming growth factor-beta action.
DACH1 inhibits transforming growth factor-beta signaling through binding Smad4.
Differential regulation of TGF-beta signaling through Smad2, Smad3 and Smad4.
A novel E1A-like inhibitor of differentiation (EID) family member, EID-2, suppresses transforming growth factor (TGF)-beta signaling by blocking TGF-beta-induced formation of Smad3-Smad4 complexes.
Atrophin-1-interacting protein 4/human Itch is a ubiquitin E3 ligase for human enhancer of filamentation 1 in transforming growth factor-beta signaling pathways.
Integration of Smad and forkhead pathways in the control of neuroepithelial and glioblastoma cell proliferation.
Activin receptor-like kinase-7 induces apoptosis through activation of MAPKs in a Smad3-dependent mechanism in hepatoma cells.
Nodal and ALK7 inhibit proliferation and induce apoptosis in human trophoblast cells.
Functional proteomics mapping of a human signaling pathway.
Hepatitis C viral proteins interact with Smad3 and differentially regulate TGF-beta/Smad3-mediated transcriptional activation.
Structural basis of heteromeric smad protein assembly in TGF-beta signaling.
Proteomics-based identification of proteins interacting with Smad3: SREBP-2 forms a complex with Smad3 and inhibits its transcriptional activity.
The Smad3 linker region contains a transcriptional activation domain.
The integral inner nuclear membrane protein MAN1 physically interacts with the R-Smad proteins to repress signaling by the transforming growth factor-{beta} superfamily of cytokines.
CHIP controls the sensitivity of transforming growth factor-beta signaling by modulating the basal level of Smad3 through ubiquitin-mediated degradation.
Nuclear targeting of transforming growth factor-beta-activated Smad complexes.
Oligomerization of Evi-1 regulated by the PR domain contributes to recruitment of corepressor CtBP.
Hepatitis C virus core variants isolated from liver tumor but not from adjacent non-tumor tissue interact with Smad3 and inhibit the TGF-beta pathway.
Cloning and functional characterization of a new Ski homolog, Fussel-18, specifically expressed in neuronal tissues.
PPM1A functions as a Smad phosphatase to terminate TGFbeta signaling.
The novel PIAS-like protein hZimp10 enhances Smad transcriptional activity.
Association of polymorphisms of IGF1R and genes in the transforming growth factor- beta /bone morphogenetic protein pathway with bacteremia in sickle cell anemia.
Potentiation of Smad-mediated transcriptional activation by the RNA-binding protein RBPMS.
The MH1 domain of Smad3 interacts with Pax6 and represses autoregulation of the Pax6 P1 promoter.
Fussel-15, a novel Ski/Sno homolog protein, antagonizes BMP signaling.
Involvement of the constitutive complex formation of c-Ski/SnoN with Smads in the impaired negative feedback regulation of transforming growth factor beta signaling in scleroderma fibroblasts.
Msk is required for nuclear import of TGF-{beta}/BMP-activated Smads.
Transforming growth factor-beta1, transforming growth factor-beta2, and transforming growth factor-beta3 enhance ovarian cancer metastatic potential by inducing a Smad3-dependent epithelial-to-mesenchymal transition.
SMAD proteins control DROSHA-mediated microRNA maturation.
TAZ controls Smad nucleocytoplasmic shuttling and regulates human embryonic stem-cell self-renewal.
Identification of novel Smad2 and Smad3 associated proteins in response to TGF-beta1.
Ligand-dependent ubiquitination of Smad3 is regulated by casein kinase 1 gamma 2, an inhibitor of TGF-beta signaling.
HMGA2 and Smads co-regulate SNAIL1 expression during induction of epithelial-to-mesenchymal transition.
Holding their own: the noncanonical roles of Smad proteins.
Ski co-repressor complexes maintain the basal repressed state of the TGF-beta target gene, SMAD7, via HDAC3 and PRMT5.
SKI and MEL1 cooperate to inhibit transforming growth factor-beta signal in gastric cancer cells.
Pin1 down-regulates transforming growth factor-beta (TGF-beta) signaling by inducing degradation of Smad proteins.
FAM/USP9x, a deubiquitinating enzyme essential for TGFbeta signaling, controls Smad4 monoubiquitination.
Nuclear export of Smad2 and Smad3 by RanBP3 facilitates termination of TGF-beta signaling.
MiR-21 regulates adipogenic differentiation through the modulation of TGF-beta signaling in mesenchymal stem cells derived from human adipose tissue.
MTMR4 attenuates transforming growth factor beta (TGFbeta) signaling by dephosphorylating R-Smads in endosomes.
TMEPAI, a transmembrane TGF-beta-inducible protein, sequesters Smad proteins from active participation in TGF-beta signaling.
An atlas of combinatorial transcriptional regulation in mouse and man.
IL-37 is a fundamental inhibitor of innate immunity.
The Crumbs complex couples cell density sensing to Hippo-dependent control of the TGF-Ξ²-SMAD pathway.
Selective targeting of activating and inhibitory Smads by distinct WWP2 ubiquitin ligase isoforms differentially modulates TGFΞ² signalling and EMT.
Homeodomain protein DLX4 counteracts key transcriptional control mechanisms of the TGF-Ξ² cytostatic program and blocks the antiproliferative effect of TGF-Ξ².
Response gene to complement 32 interacts with Smad3 to promote epithelial-mesenchymal transition of human renal tubular cells.
Nodal enhances the activity of FoxO3a and its synergistic interaction with Smads to regulate cyclin G2 transcription in ovarian cancer cells.
Antagonistic regulation of EMT by TIF1Ξ³ and Smad4 in mammary epithelial cells.
HEB and E2A function as SMAD/FOXH1 cofactors.
USP15 is a deubiquitylating enzyme for receptor-activated SMADs.
Toward an understanding of the protein interaction network of the human liver.
Ablation of Smurf2 reveals an inhibition in TGF-Ξ² signalling through multiple mono-ubiquitination of Smad3.
Bone morphogenetic protein-3b (BMP-3b) inhibits osteoblast differentiation via Smad2/3 pathway by counteracting Smad1/5/8 signaling.
Intercellular variation in signaling through the TGF-Ξ² pathway and its relation to cell density and cell cycle phase.
CSMD1 exhibits antitumor activity in A375 melanoma cells through activation of the Smad pathway.
Kindlin-2 mediates activation of TGF-Ξ²/Smad signaling and renal fibrosis.
Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathways via miR-17-92 cluster in carotid artery restenosis.
Hsp70 and Hsp90 oppositely regulate TGF-Ξ² signaling through CHIP/Stub1.
C18 ORF1, a novel negative regulator of transforming growth factor-Ξ² signaling.
Pin1 promotes degradation of Smad proteins and their interaction with phosphorylated tau in Alzheimer's disease.
A Smad3 and TTF-1/NKX2-1 complex regulates Smad4-independent gene expression.
A proteome-scale map of the human interactome network.
A massively parallel pipeline to clone DNA variants and examine molecular phenotypes of human disease mutations.
New host factors important for respiratory syncytial virus (RSV) replication revealed by a novel microfluidics screen for interactors of matrix (M) protein.
Reduction of miR-29c enhances pancreatic cancer cell migration and stem cell-like phenotype.
Proteomic analyses reveal distinct chromatin-associated and soluble transcription factor complexes.
14-3-3ΞΆ turns TGF-Ξ²'s function from tumor suppressor to metastasis promoter in breast cancer by contextual changes of Smad partners from p53 to Gli2.
Syntenin regulates TGF-Ξ²1-induced Smad activation and the epithelial-to-mesenchymal transition by inhibiting caveolin-mediated TGF-Ξ² type I receptor internalization.
Widespread macromolecular interaction perturbations in human genetic disorders.
MicroRNA-143 Activation Regulates Smooth Muscle and Endothelial Cell Crosstalk in Pulmonary Arterial Hypertension.
ShcA Protects against Epithelial-Mesenchymal Transition through Compartmentalized Inhibition of TGF-Ξ²-Induced Smad Activation.
MiRNA-199a-5p influences pulmonary artery hypertension via downregulating Smad3.
Suppression of innate inflammation and immunity by interleukin-37.
Pooled-matrix protein interaction screens using Barcode Fusion Genetics.
PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscle Cells, Regulating Cell Proliferation and Glucose Metabolism.
MiR-1 suppresses tumor cell proliferation in colorectal cancer by inhibition of Smad3-mediated tumor glycolysis.
IL-37 isoform D downregulates pro-inflammatory cytokines expression in a Smad3-dependent manner.
An interactome perturbation framework prioritizes damaging missense mutations for developmental disorders.
NEDD9 targets COL3A1 to promote endothelial fibrosis and pulmonary arterial hypertension.
LuTHy: a double-readout bioluminescence-based two-hybrid technology for quantitative mapping of protein-protein interactions in mammalian cells.
Novel role of the clustered miR-23b-3p and miR-27b-3p in enhanced expression of fibrosis-associated genes by targeting TGFBR3 in atrial fibroblasts.
LRP1 Deficiency in Vascular SMC Leads to Pulmonary Arterial Hypertension That Is Reversed by PPARΞ³ Activation.
Extensive disruption of protein interactions by genetic variants across the allele frequency spectrum in human populations.
Structural basis for distinct roles of SMAD2 and SMAD3 in FOXH1 pioneer-directed TGF-Ξ² signaling.
miR-4286/TGF-Ξ²1/Smad3-Negative Feedback Loop Ameliorated Vascular Endothelial Cell Damage by Attenuating Apoptosis and Inflammatory Response.
A reference map of the human binary protein interactome.
Interactome Mapping Provides a Network of Neurodegenerative Disease Proteins and Uncovers Widespread Protein Aggregation in Affected Brains.
Dual proteome-scale networks reveal cell-specific remodeling of the human interactome.
GDF-8 stimulates trophoblast cell invasion by inducing ALK5-SMAD2/3-mediated MMP2 expression.
The Interplay Between TGF-Ξ² Signaling and Cell Metabolism.
Systematic discovery of mutation-directed neo-protein-protein interactions in cancer.
Gain-of-function variants in SMAD4 compromise respiratory epithelial function.
Receptor-associated Mad homologues synergize as effectors of the TGF-beta response.
Heteromeric and homomeric interactions correlate with signaling activity and functional cooperativity of Smad3 and Smad4/DPC4.
TGF-beta receptor-mediated signalling through Smad2, Smad3 and Smad4.
Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
SARA, a FYVE domain protein that recruits Smad2 to the TGFbeta receptor.
Roles of pathway-specific and inhibitory Smads in activin receptor signaling.
Reactome:R-HSA-1535903
Phospho R-SMAD(SMAD2/3):CO-SMAD(SMAD4):FOXO3 binds FoxO3a-binding elements
Reactome:R-HSA-1549526
Phosphorylation of SMAD2,3 by Activin:Activin Receptor
Reactome:R-HSA-170835
An anchoring protein, ZFYVE9 (SARA), recruits SMAD2/3
Reactome:R-HSA-170847
Phosphorylated SMAD2 and SMAD3 form a complex with SMAD4
Reactome:R-HSA-170850
Phosphorylated SMAD2/3 dissociates from TGFBR
Reactome:R-HSA-170868
Activated type I receptor phosphorylates SMAD2/3 directly
Reactome:R-HSA-173481
SKI/SKIL binds SMAD complex, suppressing TGF-beta signaling
Reactome:R-HSA-173488
The SMAD2/3:SMAD4 complex transfers to the nucleus
Reactome:R-HSA-173545
Ubiquitin-dependent degradation of the SMAD complex terminates TGF-beta signaling
Reactome:R-HSA-2031355
WWTR1 binds SMAD2/3:SMAD4 heterotrimer
Reactome:R-HSA-209055
PPM1A dephosphorylates nuclear SMAD2/3
Reactome:R-HSA-2106579
WWTR1:SMAD translocates to the nucleus
Reactome:R-HSA-2127257
SMAD2/3:SMAD4 heterotrimer forms a complex with RBL1, E2F4/5 and DP1/2
Reactome:R-HSA-2176475
Phosphorylation of SMAD2 and SMAD3 linker regions by CDK8 or CDK9
Reactome:R-HSA-2176491
NEDD4L binds phosphorylated linker region of SMAD2/3
Reactome:R-HSA-2176502
NEDD4L ubiquitinates SMAD2/3
Reactome:R-HSA-2176503
Degradation of SMAD2/3:SMAD4 complex
Reactome:R-HSA-2179274
SMURF2 binds SMAD3 phosphorylated in the linker region
Reactome:R-HSA-2179276
SMURF2 monoubiquitinates SMAD3
Reactome:R-HSA-2186607
TGIF recruits HDAC1 to SMAD2/3:SMAD4 heterotrimer
Reactome:R-HSA-2186643
MEN1 binds SMAD2/3:SMAD4 heterotrimer
Reactome:R-HSA-2187309
SMAD2/3:SMAD4 heterotrimer binds SP1
Reactome:R-HSA-2187325
PARP1 ADP-ribosylates SMAD3 and SMAD4
Reactome:R-HSA-2187330
PARP1 binds SMAD2/3:SMAD4 heterotrimer
Reactome:R-HSA-2187355
PMEPA1 sequesters phosphorylated SMAD2/3
Reactome:R-HSA-2187358
PMEPA1 sequesters unphosphorylated SMAD2/3
Reactome:R-HSA-2187368
STUB1 (CHIP) ubiquitinates SMAD3
Reactome:R-HSA-2187375
SMAD3 binds STUB1 (CHIP)
Reactome:R-HSA-2187382
Degradation of ubiquitinated SMAD3
Reactome:R-HSA-2187388
PPM1A protein phosphatase binds phosphorylated SMAD2/3
Reactome:R-HSA-2187395
Dephosphorylated SMAD2/3 translocates to the cytosol
Reactome:R-HSA-2187401
MTMR4 dephosphorylates SMAD2/3
Reactome:R-HSA-2187405
MTMR4 binds phosphorylated SMAD2/3
Reactome:R-HSA-3311014
SMAD4 MH2 Domain Mutants do not bind phosphorylated SMAD2 and SMAD3
Reactome:R-HSA-3315483
Phosphorylated SMAD2/3 MH2 Domain Mutants do not bind SMAD4
Reactome:R-HSA-3656523
An anchoring protein ZFYVE9 (SARA) does not recruit SMAD2/3 to TGFB1:TGFBR2:p-TGFBR1 KD Mutants
Reactome:R-HSA-6781764
USP15 deubiquitinates SMAD1,SMAD2,SMAD3, SMAD7:SMURF,KEAP1
Reactome:R-HSA-870449
TRIM33 monoubiquitinates SMAD4
Reactome:R-HSA-870538
TRIM33 (Ectodermin) binds SMAD heterotrimer in the nucleus
Reactome:R-HSA-8878143
RUNX3 binds SMAD3 and SMAD4
Reactome:R-HSA-8878178
The complex of RUNX3, SMAD3 and SMAD4 binds the CDKN1A gene promoter
Reactome:R-HSA-8952226
RUNX3 binds the BCL2L11 (BIM) gene
Reactome:R-HSA-9008692
IL37(?-218) binds SMAD3
Reactome:R-HSA-9008928
IL37(?-218):p-S423,S425-SMAD3 translocates to the nucleus
Reactome:R-HSA-9009910
IL37(?-218) binds p-S423,S425-SMAD3
Reactome:R-HSA-9617996
FOXO1,FOXO3,FOXO4 bind p-2S-SMAD2/3:SMAD4
Reactome:R-HSA-9618004
FOXO:SMAD complex binds CDKN1A gene promoter
Reactome:R-HSA-9618021
FOXG1 binds FOXO:SMAD complexes
Reactome:R-HSA-9727922
SARS-CoV-1 SUMO1-K62-p-S177-N dimer binds to SMAD3
Reactome:R-HSA-9731111
MAPK1 and MAPK3 phosphorylate SMAD2 and SMAD3
Reactome:R-HSA-9733207
p-2S-SMAD3:p-2S-SMAD3:SMAD4 forms a complex with SP1 and EP300 at the COL1A2 gene promoter
Reactome:R-HSA-9733247
YBX1 binds COL1A2 gene promoter
Reactome:R-HSA-9736959
WWTR1:p-2S-SMAD2,3:SMAD4 binds SERPINE1 gene promoter
Reactome:R-HSA-9736970
WWTR1:p-2S-SMAD2,3:SMAD4 binds SMAD7 gene promoter
Reactome:R-HSA-9736979
p-2S-SMAD2,3:SMAD4:SP1 binds CDKN2B gene promoter
Reactome:R-HSA-9736984
p-2S-SMAD2,3:SMAD4:RBL1:E2F4,5:DP1,2 binds MYC gene promoter
Reactome:R-HSA-9736992
p-2S-SMAD2,3:SMAD4 binds JUNB gene promoter
Reactome:R-HSA-9737710
SARS-CoV-1 N binds SMAD3 and EP300 at the SERPINE1 gene promoter
Reactome:R-HSA-9823952
EOMES, SMAD2,3, and GATA6 bind the CXCR4 gene
Reactome:R-HSA-9823959
SMAD2,3 and GATA6-AS1 lncRNA bind the GATA6 gene
Reactome:R-NUL-2186736
Rnf111 binds SKI/SKIL in complex with SMAD2/3:SMAD4 upon TGF-beta stimulation
Reactome:R-NUL-2186755
Ubiquitination of SKI/SKIL by Rnf111
Reactome:R-NUL-9625758
FOXO1,FOXO3 and SMAD3 bind Trim63 gene promoter

πŸ“š Additional Documentation

Deep Research Falcon

(SMAD3-deep-research-falcon.md)

provider: falcon
model: Edison Scientific Literature
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start_time: '2026-02-14T14:34:13.797600'
end_time: '2026-02-14T14:41:48.272750'
duration_seconds: 454.48
template_file: templates/gene_research_go_focused.md
template_variables:
organism: human
gene_id: SMAD3
gene_symbol: SMAD3
uniprot_accession: P84022
protein_description: 'RecName: Full=SMAD family member 3 {ECO:0000312|HGNC:HGNC:6769};
Short=SMAD 3 {ECO:0000305}; Short=hSMAD3; AltName: Full=JV15-2; AltName: Full=Mothers
against decapentaplegic homolog 3; Short=MAD homolog 3; Short=Mad3; Short=Mothers
against DPP homolog 3; Short=hMAD-3;'
gene_info: Name=SMAD3 {ECO:0000312|HGNC:HGNC:6769}; Synonyms=MADH3;
organism_full: Homo sapiens (Human).
protein_family: Belongs to the dwarfin/SMAD family. .
protein_domains: Dwarfin. (IPR013790); MAD_homology1_Dwarfin-type. (IPR003619);
MAD_homology_MH1. (IPR013019); SMAD-like_dom_sf. (IPR017855); SMAD_dom_Dwarfin-type.
(IPR001132)
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citation_count: 26


Question

Gene Research for Functional Annotation

⚠️ CRITICAL: Gene/Protein Identification Context

BEFORE YOU BEGIN RESEARCH: You MUST verify you are researching the CORRECT gene/protein. Gene symbols can be ambiguous, especially for less well-characterized genes from non-model organisms.

Target Gene/Protein Identity (from UniProt):

  • UniProt Accession: P84022
  • Protein Description: RecName: Full=SMAD family member 3 {ECO:0000312|HGNC:HGNC:6769}; Short=SMAD 3 {ECO:0000305}; Short=hSMAD3; AltName: Full=JV15-2; AltName: Full=Mothers against decapentaplegic homolog 3; Short=MAD homolog 3; Short=Mad3; Short=Mothers against DPP homolog 3; Short=hMAD-3;
  • Gene Information: Name=SMAD3 {ECO:0000312|HGNC:HGNC:6769}; Synonyms=MADH3;
  • Organism (full): Homo sapiens (Human).
  • Protein Family: Belongs to the dwarfin/SMAD family. .
  • Key Domains: Dwarfin. (IPR013790); MAD_homology1_Dwarfin-type. (IPR003619); MAD_homology_MH1. (IPR013019); SMAD-like_dom_sf. (IPR017855); SMAD_dom_Dwarfin-type. (IPR001132)

MANDATORY VERIFICATION STEPS:

  1. Check if the gene symbol "SMAD3" matches the protein description above
  2. Verify the organism is correct: Homo sapiens (Human).
  3. Check if protein family/domains align with what you find in literature
  4. If you find literature for a DIFFERENT gene with the same or similar symbol, STOP

If Gene Symbol is Ambiguous or You Cannot Find Relevant Literature:

DO NOT PROCEED WITH RESEARCH ON A DIFFERENT GENE. Instead:
- State clearly: "The gene symbol 'SMAD3' is ambiguous or literature is limited for this specific protein"
- Explain what you found (e.g., "Found extensive literature on a different gene with the same symbol in a different organism")
- Describe the protein based ONLY on the UniProt information provided above
- Suggest that the protein function can be inferred from domain/family information

Research Target:

Please provide a comprehensive research report on the gene SMAD3 (gene ID: SMAD3, UniProt: P84022) in human.

The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.

You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.

We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.

We are interested in where in or outside the cell the gene product carries out its function.

We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.

Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.

Output

Question: You are an expert researcher providing comprehensive, well-cited information.

Provide detailed information focusing on:
1. Key concepts and definitions with current understanding
2. Recent developments and latest research (prioritize 2023-2024 sources)
3. Current applications and real-world implementations
4. Expert opinions and analysis from authoritative sources
5. Relevant statistics and data from recent studies

Format as a comprehensive research report with proper citations. Include URLs and publication dates where available.
Always prioritize recent, authoritative sources and provide specific citations for all major claims.

Gene Research for Functional Annotation

⚠️ CRITICAL: Gene/Protein Identification Context

BEFORE YOU BEGIN RESEARCH: You MUST verify you are researching the CORRECT gene/protein. Gene symbols can be ambiguous, especially for less well-characterized genes from non-model organisms.

Target Gene/Protein Identity (from UniProt):

  • UniProt Accession: P84022
  • Protein Description: RecName: Full=SMAD family member 3 {ECO:0000312|HGNC:HGNC:6769}; Short=SMAD 3 {ECO:0000305}; Short=hSMAD3; AltName: Full=JV15-2; AltName: Full=Mothers against decapentaplegic homolog 3; Short=MAD homolog 3; Short=Mad3; Short=Mothers against DPP homolog 3; Short=hMAD-3;
  • Gene Information: Name=SMAD3 {ECO:0000312|HGNC:HGNC:6769}; Synonyms=MADH3;
  • Organism (full): Homo sapiens (Human).
  • Protein Family: Belongs to the dwarfin/SMAD family. .
  • Key Domains: Dwarfin. (IPR013790); MAD_homology1_Dwarfin-type. (IPR003619); MAD_homology_MH1. (IPR013019); SMAD-like_dom_sf. (IPR017855); SMAD_dom_Dwarfin-type. (IPR001132)

MANDATORY VERIFICATION STEPS:

  1. Check if the gene symbol "SMAD3" matches the protein description above
  2. Verify the organism is correct: Homo sapiens (Human).
  3. Check if protein family/domains align with what you find in literature
  4. If you find literature for a DIFFERENT gene with the same or similar symbol, STOP

If Gene Symbol is Ambiguous or You Cannot Find Relevant Literature:

DO NOT PROCEED WITH RESEARCH ON A DIFFERENT GENE. Instead:
- State clearly: "The gene symbol 'SMAD3' is ambiguous or literature is limited for this specific protein"
- Explain what you found (e.g., "Found extensive literature on a different gene with the same symbol in a different organism")
- Describe the protein based ONLY on the UniProt information provided above
- Suggest that the protein function can be inferred from domain/family information

Research Target:

Please provide a comprehensive research report on the gene SMAD3 (gene ID: SMAD3, UniProt: P84022) in human.

The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.

You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.

We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.

We are interested in where in or outside the cell the gene product carries out its function.

We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.

Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.

Title: Functional annotation of human SMAD3 (UniProt P84022)

Plan and verification. We verified the target as human SMAD3, a receptor-regulated SMAD (R‑SMAD) in the dwarfin/SMAD family. SMAD3 comprises an N‑terminal MH1 DNA‑binding domain, a C‑terminal MH2 protein‑interaction domain, and a regulatory linker; this architecture underlies its role as a transcriptional effector of TGF‑β/Activin signaling (wang2023smadproteinsin pages 1-2, danielpour2024advancesandchallenges pages 12-13). Below we synthesize current evidence with emphasis on 2023–2025 sources and translational applications.

Key concepts and definitions. SMAD3 is the principal R‑SMAD that, upon activation by TGF‑β type I receptor (TGFBR1/ALK5), forms oligomers with the co‑SMAD, SMAD4, to regulate target genes. The MH1 domain binds short Smad‑binding elements (SBE, 5′‑GTCT/AGAC‑3β€²) and CAGA‑box variants, whereas MH2 mediates receptor docking, SMAD4 assembly, and cofactor recruitment; the linker integrates regulatory post‑translational modifications (PTMs) (wang2023smadproteinsin pages 1-2, danielpour2024advancesandchallenges pages 12-13). Because Smad core motifs are only 4 bp, SMAD3 complexes have intrinsically low DNA affinity and rely on cooperating transcription factors (e.g., AP‑1) and composite motifs to achieve specificity (itoh2024analysisofthe pages 1-2).

Recent mechanistic developments (2023–2025 priority). New analyses emphasize DNA recognition and context dependence. CASTing of endogenous Smad2/3–Smad4 complexes across cell types showed differential preference for SBE vs CAGA and frequent enrichment of Fos/Jun motifs; composite elements were required for robust TGF‑β responses, explaining cell‑type dependence (The FASEB Journal, 2024; https://doi.org/10.1096/fj.202400978r) (itoh2024analysisofthe pages 1-2). Reviews in 2024 synthesized PTM logic: ALK5 phosphorylates the SMAD3 C‑terminal SSXS motif to enable trimerization with SMAD4 and nuclear import; linker phosphorylation by ERK/MAPK and CDK8/9 tunes transcriptional output and cofactor engagement, while PPM1A dephosphorylation and RanBP3 export terminate signaling (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17040533; Pharmaceuticals, 2024; https://doi.org/10.3390/ph17030326) (danielpour2024advancesandchallenges pages 12-13, runa2024targetingsmaddependentsignaling pages 2-4, runa2024targetingsmaddependentsignaling pages 8-9). Single‑cell work highlighted distinct SMAD3 dynamics, including often higher basal nuclear localization than SMAD2 and non‑redundant ligand responses, reinforcing differential roles within the Smad2/3 pair (2024; https://doi.org/10.26083/tuprints-00028725) (huang2024investigatingsmadsignaling pages 81-84).

Biochemical role and pathway placement. In canonical TGF‑β signaling, ligand‑bound TGFBR2 phosphorylates ALK5, which then phosphorylates SMAD3 at the C‑terminal SSXS motif; phospho‑SMAD3 dissociates, oligomerizes primarily as a heterotrimer with SMAD4, and translocates to the nucleus to regulate transcription with coactivators (e.g., p300/CBP) or corepressors (e.g., TGIF, Ski/Sno) (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17040533) (danielpour2024advancesandchallenges pages 12-13). SMAD3’s linker is a regulatory hub targeted by ERK/MAPK, CDKs and other kinases; phosphatases (PPM1A, SCPs) and E3 ligases (SMURF/NEDD4L) modulate turnover and signal duration (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17030326) (runa2024targetingsmaddependentsignaling pages 8-9). Functionally, SMAD3 contributes to EMT programs and extracellular matrix gene induction; differential DNA‑binding capacity versus SMAD2 (SMAD2’s MH1 insertion restricts direct DNA binding) gives SMAD3 a prominent direct chromatin‑engagement role (danielpour2024advancesandchallenges pages 12-13).

Subcellular localization and dynamics. SMAD3 shuttles constitutively between cytoplasm and nucleus; receptor‑mediated phosphorylation exposes/imports MH1‑encoded NLSs via importin‑β pathways. Export is driven by RanBP3 and exportins when SMAD3 is dephosphorylated by PPM1A; SMAD4 import relies on importin‑α and nuclear pore components participate in flux (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17030326) (runa2024targetingsmaddependentsignaling pages 2-4). Single‑cell studies report that SMAD3 can be constitutively more nuclear than SMAD2 and that its interaction with SMAD4 differs dynamically even when SSXS sites are phosphorylated, contributing to distinct signaling kinetics (2024; https://doi.org/10.26083/tuprints-00028725) (huang2024investigatingsmadsignaling pages 81-84).

Genome binding and enhancer logic. Genome‑wide and CASTing studies show that Smad complexes bind short SBEs/CAGA motifs with low affinity and typically require adjacent TF motifs (e.g., AP‑1) at enhancers to stabilize occupancy and drive transcription, explaining strong context dependence. Reporter assays validated that composite elements potentiate cell type–specific TGF‑β responses (FASEB J, 2024; https://doi.org/10.1096/fj.202400978r) (itoh2024analysisofthe pages 1-2). Methodologically, a multiplexed TR‑FRET uHTS assay now enables simultaneous monitoring of SMAD4–SMAD3 PPI and SBE DNA binding at single‑amino‑acid resolution, and has identified initial small‑molecule hits that disrupt the SMAD4–SMAD3–DNA complex (J Mol Cell Biol, 2023; https://doi.org/10.1093/jmcb/mjad068) (ouyang2023amultiplexedtimeresolved pages 1-2).

Cellular location of function. SMAD3 functions in both cytoplasm (activation and complex assembly) and nucleus (transcriptional regulation). Its transcriptional activity depends on nuclear accumulation with SMAD4 and on engagement of coactivators/corepressors; termination involves nuclear export following dephosphorylation (runa2024targetingsmaddependentsignaling pages 2-4, danielpour2024advancesandchallenges pages 12-13).

Disease relevance and phenotypes. SMAD3 is integral to fibrotic programs and cancer biology via TGF‑β‑driven EMT and matrix deposition; perturbations in TGF‑β/SMAD signaling can support tumor progression and immune evasion, while intact signaling provides tumor‑suppressive effects in early diseaseβ€”underscoring context‑dependent roles (Diagnostics, 2023; https://doi.org/10.3390/diagnostics13172769) (wang2023smadproteinsin pages 17-19). Inherited defects within the TGF‑β–SMAD axis cause syndromic aortopathy; SMAD3 is the defining gene for Loeys–Dietz syndrome type 3, and its downstream signaling contributes to vascular remodeling and aneurysm risk across the TGF‑β pathway disorders (wang2023smadproteinsin pages 17-19). Expert immuno‑oncology perspectives further highlight TGF‑β’s control of anti‑tumor immunity, with SMAD2/3 as key intracellular mediators (Nature Reviews Immunology, 2023; https://doi.org/10.1038/s41577-022-00796-z) (wang2023smadproteinsin pages 17-19).

Therapeutics and real‑world implementations. Upstream TGF‑β pathway inhibitors include ALK5 small‑molecule inhibitors (e.g., galunisertib/LY2157299; vactosertib/TEW‑7197) and pan‑TGF‑β neutralizing antibodies/ligand traps (e.g., fresolimumab). Reviews in 2024–2025 outline their clinical exploration in oncology and fibrosis and emphasize challenges from pathway pleiotropy (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17040533) (danielpour2024advancesandchallenges pages 12-13). Novel combinatorial strategies include a dual ALK5/VEGFR2 inhibitor (TU2218) that enhances immune‑mediated antitumor effects and synergizes with checkpoint blockade in preclinical models, illustrating rational co‑targeting of TGF‑β/VEGF axes (Cancer Immunol Immunother, 2024; https://doi.org/10.1007/s00262-024-03777-4) (danielpour2024advancesandchallenges pages 12-13). Downstream targeting is emerging: a uHTS TR‑FRET platform directly screens for inhibitors of the SMAD4–SMAD3–DNA complex, identifying initial hit chemotypes and providing a route to modulate nuclear SMAD function selectively (J Mol Cell Biol, 2023; https://doi.org/10.1093/jmcb/mjad068) (ouyang2023amultiplexedtimeresolved pages 1-2). Expert reviews recommend cell‑type– and context‑guided strategies and, where possible, selective interference at SMAD nodes to mitigate systemic toxicity (wang2023smadproteinsin pages 17-19, danielpour2024advancesandchallenges pages 12-13).

Relevant statistics and data highlights. Quantitative DNA‑binding studies show Smad complexes’ low intrinsic affinity for 4‑bp SBE cores (Kd on the order of ~10^βˆ’7 M), necessitating cooperative binding at composite motifs; CASTing revealed motif‑usage shifts among A549, HepG2, and HaCaT cells, aligning with changes in Smad complex composition and partner TF availability (FASEB J, 2024; https://doi.org/10.1096/fj.202400978r) (itoh2024analysisofthe pages 1-2). Screening statistics from an FDA‑approved bioactive library using the uHTS TR‑FRET assay yielded tractable hits (gambogic/gambogenic acids), validating feasibility of high‑throughput disruption of SMAD4–SMAD3–DNA assemblies (J Mol Cell Biol, 2023; https://doi.org/10.1093/jmcb/mjad068) (ouyang2023amultiplexedtimeresolved pages 1-2).

Expert analyses and opinions. Contemporary reviews argue that because systemic TGF‑β blockade can impair tissue homeostasis, precise targeting of SMAD‑dependent mechanismsβ€”guided by single‑cell context, enhancer grammar, and PTM stateβ€”may produce better therapeutic indices in cancer and fibrosis (Pharmaceuticals, 2024; https://doi.org/10.3390/ph17040533; Diagnostics, 2023; https://doi.org/10.3390/diagnostics13172769) (wang2023smadproteinsin pages 17-19, danielpour2024advancesandchallenges pages 12-13). Immuno‑oncology perspectives underscore integrating TGF‑β/SMAD inhibition with immune checkpoint therapies to reverse exclusion and stromal barriers (Nature Reviews Immunology, 2023; https://doi.org/10.1038/s41577-022-00796-z) (wang2023smadproteinsin pages 17-19).

Primary function and cellular site of action summarized. SMAD3 is a sequence‑specific transcription factor and signal transducer. It is not an enzyme or transporter; rather, it is phosphorylated at its C‑terminal SSXS motif by ALK5, complexes with SMAD4, and regulates gene expression by binding SBE/CAGA‑containing enhancers and promoters with cofactors. Its functional activity occurs in the nucleus following cytoplasmic activation and translocation (runa2024targetingsmaddependentsignaling pages 2-4, danielpour2024advancesandchallenges pages 12-13, itoh2024analysisofthe pages 1-2).

Aspect Key facts Primary sources (authors, year; journal) URL/DOI
Identity & domains - Receptor-regulated SMAD (R-SMAD) belonging to the dwarfin/SMAD family
- Conserved MH1 (N-term, DNA-binding) and MH2 (C-term, protein interaction) domains; regulatory linker between them
Wang Q. et al., 2023; Diagnostics; Danielpour D., 2024; Pharmaceuticals (wang2023smadproteinsin pages 1-2, danielpour2024advancesandchallenges pages 12-13) doi:10.3390/diagnostics13172769; https://doi.org/10.3390/ph17040533
Canonical activation - TGF-Ξ²RI/ALK5 phosphorylates the C-terminal SSXS motif on SMAD3
- Phosphorylated SMAD3 oligomerizes with SMAD4 and translocates to nucleus; recruitment aided by adaptors (e.g., SARA)
Runa F. et al., 2024; Pharmaceuticals; Danielpour D., 2024; Pharmaceuticals (runa2024targetingsmaddependentsignaling pages 2-4, danielpour2024advancesandchallenges pages 12-13) 10.3390/ph17030326; https://doi.org/10.3390/ph17040533
DNA recognition - SMAD3/SMAD4 complexes bind SBE (GTCT/AGAC) and CAGA motifs but have low intrinsic DNA affinity
- Require cooperating TFs (e.g., AP-1) and composite motifs for stable, cell-type-specific binding
Itoh Y. et al., 2024; The FASEB Journal; Ouyang W. et al., 2023; J Mol Cell Biol (itoh2024analysisofthe pages 1-2, ouyang2023amultiplexedtimeresolved pages 1-2) https://doi.org/10.1096/fj.202400978r; https://doi.org/10.1093/jmcb/mjad068
Nuclear shuttling - Dynamic cytoplasm↔nucleus shuttling mediated by importins/exportins (importin-Ξ²/Ξ±, RanBP3) and nuclear pores
- Termination/export promoted by phosphatase PPM1A (dephosphorylates C-term)
Runa F. et al., 2024; Pharmaceuticals; Huang Z., 2024 (runa2024targetingsmaddependentsignaling pages 2-4, huang2024investigatingsmadsignaling pages 81-84) 10.3390/ph17030326; https://doi.org/10.26083/tuprints-00028725
Post-translational regulation - Linker phosphorylation by ERK/MAPK and CDK8/9 modulates transcriptional output
- Ubiquitin ligases (SMURF family, NEDD4L, etc.) drive proteasomal turnover; phosphatases (PPM1A, SCPs) reverse phosphorylation
Runa F. et al., 2024; Pharmaceuticals; Danielpour D., 2024; Pharmaceuticals (runa2024targetingsmaddependentsignaling pages 8-9, danielpour2024advancesandchallenges pages 12-13) 10.3390/ph17030326; https://doi.org/10.3390/ph17040533
Enhancer logic & context dependence - SMAD complexes often act at distal enhancers and require nearby TF motifs (AP‑1, FOXO1 etc.) to drive context-specific transcription
- Genome-wide assays (CASTing/ChIP) reveal cell-type motif preferences and composite-site dependence
Itoh Y. et al., 2024; The FASEB Journal; Ouyang W. et al., 2023; J Mol Cell Biol (itoh2024analysisofthe pages 1-2, ouyang2023amultiplexedtimeresolved pages 1-2) https://doi.org/10.1096/fj.202400978r; https://doi.org/10.1093/jmcb/mjad068
Disease genetics - Pathogenic SMAD3/TGF‑β axis variants underlie vascular/connective-tissue phenotypes (Loeys–Dietz–related aortopathy) and contribute to fibrotic and cancer biology
- Dysregulated SMAD3 signaling implicated in fibrosis, tumor progression, and immune modulation
Wang Q. et al., 2023; Diagnostics; Danielpour D., 2024; Pharmaceuticals (wang2023smadproteinsin pages 17-19, danielpour2024advancesandchallenges pages 12-13) doi:10.3390/diagnostics13172769; https://doi.org/10.1186/s12967-024-05411-4
Therapeutic applications - Upstream inhibitors: ALK5 inhibitors (galunisertib, vactosertib) and pan‑TGF‑β traps (fresolimumab) in clinical/preclinical exploration
- Dual-target strategies (e.g., ALK5/VEGFR TU2218) and direct SMAD4–SMAD3–DNA screening (TR‑FRET uHTS) enable downstream/combination approaches
Danielpour D., 2024; Pharmaceuticals; Kim N.-H. et al., 2024; Cancer Immunol Immunother; Ouyang W. et al., 2023; J Mol Cell Biol (danielpour2024advancesandchallenges pages 12-13, ouyang2023amultiplexedtimeresolved pages 1-2) https://doi.org/10.3390/ph17040533; https://doi.org/10.1007/s00262-024-03777-4; https://doi.org/10.1093/jmcb/mjad068

Table: Compact reference table summarizing SMAD3 identity, mechanism, regulation, genomic binding logic, disease links, and therapeutic strategies with primary 2023–2025 sources (pqac context IDs). This provides a quick evidence‑linked overview for functional annotation and research planning.

Citations with URLs and dates
- Wang Q. et al., 2023, Diagnostics 13:2769. SMAD proteins in TGF‑β signalling pathway in cancer. https://doi.org/10.3390/diagnostics13172769 (Accessed 2026) (wang2023smadproteinsin pages 1-2, wang2023smadproteinsin pages 17-19).
- Danielpour D., 2024, Pharmaceuticals. Advances and challenges in targeting TGF‑β isoforms. https://doi.org/10.3390/ph17040533 (Accessed 2026) (danielpour2024advancesandchallenges pages 12-13).
- Runa F. et al., 2024, Pharmaceuticals. Targeting SMAD‑dependent signaling. https://doi.org/10.3390/ph17030326 (Accessed 2026) (runa2024targetingsmaddependentsignaling pages 2-4, runa2024targetingsmaddependentsignaling pages 8-9).
- Itoh Y. et al., 2024, The FASEB Journal. DNA‑binding properties of TGF‑β‑activated Smad complexes. https://doi.org/10.1096/fj.202400978r (Accessed 2026) (itoh2024analysisofthe pages 1-2).
- Ouyang W. et al., 2023, Journal of Molecular Cell Biology. TR‑FRET uHTS assay targeting SMAD4–SMAD3–DNA. https://doi.org/10.1093/jmcb/mjad068 (Accessed 2026) (ouyang2023amultiplexedtimeresolved pages 1-2).
- Huang Z., 2024. Investigating SMAD signaling dynamics and function in single cells. https://doi.org/10.26083/tuprints-00028725 (Accessed 2026) (huang2024investigatingsmadsignaling pages 81-84).
- Nixon B.G. et al., 2023, Nature Reviews Immunology. TGFΞ² control of immune responses in cancer. https://doi.org/10.1038/s41577-022-00796-z (Accessed 2026) (wang2023smadproteinsin pages 17-19).

Limitations. While core mechanistic features are well supported, quantitative PTM kinetics and complete maps of human disease variants specific to SMAD3 were not exhaustively retrieved here; future updates should integrate curated variant databases and clinical genetics cohorts specifically for SMAD3.

References

  1. (wang2023smadproteinsin pages 1-2): Qi Wang, Fei Xiong, Guanhua Wu, Da Wang, Wenzheng Liu, Junsheng Chen, Yongqiang Qi, Bing Wang, and Yongjun Chen. Smad proteins in tgf-Ξ² signalling pathway in cancer: regulatory mechanisms and clinical applications. Diagnostics, 13:2769, Aug 2023. URL: https://doi.org/10.3390/diagnostics13172769, doi:10.3390/diagnostics13172769. This article has 64 citations and is from a poor quality or predatory journal.

  2. (danielpour2024advancesandchallenges pages 12-13): David Danielpour. Advances and challenges in targeting tgf-Ξ² isoforms for therapeutic intervention of cancer: a mechanism-based perspective. Pharmaceuticals, Apr 2024. URL: https://doi.org/10.3390/ph17040533, doi:10.3390/ph17040533. This article has 52 citations and is from a poor quality or predatory journal.

  3. (itoh2024analysisofthe pages 1-2): Yuka Itoh, Kunio Miyake, Daizo Koinuma, Chiho Omata, Masao Saitoh, and Keiji Miyazawa. Analysis of the dna‐binding properties of tgf‐β‐activated smad complexes unveils a possible molecular basis for cellular context‐dependent signaling. The FASEB Journal, Aug 2024. URL: https://doi.org/10.1096/fj.202400978r, doi:10.1096/fj.202400978r. This article has 2 citations.

  4. (runa2024targetingsmaddependentsignaling pages 2-4): Farhana Runa, Gabriela Ortiz-Soto, Natan Roberto de Barros, and Jonathan A. Kelber. Targeting smad-dependent signaling: considerations in epithelial and mesenchymal solid tumors. Pharmaceuticals, 17:326, Mar 2024. URL: https://doi.org/10.3390/ph17030326, doi:10.3390/ph17030326. This article has 17 citations and is from a poor quality or predatory journal.

  5. (runa2024targetingsmaddependentsignaling pages 8-9): Farhana Runa, Gabriela Ortiz-Soto, Natan Roberto de Barros, and Jonathan A. Kelber. Targeting smad-dependent signaling: considerations in epithelial and mesenchymal solid tumors. Pharmaceuticals, 17:326, Mar 2024. URL: https://doi.org/10.3390/ph17030326, doi:10.3390/ph17030326. This article has 17 citations and is from a poor quality or predatory journal.

  6. (huang2024investigatingsmadsignaling pages 81-84): Zixin Huang. Investigating smad signaling dynamics and function in single cells. Text, Jan 2024. URL: https://doi.org/10.26083/tuprints-00028725, doi:10.26083/tuprints-00028725. This article has 0 citations and is from a peer-reviewed journal.

  7. (ouyang2023amultiplexedtimeresolved pages 1-2): Wukun Ouyang, Qianjin Li, Qiankun Niu, Min Qui, Haian Fu, Yuhong Du, and Xiulei Mo. A multiplexed time-resolved fluorescence resonance energy transfer ultrahigh-throughput screening assay for targeting the smad4–smad3–dna complex. Journal of Molecular Cell Biology, Nov 2023. URL: https://doi.org/10.1093/jmcb/mjad068, doi:10.1093/jmcb/mjad068. This article has 11 citations and is from a peer-reviewed journal.

  8. (wang2023smadproteinsin pages 17-19): Qi Wang, Fei Xiong, Guanhua Wu, Da Wang, Wenzheng Liu, Junsheng Chen, Yongqiang Qi, Bing Wang, and Yongjun Chen. Smad proteins in tgf-Ξ² signalling pathway in cancer: regulatory mechanisms and clinical applications. Diagnostics, 13:2769, Aug 2023. URL: https://doi.org/10.3390/diagnostics13172769, doi:10.3390/diagnostics13172769. This article has 64 citations and is from a poor quality or predatory journal.

Citations

  1. itoh2024analysisofthe pages 1-2
  2. huang2024investigatingsmadsignaling pages 81-84
  3. danielpour2024advancesandchallenges pages 12-13
  4. runa2024targetingsmaddependentsignaling pages 8-9
  5. runa2024targetingsmaddependentsignaling pages 2-4
  6. ouyang2023amultiplexedtimeresolved pages 1-2
  7. wang2023smadproteinsin pages 17-19
  8. wang2023smadproteinsin pages 1-2
  9. https://doi.org/10.1096/fj.202400978r
  10. https://doi.org/10.3390/ph17040533;
  11. https://doi.org/10.3390/ph17030326
  12. https://doi.org/10.26083/tuprints-00028725
  13. https://doi.org/10.3390/ph17040533
  14. https://doi.org/10.1093/jmcb/mjad068
  15. https://doi.org/10.3390/diagnostics13172769
  16. https://doi.org/10.1038/s41577-022-00796-z
  17. https://doi.org/10.1007/s00262-024-03777-4
  18. https://doi.org/10.1096/fj.202400978r;
  19. https://doi.org/10.1186/s12967-024-05411-4
  20. https://doi.org/10.1007/s00262-024-03777-4;
  21. https://doi.org/10.3390/diagnostics13172769,
  22. https://doi.org/10.3390/ph17040533,
  23. https://doi.org/10.1096/fj.202400978r,
  24. https://doi.org/10.3390/ph17030326,
  25. https://doi.org/10.26083/tuprints-00028725,
  26. https://doi.org/10.1093/jmcb/mjad068,

πŸ“„ View Raw YAML

id: P84022
gene_symbol: SMAD3
product_type: PROTEIN
status: IN_PROGRESS
taxon:
  id: NCBITaxon:9606
  label: Homo sapiens
description: 'SMAD3 (Mothers against decapentaplegic homolog 3) is a receptor-regulated SMAD (R-SMAD) that serves as the principal
  intracellular signal transducer and transcriptional modulator in the TGF-beta/activin signaling pathway. Upon TGF-beta receptor
  activation, SMAD3 is phosphorylated at its C-terminal SSXS motif by TGF-beta type I receptor (ALK5/TGFBR1), forms heteromeric
  complexes with SMAD4 (co-SMAD), and translocates to the nucleus where it directly binds DNA at SMAD binding elements (SBE:
  GTCT/AGAC) and CAGA-box variants through its MH1 domain. SMAD3 cooperates with transcription factors such as c-Jun/c-Fos
  at composite AP-1/SMAD elements. It also participates in activin and nodal signaling. Key biological outcomes include regulation
  of cell proliferation, EMT, fibrosis, immune regulation, and cell differentiation. Germline mutations cause Loeys-Dietz
  syndrome type 3.'
alternative_products:
- name: '1'
  id: P84022-1
- name: '2'
  id: P84022-2
  sequence_note: VSP_042900
- name: '3'
  id: P84022-3
  sequence_note: VSP_043793
- name: '4'
  id: P84022-4
  sequence_note: VSP_045348
existing_annotations:
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 directly binds DNA at SMAD binding elements (SBEs) in cis-regulatory regions of target genes, demonstrated
      by crystal structure (PMID:9741623), CASTing assays (Itoh et al. 2024), and ChIP-seq studies. IBA is well-supported
      across SMAD family.
    action: ACCEPT
    reason: Core molecular function of SMAD3. The MH1 domain directly binds SBE sequences (GTCT/AGAC) in promoter/enhancer
      regions, extensively validated by structural and functional studies.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was
        phosphorylated by TbetaR-II kinase.
    - reference_id: PMID:9732876
      supporting_text: Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription
        from the TRE
- term:
    id: GO:0009653
    label: anatomical structure morphogenesis
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 is involved in anatomical structure morphogenesis via TGF-beta signaling. IBA supported across species.
    action: KEEP_AS_NON_CORE
    reason: This is a broad downstream biological process. While SMAD3 contributes to morphogenesis through TGF-beta signaling,
      this is a pleiotropic outcome rather than a core function. IBA is valid at this level for SMAD family.
- term:
    id: GO:0030154
    label: cell differentiation
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 is involved in cell differentiation through TGF-beta signaling. IBA supported.
    action: KEEP_AS_NON_CORE
    reason: Broad downstream process. TGF-beta/SMAD3 signaling regulates differentiation in many contexts (chondrocyte, osteoblast,
      T cell, etc.), but this is a pleiotropic outcome rather than core function.
- term:
    id: GO:0032924
    label: activin receptor signaling pathway
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 also mediates activin receptor signaling, being phosphorylated by ACVR1B. IBA well-supported.
    action: ACCEPT
    reason: Core biological process. SMAD3 is activated by activin type I receptor (ACVR1B/ALK4) in addition to TGF-beta receptors.
      UniProt confirms interaction with ACVR1B (PMID:9892009).
    supported_by:
    - reference_id: PMID:9892009
      supporting_text: Roles of pathway-specific and inhibitory Smads in activin receptor signaling
- term:
    id: GO:0000981
    label: DNA-binding transcription factor activity, RNA polymerase II-specific
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 is a sequence-specific DNA-binding transcription factor that activates and represses RNA polymerase II
      target genes in response to TGF-beta signaling. IBA supported across the SMAD family.
    action: ACCEPT
    reason: Core molecular function. SMAD3 directly binds DNA through its MH1 domain and recruits transcriptional coactivators
      (p300/CBP) or corepressors (Ski/SnoN, TGIF) to regulate Pol II transcription.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription
    - reference_id: PMID:21947082
      supporting_text: USP15 is required for TGFbeta and BMP responses in mammalian cells
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 is a core component of the SMAD protein signal transduction pathway. Upon TGF-beta receptor activation,
      SMAD3 is phosphorylated, complexes with SMAD4, and translocates to the nucleus. IBA well-supported.
    action: ACCEPT
    reason: Core biological process. SMAD3 is one of the principal R-SMADs mediating TGF-beta/activin signal transduction
      through the canonical SMAD pathway.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: TGF-beta induces heteromeric complexes of Smads 2, 3 and 4, and their concomitant translocation to
        the nucleus, which is required for efficient TGF-beta signal transduction
    - reference_id: PMID:8774881
      supporting_text: hMAD-3 but not hMAD-4 was phosphorylated and associated with the ligand-bound receptor complex
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 is a key intracellular mediator of TGF-beta receptor signaling. Activated by TGFBR1/ALK5 phosphorylation.
      IBA well-supported.
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated directly by the TGF-beta type I receptor and is essential for
      canonical TGF-beta signal transduction.
    supported_by:
    - reference_id: PMID:8774881
      supporting_text: The activity of hMAD-3 and -4 was regulated by the TGF-beta receptors, and hMAD-3 but not hMAD-4 was
        phosphorylated and associated with the ligand-bound receptor complex
    - reference_id: PMID:9311995
      supporting_text: Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor
- term:
    id: GO:0071144
    label: heteromeric SMAD protein complex
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 forms heteromeric SMAD complexes with SMAD4 upon TGF-beta stimulation. IBA well-supported across species.
    action: ACCEPT
    reason: Core cellular component. The SMAD3/SMAD4 heteromeric complex is the functional unit of TGF-beta signaling. Crystal
      structure solved (PMID:15350224).
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: TGF-beta induces heteromeric complexes of Smads 2, 3 and 4
- term:
    id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 positively regulates transcription by RNA polymerase II at TGF-beta target gene promoters. IBA supported.
    action: ACCEPT
    reason: Core biological process. SMAD3/SMAD4 complexes recruit p300/CBP coactivators to activate transcription of TGF-beta
      responsive genes.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta
- term:
    id: GO:0070411
    label: I-SMAD binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: SMAD3 binds inhibitory SMADs (SMAD6/SMAD7). IBA supported across the SMAD family.
    action: ACCEPT
    reason: Well-established interaction. Inhibitory SMADs compete with R-SMADs for receptor binding and recruit phosphatases/ubiquitin
      ligases. SMAD3 binds SMAD7 as part of pathway regulation.
- term:
    id: GO:0003677
    label: DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for DNA binding. SMAD3 directly binds DNA through MH1 domain at SBE sequences.
    action: ACCEPT
    reason: Correct but overly broad. More specific DNA binding terms (GO:0000978, GO:0000987) are already annotated with
      experimental evidence. This IEA is acceptable as a general parent term.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for nucleus localization. SMAD3 translocates to nucleus upon TGF-beta stimulation.
    action: ACCEPT
    reason: Well-supported. SMAD3 is found in nucleus upon activation. Multiple IDA annotations also confirm this.
- term:
    id: GO:0005667
    label: transcription regulator complex
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for transcription regulator complex. SMAD3 forms complexes with SMAD4 and other transcription factors.
    action: ACCEPT
    reason: Correct. SMAD3/SMAD4 complexes function as transcription regulator complexes. Also supported by IDA (PMID:21947082).
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for cytoplasm localization. SMAD3 resides in cytoplasm when inactive.
    action: ACCEPT
    reason: Well-supported. SMAD3 shuttles between cytoplasm and nucleus, residing primarily in cytoplasm before TGF-beta
      stimulation.
- term:
    id: GO:0006355
    label: regulation of DNA-templated transcription
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for regulation of DNA-templated transcription based on InterPro domain mapping.
    action: ACCEPT
    reason: Correct. SMAD3 is a transcription factor that regulates DNA-templated transcription. Consistent with experimental
      annotations.
- term:
    id: GO:0032502
    label: developmental process
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  review:
    summary: IEA for developmental process. Very broad term from ARBA prediction.
    action: MARK_AS_OVER_ANNOTATED
    reason: Excessively broad. While TGF-beta/SMAD3 signaling contributes to development, GO:0032502 is too general to be
      informative. More specific developmental processes are annotated elsewhere.
- term:
    id: GO:0046872
    label: metal ion binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000043
  review:
    summary: IEA for metal ion binding from UniProt keyword mapping. SMAD3 MH1 domain coordinates a zinc ion essential for
      DNA binding.
    action: MODIFY
    reason: The annotation is correct in that SMAD3 binds metal ions, but the term is too broad. The MH1 domain contains a
      zinc-binding site essential for structural integrity and DNA binding (PMID:12686552). Should be zinc ion binding (GO:0008270)
      which is already annotated with IDA.
    proposed_replacement_terms:
    - id: GO:0008270
      label: zinc ion binding
- term:
    id: GO:0071363
    label: cellular response to growth factor stimulus
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  review:
    summary: IEA for cellular response to growth factor stimulus. SMAD3 mediates TGF-beta responses.
    action: ACCEPT
    reason: Correct but broad. SMAD3 mediates cellular responses to TGF-beta and activin, which are growth factors. More specific
      pathway terms are also annotated.
- term:
    id: GO:0141091
    label: transforming growth factor beta receptor superfamily signaling pathway
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  review:
    summary: IEA for TGF-beta receptor superfamily signaling pathway from ARBA.
    action: ACCEPT
    reason: Correct. SMAD3 is a core mediator of TGF-beta receptor superfamily signaling. More specific TGF-beta pathway terms
      are also present.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:11278756
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11278756). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:11387212
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11387212). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:12154125
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12154125). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:12650946
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12650946). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:12857746
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:12857746). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:14525983
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:14525983). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15084259
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15084259). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15231748
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15231748). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15350224
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15350224). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15527767
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15527767). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:16007207
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:16007207). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:17785517
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17785517). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18548003
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18548003). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18729074
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18729074). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19032343
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19032343). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19135894
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19135894). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:20061380
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20061380). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:20211142
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20211142). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21258410
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21258410). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21297662
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21297662). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21532621
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21532621). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21597466
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21597466). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21828274
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21828274). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21988832
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21988832). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22045334
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22045334). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22442258
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22442258). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22538441
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:22538441). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25416956
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25416956). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25502805
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25502805). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25609649
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25609649). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25670079
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25670079). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25910212
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25910212). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:26680585
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:26680585). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:27107012
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:27107012). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:28471448
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:28471448). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:29892012
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29892012). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:29997244
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29997244). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:31515488
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:31515488). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:32296183
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:32296183). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:32814053
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:32814053). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:33961781
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:33961781). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:35512704
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:35512704). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:39243984
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:39243984). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:18729074
  review:
    summary: SMAD3 forms homo-oligomers. Confirmed by co-immunoprecipitation and structural studies.
    action: ACCEPT
    reason: SMAD3 homo-oligomerization is well-documented. Crystal structure shows SMAD3 trimeric assembly (PMID:11224571).
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:22045334
  review:
    summary: SMAD3 forms homo-oligomers. Confirmed by co-immunoprecipitation and structural studies.
    action: ACCEPT
    reason: SMAD3 homo-oligomerization is well-documented. Crystal structure shows SMAD3 trimeric assembly (PMID:11224571).
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
- term:
    id: GO:0000122
    label: negative regulation of transcription by RNA polymerase II
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of Pol II transcription. SMAD3 can also repress transcription
      at certain loci.
    action: ACCEPT
    reason: SMAD3 acts as both transcriptional activator and repressor depending on context and cofactors (Ski/SnoN, TGIF).
      Also supported by IDA (PMID:28467929).
- term:
    id: GO:0000165
    label: MAPK cascade
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for MAPK cascade.
    action: MARK_AS_OVER_ANNOTATED
    reason: SMAD3 is not a core component of the MAPK cascade. While MAPK phosphorylates SMAD3 linker region, and SMAD3/AP-1
      cooperate at promoters, SMAD3 does not directly participate in the MAPK cascade itself. Cross-talk annotation is misleading.
- term:
    id: GO:0000977
    label: RNA polymerase II transcription regulatory region sequence-specific DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA (Ensembl/UniProt) for RNA polymerase II transcription regulatory region sequence-specific DNA binding.
    action: ACCEPT
    reason: Correct. SMAD3 binds SBE sequences in Pol II regulatory regions. Also supported by IDA annotations.
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for DNA-binding transcription activator activity, Pol II-specific.
    action: ACCEPT
    reason: Correct. SMAD3 activates Pol II transcription at TGF-beta target genes. Also supported by multiple IDA annotations.
- term:
    id: GO:0001649
    label: osteoblast differentiation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for osteoblast differentiation from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. SMAD3 regulates osteoblast differentiation and chondrogenesis (UniProt confirms SMAD3
      is a regulator of osteogenesis). Non-core pleiotropic process.
- term:
    id: GO:0001657
    label: ureteric bud development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for ureteric bud development from mouse data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Very specific developmental process likely reflecting pleiotropic TGF-beta pathway effects in kidney development.
      No direct evidence for SMAD3-specific role in ureteric bud development in humans.
- term:
    id: GO:0001836
    label: release of cytochrome c from mitochondria
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for release of cytochrome c from mitochondria from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect. TGF-beta/SMAD3 signaling can trigger apoptosis in some contexts, which may involve
      cytochrome c release, but this is not a direct function of SMAD3.
- term:
    id: GO:0003682
    label: chromatin binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for chromatin binding from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 binds chromatin at enhancer/promoter regions. Also supported by IDA chromatin localization (PMID:21828274).
- term:
    id: GO:0003690
    label: double-stranded DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for double-stranded DNA binding from mouse data.
    action: ACCEPT
    reason: Correct. Crystal structure confirms SMAD3 MH1 domain binds dsDNA at SBE sequences (PMID:9741623, PMID:12686552).
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: IEA for DNA-binding transcription factor activity.
    action: ACCEPT
    reason: Correct. SMAD3 is a bona fide DNA-binding transcription factor. Also supported by multiple IDA annotations.
- term:
    id: GO:0005518
    label: collagen binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for collagen binding from mouse data.
    action: MARK_AS_OVER_ANNOTATED
    reason: SMAD3 regulates collagen gene expression as a transcription factor, but there is no evidence it directly binds
      collagen protein. This appears to be a misannotation or conflation of transcriptional regulation with physical binding.
- term:
    id: GO:0005886
    label: plasma membrane
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for plasma membrane localization from mouse data.
    action: KEEP_AS_NON_CORE
    reason: SMAD3 can associate with TGF-beta receptors at the plasma membrane during initial activation, but its primary
      functional locations are cytoplasm and nucleus. Plasma membrane localization is transient.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for TGF-beta receptor signaling pathway from rat orthologs.
    action: ACCEPT
    reason: Correct. Core biological process for SMAD3. Also supported by multiple IDA and IMP annotations.
- term:
    id: GO:0007254
    label: JNK cascade
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for JNK cascade from mouse data.
    action: MARK_AS_OVER_ANNOTATED
    reason: SMAD3 is not a component of the JNK cascade. While TGF-beta can activate JNK through non-canonical signaling and
      SMAD3/JUN cooperate at AP-1 sites, SMAD3 does not participate directly in JNK signal transduction.
- term:
    id: GO:0008013
    label: beta-catenin binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA annotation for beta-catenin binding via Ensembl Compara ortholog transfer. SMAD3 has been reported to interact
      with beta-catenin in the context of Wnt/TGF-beta crosstalk, but this is not a primary binding activity.
    action: MARK_AS_OVER_ANNOTATED
    reason: While SMAD3 can interact with beta-catenin in certain cellular contexts as part of Wnt/TGF-beta signaling crosstalk,
      beta-catenin binding is not a core molecular function of SMAD3. This IEA annotation likely represents a context-dependent
      interaction rather than an intrinsic binding activity.
- term:
    id: GO:0008285
    label: negative regulation of cell population proliferation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of cell population proliferation from rat data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 signaling inhibits cell proliferation through transcriptional regulation
      of CDK inhibitors (p15, p21). Also supported by IMP (PMID:14555988). Non-core pleiotropic process.
- term:
    id: GO:0010332
    label: response to gamma radiation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for response to gamma radiation from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect response. While SMAD3 may be activated or its expression altered by gamma radiation in some cell types,
      this is not a direct or core function of SMAD3.
- term:
    id: GO:0010467
    label: gene expression
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for gene expression from mouse data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Excessively broad. 'Gene expression' is too general. SMAD3 regulates transcription, which is a component of gene
      expression, but more specific terms are already annotated.
- term:
    id: GO:0019899
    label: enzyme binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for enzyme binding from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: While SMAD3 binds kinases (TGFBR1, CDKs, PDPK1), phosphatases (PPM1A), and E3 ligases, 'enzyme binding' is too
      broad. More specific binding terms are already present.
- term:
    id: GO:0023019
    label: signal transduction involved in regulation of gene expression
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for signal transduction involved in regulation of gene expression from rat data.
    action: ACCEPT
    reason: 'Correct description of SMAD3 core function: signal transduction (TGF-beta pathway) that directly regulates gene
      expression.'
- term:
    id: GO:0030325
    label: adrenal gland development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for adrenal gland development from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Very specific developmental process likely reflecting broad TGF-beta pathway pleiotropic effects. No direct evidence
      for SMAD3-specific role in human adrenal development.
- term:
    id: GO:0030335
    label: positive regulation of cell migration
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of cell migration from rat data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 signaling promotes cell migration, particularly in EMT context. UniProt
      notes SMAD3 role in EMT. Non-core pleiotropic process.
- term:
    id: GO:0030501
    label: positive regulation of bone mineralization
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of bone mineralization from rat data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. UniProt notes SMAD3 is a regulator of osteogenesis. Non-core tissue-specific process.
- term:
    id: GO:0031490
    label: chromatin DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for chromatin DNA binding from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 binds DNA within chromatin context at enhancers and promoters. Supported by ChIP studies.
- term:
    id: GO:0032332
    label: positive regulation of chondrocyte differentiation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of chondrocyte differentiation from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. UniProt notes SMAD3 is a regulator of chondrogenesis. Non-core developmental process.
- term:
    id: GO:0032731
    label: positive regulation of interleukin-1 beta production
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of IL-1 beta production from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect of TGF-beta signaling in immune regulation. Not a direct function of SMAD3 as a transcription
      factor.
- term:
    id: GO:0032916
    label: positive regulation of transforming growth factor beta3 production
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of TGF-beta3 production from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: While SMAD3 could transcriptionally regulate TGF-beta3 expression as a feedback mechanism, this is a very specific
      downstream effect. Limited direct human evidence.
- term:
    id: GO:0032993
    label: protein-DNA complex
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for protein-DNA complex from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 forms protein-DNA complexes at SBE-containing regulatory regions. Crystal structure of SMAD3 MH1-DNA
      complex solved (PMID:12686552).
- term:
    id: GO:0033689
    label: negative regulation of osteoblast proliferation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of osteoblast proliferation from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect consistent with SMAD3 role in osteogenesis regulation. Non-core tissue-specific process.
- term:
    id: GO:0036120
    label: cellular response to platelet-derived growth factor stimulus
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for cellular response to PDGF stimulus from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect. PDGF may modulate TGF-beta/SMAD signaling through MAPK crosstalk, but SMAD3 is not a direct mediator
      of PDGF responses.
- term:
    id: GO:0042110
    label: T cell activation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for T cell activation from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 signaling is critical for T cell differentiation and immune regulation.
      SMAD3 knockout mice have T cell activation defects. Non-core immune process.
- term:
    id: GO:0042177
    label: negative regulation of protein catabolic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of protein catabolic process from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect. While TGF-beta signaling can affect protein stability, this is not a direct SMAD3
      function.
- term:
    id: GO:0042220
    label: response to cocaine
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for response to cocaine from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Highly specific environmental response likely from rat neurological studies. No evidence SMAD3 directly mediates
      cocaine responses. Over-annotation from ortholog transfer.
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for identical protein binding from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 forms homomeric complexes. SMAD3 homo-oligomerization is well-documented (PMID:9670020).
- term:
    id: GO:0043066
    label: negative regulation of apoptotic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of apoptotic process from rat data.
    action: KEEP_AS_NON_CORE
    reason: Context-dependent downstream effect. TGF-beta/SMAD3 can both promote and inhibit apoptosis depending on cell type.
      Non-core pleiotropic process.
- term:
    id: GO:0043565
    label: sequence-specific DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for sequence-specific DNA binding from rat data.
    action: ACCEPT
    reason: 'Correct. SMAD3 MH1 domain binds specific DNA sequences (SBE: GTCT/AGAC). Crystal structure confirms sequence-specific
      binding (PMID:9741623).'
- term:
    id: GO:0045429
    label: positive regulation of nitric oxide biosynthetic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of nitric oxide biosynthesis from rat data.
    action: KEEP_AS_NON_CORE
    reason: Context-dependent downstream effect. TGF-beta/SMAD3 can regulate iNOS expression in certain cell types. Also supported
      by IDA (PMID:27038547). Non-core process.
- term:
    id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of Pol II transcription from rat data.
    action: ACCEPT
    reason: Correct. SMAD3 activates Pol II transcription. Also supported by IBA and IMP annotations.
- term:
    id: GO:0050728
    label: negative regulation of inflammatory response
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of inflammatory response from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 is a key anti-inflammatory pathway. Also supported by NAS (ComplexPortal).
      Non-core immune process.
- term:
    id: GO:0050776
    label: regulation of immune response
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for regulation of immune response from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid broad downstream effect. TGF-beta/SMAD3 is critical for immune regulation. Non-core pleiotropic process.
- term:
    id: GO:0050821
    label: protein stabilization
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for protein stabilization from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect effect. While SMAD3 can affect protein stability through transcriptional regulation of ubiquitin pathway
      components, protein stabilization is not a direct function.
- term:
    id: GO:0050927
    label: positive regulation of positive chemotaxis
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of positive chemotaxis from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect of TGF-beta signaling. Not a direct SMAD3 function.
- term:
    id: GO:0051496
    label: positive regulation of stress fiber assembly
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of stress fiber assembly from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect. TGF-beta can promote stress fiber formation through EMT-related cytoskeletal remodeling,
      but this is not a direct SMAD3 transcription factor function.
- term:
    id: GO:0051881
    label: regulation of mitochondrial membrane potential
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for regulation of mitochondrial membrane potential from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect. Not a direct function of SMAD3 as a transcription factor.
- term:
    id: GO:0051894
    label: positive regulation of focal adhesion assembly
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of focal adhesion assembly from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect of TGF-beta-mediated EMT or cytoskeletal changes. Not a direct SMAD3 function.
- term:
    id: GO:0060290
    label: transdifferentiation
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for transdifferentiation from rat data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 drives EMT (a form of transdifferentiation). Non-core pleiotropic process.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for SMAD protein signal transduction from rat data.
    action: ACCEPT
    reason: Correct. Core biological process. Also supported by multiple IBA and IDA annotations.
- term:
    id: GO:0061001
    label: regulation of dendritic spine morphogenesis
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for regulation of dendritic spine morphogenesis from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Very specific neuronal process. Likely indirect TGF-beta pathway effect. No direct evidence for SMAD3-specific
      role in dendritic spine morphogenesis in humans.
- term:
    id: GO:0061045
    label: negative regulation of wound healing
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of wound healing from mouse data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. UniProt notes SMAD3 has inhibitory effect on wound healing. Non-core tissue-specific
      process.
- term:
    id: GO:0061629
    label: RNA polymerase II-specific DNA-binding transcription factor binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for RNA polymerase II-specific DNA-binding TF binding from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 binds other Pol II TFs (c-Jun, c-Fos, FOXH1, etc.). Also supported by IPI annotations.
- term:
    id: GO:0061767
    label: negative regulation of lung blood pressure
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of lung blood pressure from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Very specific physiological process likely from rat cardiovascular studies. Indirect downstream effect of TGF-beta
      signaling.
- term:
    id: GO:0071333
    label: cellular response to glucose stimulus
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for cellular response to glucose stimulus from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect. TGF-beta/SMAD signaling may be modulated by metabolic conditions, but SMAD3 is not a direct glucose
      sensor.
- term:
    id: GO:0071560
    label: cellular response to transforming growth factor beta stimulus
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for cellular response to TGF-beta stimulus from rat data.
    action: ACCEPT
    reason: Correct. Core biological process. SMAD3 is a primary mediator of cellular responses to TGF-beta. Also supported
      by IDA (PMID:12902338).
- term:
    id: GO:0090263
    label: positive regulation of canonical Wnt signaling pathway
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for positive regulation of canonical Wnt signaling pathway from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: SMAD3 is not a direct component of the Wnt pathway. While there is TGF-beta/Wnt pathway crosstalk, annotating
      SMAD3 to Wnt pathway regulation is misleading.
- term:
    id: GO:0097191
    label: extrinsic apoptotic signaling pathway
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for extrinsic apoptotic signaling pathway from rat data.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta can trigger apoptosis through SMAD3-dependent transcriptional programs. Also
      supported by IMP (PMID:15334054). Non-core process.
- term:
    id: GO:0097305
    label: response to alcohol
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for response to alcohol from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Likely from rat liver/fibrosis studies where alcohol induces TGF-beta signaling. Not a direct SMAD3 function.
- term:
    id: GO:0098586
    label: cellular response to virus
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for cellular response to virus from mouse data.
    action: KEEP_AS_NON_CORE
    reason: TGF-beta/SMAD3 signaling is involved in antiviral responses and immune regulation. SARS-CoV nucleoprotein interacts
      with SMAD3 (PMID:18055455). Non-core immune process.
- term:
    id: GO:1903243
    label: negative regulation of cardiac muscle hypertrophy in response to stress
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for negative regulation of cardiac muscle hypertrophy from rat data.
    action: MARK_AS_OVER_ANNOTATED
    reason: Very specific cardiac process. Indirect TGF-beta pathway effect. Not a direct SMAD3 function.
- term:
    id: GO:1990841
    label: promoter-specific chromatin binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: IEA (Ensembl orthologs) for promoter-specific chromatin binding from mouse data.
    action: ACCEPT
    reason: Correct. SMAD3 binds chromatin at specific promoters of TGF-beta target genes.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: IDA
  original_reference_id: GO_REF:0000052
  review:
    summary: Nucleoplasm localization (GO_REF:0000052).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0006355
    label: regulation of DNA-templated transcription
  evidence_type: NAS
  original_reference_id: PMID:35359452
  review:
    summary: Regulation of DNA-templated transcription confirmed (PMID:35359452).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a transcription factor that regulates gene expression.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: NAS
  original_reference_id: PMID:35359452
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:35359452).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0032924
    label: activin receptor signaling pathway
  evidence_type: NAS
  original_reference_id: PMID:15150278
  review:
    summary: Activin receptor signaling pathway confirmed (PMID:15150278).
    action: ACCEPT
    reason: Core biological process. SMAD3 mediates activin signaling through ACVR1B.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IPI
  original_reference_id: PMID:20935647
  review:
    summary: Nucleus localization confirmed (PMID:20935647). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IPI
  original_reference_id: PMID:20935647
  review:
    summary: Cytoplasm localization confirmed (PMID:20935647). SMAD3 resides in cytoplasm before activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
- term:
    id: GO:0006355
    label: regulation of DNA-templated transcription
  evidence_type: IDA
  original_reference_id: PMID:25060702
  review:
    summary: Regulation of DNA-templated transcription confirmed (PMID:25060702).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a transcription factor that regulates gene expression.
- term:
    id: GO:0031665
    label: negative regulation of lipopolysaccharide-mediated signaling pathway
  evidence_type: NAS
  original_reference_id: PMID:29789615
  review:
    summary: Negative regulation of LPS-mediated signaling pathway (PMID:29789615). NAS from ComplexPortal.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect reflecting anti-inflammatory role of TGF-beta/SMAD3. Non-core immune process.
- term:
    id: GO:0050728
    label: negative regulation of inflammatory response
  evidence_type: NAS
  original_reference_id: PMID:27060871
  review:
    summary: Negative regulation of inflammatory response (PMID:27060871). NAS from ComplexPortal.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 is a key anti-inflammatory pathway. Non-core immune process.
- term:
    id: GO:0071635
    label: negative regulation of transforming growth factor beta production
  evidence_type: NAS
  original_reference_id: PMID:20935647
  review:
    summary: Negative regulation of TGF-beta production (PMID:20935647). NAS from ComplexPortal.
    action: KEEP_AS_NON_CORE
    reason: Valid feedback regulation. SMAD3 can modulate TGF-beta expression as part of signaling feedback. Non-core regulatory
      process.
- term:
    id: GO:0061450
    label: trophoblast cell migration
  evidence_type: IDA
  original_reference_id: PMID:34432647
  review:
    summary: Trophoblast cell migration (PMID:34432647).
    action: KEEP_AS_NON_CORE
    reason: Specific developmental process. TGF-beta/SMAD3 promotes trophoblast migration. Non-core tissue-specific process.
- term:
    id: GO:0045596
    label: negative regulation of cell differentiation
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Negative regulation of cell differentiation (GO_REF:0000024). ISS from mouse ortholog.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 can inhibit differentiation in certain contexts. Non-core pleiotropic
      process.
- term:
    id: GO:0060391
    label: positive regulation of SMAD protein signal transduction
  evidence_type: IMP
  original_reference_id: PMID:15107418
  review:
    summary: Positive regulation of SMAD protein signal transduction (PMID:15107418).
    action: ACCEPT
    reason: SMAD3 positively promotes SMAD signal transduction as an R-SMAD that forms complexes with SMAD4.
- term:
    id: GO:0097190
    label: apoptotic signaling pathway
  evidence_type: IMP
  original_reference_id: PMID:15107418
  review:
    summary: Apoptotic signaling pathway (PMID:15107418).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 can trigger apoptosis in certain contexts. Non-core pleiotropic process.
- term:
    id: GO:0030279
    label: negative regulation of ossification
  evidence_type: IDA
  original_reference_id: PMID:22155034
  review:
    summary: Negative regulation of ossification (PMID:22155034).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. UniProt notes SMAD3 inhibits early healing of bone fractures. Non-core tissue-specific
      process.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IMP
  original_reference_id: PMID:8774881
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:8774881).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IMP
  original_reference_id: PMID:30729664
  review:
    summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:30729664).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
- term:
    id: GO:1901203
    label: positive regulation of extracellular matrix assembly
  evidence_type: IMP
  original_reference_id: PMID:30729664
  review:
    summary: Positive regulation of extracellular matrix assembly (PMID:30729664).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 transcriptionally activates ECM genes (collagen, fibronectin). Non-core
      but well-documented in fibrosis context.
- term:
    id: GO:1902895
    label: positive regulation of miRNA transcription
  evidence_type: IMP
  original_reference_id: PMID:30729664
  review:
    summary: Positive regulation of miRNA transcription (PMID:30729664).
    action: KEEP_AS_NON_CORE
    reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
- term:
    id: GO:1990776
    label: response to angiotensin
  evidence_type: IMP
  original_reference_id: PMID:30729664
  review:
    summary: Response to angiotensin (PMID:30729664). Acts upstream of.
    action: KEEP_AS_NON_CORE
    reason: TGF-beta/SMAD3 signaling cross-talks with angiotensin pathways in cardiovascular/fibrotic contexts. Non-core.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IDA
  original_reference_id: PMID:9732876
  review:
    summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9732876).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
- term:
    id: GO:0031625
    label: ubiquitin protein ligase binding
  evidence_type: IPI
  original_reference_id: PMID:15781469
  review:
    summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP
      (PMID:15781469).
    action: ACCEPT
    reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:9311995
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:9311995).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IDA
  original_reference_id: PMID:9311995
  review:
    summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9311995).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
- term:
    id: GO:0001222
    label: transcription corepressor binding
  evidence_type: IPI
  original_reference_id: PMID:14612439
  review:
    summary: Transcription corepressor binding confirmed. SMAD3 interacts with corepressors including Ski, SnoN, TGIF (PMID:14612439).
    action: ACCEPT
    reason: Well-supported. SMAD3 recruits transcriptional corepressors at certain target genes.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IDA
  original_reference_id: PMID:10823886
  review:
    summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:10823886).
    action: ACCEPT
    reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:10823886
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:10823886).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IDA
  original_reference_id: PMID:10823886
  review:
    summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:10823886).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
- term:
    id: GO:0060395
    label: SMAD protein signal transduction
  evidence_type: IDA
  original_reference_id: PMID:9111321
  review:
    summary: SMAD protein signal transduction confirmed by experimental evidence (PMID:9111321).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a principal mediator of SMAD signal transduction.
- term:
    id: GO:0000987
    label: cis-regulatory region sequence-specific DNA binding
  evidence_type: IMP
  original_reference_id: PMID:32141990
  review:
    summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:32141990).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IMP
  original_reference_id: PMID:32141990
  review:
    summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:32141990).
    action: ACCEPT
    reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
- term:
    id: GO:1902894
    label: negative regulation of miRNA transcription
  evidence_type: IMP
  original_reference_id: PMID:32141990
  review:
    summary: Negative regulation of miRNA transcription (PMID:32141990).
    action: KEEP_AS_NON_CORE
    reason: Valid specific function. SMAD3 can also repress miRNA transcription at certain loci. Non-core regulatory process.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:24378993
  review:
    summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:24378993).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: IDA
  original_reference_id: PMID:24378993
  review:
    summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24378993).
    action: ACCEPT
    reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: IMP
  original_reference_id: PMID:24378993
  review:
    summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24378993).
    action: ACCEPT
    reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
- term:
    id: GO:0010629
    label: negative regulation of gene expression
  evidence_type: IMP
  original_reference_id: PMID:24378993
  review:
    summary: Negative regulation of gene expression (PMID:24378993).
    action: ACCEPT
    reason: Valid. SMAD3 can repress gene expression at certain loci through corepressor recruitment.
- term:
    id: GO:1902895
    label: positive regulation of miRNA transcription
  evidence_type: IDA
  original_reference_id: PMID:24378993
  review:
    summary: Positive regulation of miRNA transcription (PMID:24378993).
    action: KEEP_AS_NON_CORE
    reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
- term:
    id: GO:1902895
    label: positive regulation of miRNA transcription
  evidence_type: IMP
  original_reference_id: PMID:24378993
  review:
    summary: Positive regulation of miRNA transcription (PMID:24378993).
    action: KEEP_AS_NON_CORE
    reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2179276
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2179276).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15051726
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15051726). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:29899023
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:29899023). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: ISS
  original_reference_id: PMID:24964035
  review:
    summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:24964035).
    action: ACCEPT
    reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: IDA
  original_reference_id: PMID:25605017
  review:
    summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:25605017).
    action: ACCEPT
    reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
- term:
    id: GO:0032810
    label: sterol response element binding
  evidence_type: IGI
  original_reference_id: PMID:25605017
  review:
    summary: Sterol response element binding. SMAD3 binds SRE in IGI with TGF-beta1 (PMID:25605017).
    action: KEEP_AS_NON_CORE
    reason: Specialized binding activity at specific genomic loci. Non-core but documented.
- term:
    id: GO:0061629
    label: RNA polymerase II-specific DNA-binding transcription factor binding
  evidence_type: IPI
  original_reference_id: PMID:9732876
  review:
    summary: SMAD3 binds Pol II-specific DNA-binding TFs including c-Jun, c-Fos, PAX6 (PMID:9732876).
    action: ACCEPT
    reason: Core molecular function. SMAD3 cooperates with other TFs at composite promoter elements.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta,
        through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos
- term:
    id: GO:0140297
    label: DNA-binding transcription factor binding
  evidence_type: IPI
  original_reference_id: PMID:21828274
  review:
    summary: DNA-binding transcription factor binding. SMAD3 binds FOXH1 (PMID:21828274).
    action: ACCEPT
    reason: Well-supported. SMAD3 cooperates with multiple DNA-binding TFs.
- term:
    id: GO:0016922
    label: nuclear receptor binding
  evidence_type: IPI
  original_reference_id: PMID:31023188
  review:
    summary: Nuclear receptor binding. SMAD3 binds PPARgamma (PMID:31023188).
    action: ACCEPT
    reason: Well-documented interaction. SMAD3 interacts with nuclear receptors as part of transcriptional cross-talk.
- term:
    id: GO:0008285
    label: negative regulation of cell population proliferation
  evidence_type: IMP
  original_reference_id: PMID:14555988
  review:
    summary: Negative regulation of cell population proliferation confirmed (PMID:14555988).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 mediates growth arrest through CDK inhibitor induction. Non-core but important
      pleiotropic process.
- term:
    id: GO:1902893
    label: regulation of miRNA transcription
  evidence_type: IC
  original_reference_id: PMID:25605017
  review:
    summary: Regulation of miRNA transcription (PMID:25605017).
    action: KEEP_AS_NON_CORE
    reason: Valid. SMAD3 regulates miRNA transcription. Parent term of both positive and negative regulation annotations.
      Non-core.
- term:
    id: GO:0000122
    label: negative regulation of transcription by RNA polymerase II
  evidence_type: IDA
  original_reference_id: PMID:28467929
  review:
    summary: Negative regulation of transcription by Pol II confirmed (PMID:28467929). SMAD3 can repress transcription.
    action: ACCEPT
    reason: Valid. SMAD3 acts as transcriptional repressor at certain loci with Ski/SnoN/TGIF cofactors.
- term:
    id: GO:0001217
    label: DNA-binding transcription repressor activity
  evidence_type: IDA
  original_reference_id: PMID:28467929
  review:
    summary: DNA-binding transcription repressor activity confirmed (PMID:28467929). SMAD3 can also repress transcription
      at certain loci.
    action: ACCEPT
    reason: Valid. SMAD3 acts as both activator and repressor depending on cofactor context (Ski/SnoN, TGIF for repression).
- term:
    id: GO:0003677
    label: DNA binding
  evidence_type: IDA
  original_reference_id: PMID:28467929
  review:
    summary: DNA binding confirmed by direct assay (PMID:28467929).
    action: ACCEPT
    reason: Core molecular function. SMAD3 MH1 domain directly binds DNA.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:28467929
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:28467929).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9731111
  review:
    summary: Cytosol localization (Reactome:R-HSA-9731111).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0001228
    label: DNA-binding transcription activator activity, RNA polymerase II-specific
  evidence_type: IMP
  original_reference_id: PMID:26311719
  review:
    summary: DNA-binding transcription activator activity, Pol II-specific, confirmed (PMID:26311719).
    action: ACCEPT
    reason: Core molecular function. SMAD3 activates transcription of TGF-beta target genes.
- term:
    id: GO:1902895
    label: positive regulation of miRNA transcription
  evidence_type: IMP
  original_reference_id: PMID:26311719
  review:
    summary: Positive regulation of miRNA transcription (PMID:26311719).
    action: KEEP_AS_NON_CORE
    reason: Valid specific function. SMAD3 regulates miRNA transcription at specific loci. Non-core regulatory process.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:23723426
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:23723426). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IGI
  original_reference_id: PMID:23723426
  review:
    summary: Nucleus localization confirmed (PMID:23723426). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IGI
  original_reference_id: PMID:23723426
  review:
    summary: Cytosol localization (PMID:23723426).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:18832382
  review:
    summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:18832382).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:21145499
  review:
    summary: Nucleus localization confirmed (PMID:21145499). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IDA
  original_reference_id: PMID:21145499
  review:
    summary: Cytoplasm localization confirmed (PMID:21145499). SMAD3 resides in cytoplasm before activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:31582430
  review:
    summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:31582430).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
- term:
    id: GO:0001223
    label: transcription coactivator binding
  evidence_type: IPI
  original_reference_id: PMID:16777850
  review:
    summary: Transcription coactivator binding confirmed. SMAD3 interacts with coactivators including p300/CBP, ZMIZ1 (PMID:16777850).
    action: ACCEPT
    reason: Well-supported. SMAD3 linker region and MH2 domain recruit p300/CBP coactivators.
- term:
    id: GO:0000987
    label: cis-regulatory region sequence-specific DNA binding
  evidence_type: IC
  original_reference_id: PMID:21828274
  review:
    summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:21828274).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
- term:
    id: GO:0000785
    label: chromatin
  evidence_type: ISA
  original_reference_id: GO_REF:0000113
  review:
    summary: Chromatin localization (GO_REF:0000113). SMAD3 is found at chromatin at TGF-beta target gene loci.
    action: ACCEPT
    reason: Correct. ChIP studies confirm SMAD3 binding to chromatin at enhancer/promoter regions.
- term:
    id: GO:0000981
    label: DNA-binding transcription factor activity, RNA polymerase II-specific
  evidence_type: ISA
  original_reference_id: GO_REF:0000113
  review:
    summary: ISA from TFClass database for DNA-binding TF activity Pol II-specific.
    action: ACCEPT
    reason: Correct. SMAD3 is classified as a TF in TFClass. Also supported by IDA and IBA.
- term:
    id: GO:0000981
    label: DNA-binding transcription factor activity, RNA polymerase II-specific
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: 'SMAD3 functions as a DNA-binding transcription factor that directly binds SMAD binding elements (SBE: GTCT/AGAC)
      and CAGA-box motifs through its MH1 domain, and regulates RNA polymerase II-dependent transcription of TGF-beta responsive
      genes.'
    action: ACCEPT
    reason: Core molecular function. SMAD3 is a bona fide sequence-specific DNA-binding transcription factor. Its MH1 domain
      directly contacts DNA at SBE sequences, and it recruits transcriptional coactivators (p300/CBP) or corepressors (Ski/SnoN)
      to regulate Pol II transcription. Consistent with IBA and ISA evidence already accepted for this term.
- term:
    id: GO:0042307
    label: positive regulation of protein import into nucleus
  evidence_type: NAS
  original_reference_id: PMID:15799969
  review:
    summary: Positive regulation of protein import into nucleus (PMID:15799969).
    action: ACCEPT
    reason: SMAD3 promotes nuclear import of the SMAD2/3-SMAD4 complex. Part of core signaling mechanism.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25556234
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:25556234). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:18548003
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:18548003).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0017151
    label: DEAD/H-box RNA helicase binding
  evidence_type: IPI
  original_reference_id: PMID:18548003
  review:
    summary: DEAD/H-box RNA helicase binding. SMAD3 interacts with DDX5 (p68) (PMID:18548003).
    action: ACCEPT
    reason: Documented interaction. DDX5/p68 modulates SMAD3-dependent transcription.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:24613385
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:24613385). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0031962
    label: nuclear mineralocorticoid receptor binding
  evidence_type: IPI
  original_reference_id: PMID:12902338
  review:
    summary: Nuclear mineralocorticoid receptor binding. SMAD3 interacts with NR3C2 (PMID:12902338).
    action: KEEP_AS_NON_CORE
    reason: Documented interaction representing transcriptional cross-talk between TGF-beta and mineralocorticoid signaling.
- term:
    id: GO:0035259
    label: nuclear glucocorticoid receptor binding
  evidence_type: IPI
  original_reference_id: PMID:12902338
  review:
    summary: Nuclear glucocorticoid receptor binding. SMAD3 interacts with NR3C1 (PMID:12902338).
    action: KEEP_AS_NON_CORE
    reason: Documented interaction representing transcriptional cross-talk between TGF-beta and glucocorticoid signaling.
- term:
    id: GO:0071560
    label: cellular response to transforming growth factor beta stimulus
  evidence_type: IDA
  original_reference_id: PMID:12902338
  review:
    summary: Cellular response to TGF-beta stimulus confirmed (PMID:12902338).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a primary mediator of cellular TGF-beta responses.
- term:
    id: GO:0045429
    label: positive regulation of nitric oxide biosynthetic process
  evidence_type: IDA
  original_reference_id: PMID:27038547
  review:
    summary: Positive regulation of nitric oxide biosynthetic process (PMID:27038547).
    action: KEEP_AS_NON_CORE
    reason: Specific downstream effect in certain cell types. Non-core pleiotropic process.
- term:
    id: GO:0051481
    label: negative regulation of cytosolic calcium ion concentration
  evidence_type: IDA
  original_reference_id: PMID:27038547
  review:
    summary: Negative regulation of cytosolic calcium ion concentration (PMID:27038547).
    action: MARK_AS_OVER_ANNOTATED
    reason: Indirect downstream effect. SMAD3 as a transcription factor does not directly regulate calcium concentrations.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-6781764
  review:
    summary: Cytosol localization (Reactome:R-HSA-6781764).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:25893292
  review:
    summary: Nucleus localization confirmed (PMID:25893292). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0045599
    label: negative regulation of fat cell differentiation
  evidence_type: IDA
  original_reference_id: PMID:19816956
  review:
    summary: Negative regulation of fat cell differentiation (PMID:19816956).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 inhibits adipogenesis. Non-core tissue-specific process.
- term:
    id: GO:1901203
    label: positive regulation of extracellular matrix assembly
  evidence_type: IDA
  original_reference_id: PMID:21307346
  review:
    summary: Positive regulation of extracellular matrix assembly (PMID:21307346).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 transcriptionally activates ECM genes (collagen, fibronectin). Non-core
      but well-documented in fibrosis context.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:16007207
  review:
    summary: Nucleus localization confirmed (PMID:16007207). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: RNA polymerase II cis-regulatory region sequence-specific DNA binding confirmed (PMID:21947082).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBEs in Pol II-regulated cis-regulatory regions.
- term:
    id: GO:0006357
    label: regulation of transcription by RNA polymerase II
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: Regulation of transcription by RNA polymerase II confirmed (PMID:21947082).
    action: ACCEPT
    reason: Core biological process. SMAD3 regulates Pol II transcription at target genes.
- term:
    id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  evidence_type: IMP
  original_reference_id: PMID:14555988
  review:
    summary: Positive regulation of Pol II transcription confirmed (PMID:14555988).
    action: ACCEPT
    reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
- term:
    id: GO:0010628
    label: positive regulation of gene expression
  evidence_type: IDA
  original_reference_id: PMID:21307346
  review:
    summary: Positive regulation of gene expression (PMID:21307346).
    action: ACCEPT
    reason: Core biological process. SMAD3 activates expression of TGF-beta target genes.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:20129061
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:20129061). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:24627487
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:24627487). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0070412
    label: R-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:20129061
  review:
    summary: R-SMAD binding confirmed. SMAD3 binds other R-SMADs (SMAD2) and itself (PMID:20129061).
    action: ACCEPT
    reason: Well-supported interaction. SMAD3 interacts with SMAD2 as part of the TGF-beta signaling pathway.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: we observed TbetaR-activation-dependent interaction between Smad2 and Smad3
- term:
    id: GO:0000785
    label: chromatin
  evidence_type: IDA
  original_reference_id: PMID:21828274
  review:
    summary: Chromatin localization (PMID:21828274). SMAD3 is found at chromatin at TGF-beta target gene loci.
    action: ACCEPT
    reason: Correct. ChIP studies confirm SMAD3 binding to chromatin at enhancer/promoter regions.
- term:
    id: GO:0043425
    label: bHLH transcription factor binding
  evidence_type: IPI
  original_reference_id: PMID:21828274
  review:
    summary: bHLH transcription factor binding. SMAD3 binds HEB/TCF12 (PMID:21828274).
    action: ACCEPT
    reason: Documented interaction between SMAD3 and bHLH TFs at target gene promoters.
- term:
    id: GO:0071144
    label: heteromeric SMAD protein complex
  evidence_type: IDA
  original_reference_id: PMID:21828274
  review:
    summary: Heteromeric SMAD protein complex confirmed (PMID:21828274).
    action: ACCEPT
    reason: Core cellular component. SMAD3/SMAD4 heterotrimer is the functional transcriptional complex.
- term:
    id: GO:0097191
    label: extrinsic apoptotic signaling pathway
  evidence_type: IMP
  original_reference_id: PMID:15334054
  review:
    summary: Extrinsic apoptotic signaling pathway (PMID:15334054).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. Non-core pleiotropic process.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2176475
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2176475).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2176491
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2176491).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2176502
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2176502).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2176503
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2176503).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2179274
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2179274).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-1535903
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-1535903).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-173481
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-173481).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-173488
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-173488).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-173545
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-173545).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-209055
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-209055).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2106579
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2106579).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2127257
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2127257).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2186607
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2186607).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2186643
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2186643).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187309
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2187309).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187325
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2187325).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187330
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2187330).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187388
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2187388).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-870449
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-870449).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-870538
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-870538).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-8878143
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-8878143).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-8878178
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-8878178).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-8952226
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-8952226).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9617996
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9617996).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9618004
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9618004).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9618021
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9618021).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9733207
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9733207).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9733247
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9733247).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9736959
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9736959).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9736970
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9736970).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9736979
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9736979).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9736984
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9736984).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9736992
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9736992).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9737710
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9737710).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9823952
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9823952).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9823959
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-9823959).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-NUL-2186736
  review:
    summary: Nucleoplasm localization (Reactome:R-NUL-2186736).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-NUL-2186755
  review:
    summary: Nucleoplasm localization (Reactome:R-NUL-2186755).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-NUL-9625758
  review:
    summary: Nucleoplasm localization (Reactome:R-NUL-9625758).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-3315483
  review:
    summary: Cytosol localization (Reactome:R-HSA-3315483).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187395
  review:
    summary: Nucleoplasm localization (Reactome:R-HSA-2187395).
    action: ACCEPT
    reason: Correct. SMAD3 is found in nucleoplasm when in the nucleus. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-1549526
  review:
    summary: Cytosol localization (Reactome:R-HSA-1549526).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-170847
  review:
    summary: Cytosol localization (Reactome:R-HSA-170847).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-170850
  review:
    summary: Cytosol localization (Reactome:R-HSA-170850).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-170868
  review:
    summary: Cytosol localization (Reactome:R-HSA-170868).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-173488
  review:
    summary: Cytosol localization (Reactome:R-HSA-173488).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2031355
  review:
    summary: Cytosol localization (Reactome:R-HSA-2031355).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2106579
  review:
    summary: Cytosol localization (Reactome:R-HSA-2106579).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187355
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187355).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187401
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187401).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187405
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187405).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-3311014
  review:
    summary: Cytosol localization (Reactome:R-HSA-3311014).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9008928
  review:
    summary: Cytosol localization (Reactome:R-HSA-9008928).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9009910
  review:
    summary: Cytosol localization (Reactome:R-HSA-9009910).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0000987
    label: cis-regulatory region sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: Cis-regulatory region sequence-specific DNA binding confirmed (PMID:21947082).
    action: ACCEPT
    reason: Core molecular function. SMAD3 binds SBE sequences in cis-regulatory regions.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:21947082).
    action: ACCEPT
    reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21947082
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21947082). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: Nucleus localization confirmed (PMID:21947082). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005667
    label: transcription regulator complex
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: Transcription regulator complex confirmed (PMID:21947082). SMAD3 forms complexes with SMAD4 and cofactors.
    action: ACCEPT
    reason: Correct. SMAD3/SMAD4 heteromeric complexes function as transcription regulator complexes.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: Cytoplasm localization confirmed (PMID:21947082). SMAD3 resides in cytoplasm before activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:21947082).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0071141
    label: SMAD protein complex
  evidence_type: IDA
  original_reference_id: PMID:21947082
  review:
    summary: SMAD protein complex confirmed (PMID:21947082).
    action: ACCEPT
    reason: Correct. SMAD3 forms SMAD protein complexes including homo-oligomers and hetero-complexes with SMAD4.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21307346
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:21307346). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0010718
    label: positive regulation of epithelial to mesenchymal transition
  evidence_type: IDA
  original_reference_id: PMID:21307346
  review:
    summary: Positive regulation of EMT (PMID:21307346).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 is a major driver of EMT. Non-core but well-documented process.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19289081
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19289081). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:9865696
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:9865696). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:9892009
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:9892009). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0032924
    label: activin receptor signaling pathway
  evidence_type: IMP
  original_reference_id: PMID:15150278
  review:
    summary: Activin receptor signaling pathway confirmed (PMID:15150278).
    action: ACCEPT
    reason: Core biological process. SMAD3 mediates activin signaling through ACVR1B.
- term:
    id: GO:0038092
    label: nodal signaling pathway
  evidence_type: IMP
  original_reference_id: PMID:15150278
  review:
    summary: Nodal signaling pathway (PMID:15150278).
    action: ACCEPT
    reason: SMAD3 also mediates Nodal signaling, which uses the same R-SMAD pathway as activin/TGF-beta.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-170835
  review:
    summary: Cytosol localization (Reactome:R-HSA-170835).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187358
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187358).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187368
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187368).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187375
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187375).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187382
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187382).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-2187395
  review:
    summary: Cytosol localization (Reactome:R-HSA-2187395).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-3656523
  review:
    summary: Cytosol localization (Reactome:R-HSA-3656523).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9008692
  review:
    summary: Cytosol localization (Reactome:R-HSA-9008692).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-9727922
  review:
    summary: Cytosol localization (Reactome:R-HSA-9727922).
    action: ACCEPT
    reason: Correct. SMAD3 is found in the cytosol before activation. Multiple Reactome pathways confirm.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:10681527
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:10681527). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0043130
    label: ubiquitin binding
  evidence_type: IDA
  original_reference_id: PMID:18794808
  review:
    summary: Ubiquitin binding confirmed (PMID:18794808). SMAD3 is subject to monoubiquitination that regulates DNA binding.
    action: ACCEPT
    reason: Well-supported by PMID:18794808 and PMID:21947082. Monoubiquitination of SMAD3 prevents DNA binding; USP15 deubiquitination
      restores it.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:17469184
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17469184). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18832382
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18832382). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  evidence_type: IDA
  original_reference_id: PMID:18832382
  review:
    summary: Positive regulation of Pol II transcription confirmed (PMID:18832382).
    action: ACCEPT
    reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
- term:
    id: GO:0071141
    label: SMAD protein complex
  evidence_type: IDA
  original_reference_id: PMID:18832382
  review:
    summary: SMAD protein complex confirmed (PMID:18832382).
    action: ACCEPT
    reason: Correct. SMAD3 forms SMAD protein complexes including homo-oligomers and hetero-complexes with SMAD4.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15588252
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15588252). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0008270
    label: zinc ion binding
  evidence_type: IDA
  original_reference_id: PMID:12686552
  review:
    summary: Zinc ion binding confirmed (PMID:12686552). SMAD3 MH1 domain coordinates zinc via Cys and His residues.
    action: ACCEPT
    reason: Crystal structure confirms zinc coordination in MH1 domain essential for DNA binding (PMID:12686552).
- term:
    id: GO:0019902
    label: phosphatase binding
  evidence_type: IPI
  original_reference_id: PMID:16751101
  review:
    summary: Phosphatase binding. SMAD3 binds PPM1A (PMID:16751101).
    action: ACCEPT
    reason: Well-documented. PPM1A dephosphorylates SMAD3 C-terminal SSXS motif to terminate TGF-beta signaling (PMID:16751101).
- term:
    id: GO:0061629
    label: RNA polymerase II-specific DNA-binding transcription factor binding
  evidence_type: IPI
  original_reference_id: PMID:17251190
  review:
    summary: SMAD3 binds Pol II-specific DNA-binding TFs including c-Jun, c-Fos, PAX6 (PMID:17251190).
    action: ACCEPT
    reason: Core molecular function. SMAD3 cooperates with other TFs at composite promoter elements.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta,
        through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18568018
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:18568018). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0031625
    label: ubiquitin protein ligase binding
  evidence_type: IPI
  original_reference_id: PMID:11278251
  review:
    summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP
      (PMID:11278251).
    action: ACCEPT
    reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15897867
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15897867). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:17099224
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17099224). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:15647271
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:15647271). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005637
    label: nuclear inner membrane
  evidence_type: IDA
  original_reference_id: PMID:15647271
  review:
    summary: Nuclear inner membrane localization (PMID:15647271). SMAD3 co-localizes with LEMD3/MAN1.
    action: KEEP_AS_NON_CORE
    reason: SMAD3 interacts with LEMD3/MAN1 at the nuclear inner membrane, which sequesters SMAD3 to antagonize signaling.
      Specialized localization.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19049980
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19049980). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0019901
    label: protein kinase binding
  evidence_type: IPI
  original_reference_id: PMID:12874272
  review:
    summary: Protein kinase binding. SMAD3 binds CDK8/CDK9 (PMID:12874272).
    action: ACCEPT
    reason: SMAD3 interacts with kinases including CDK8/9 that phosphorylate its linker region.
- term:
    id: GO:0010718
    label: positive regulation of epithelial to mesenchymal transition
  evidence_type: IMP
  original_reference_id: PMID:18505915
  review:
    summary: Positive regulation of EMT (PMID:18505915).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect. TGF-beta/SMAD3 is a major driver of EMT. Non-core but well-documented process.
- term:
    id: GO:0045216
    label: cell-cell junction organization
  evidence_type: IMP
  original_reference_id: PMID:18505915
  review:
    summary: Cell-cell junction organization (PMID:18505915). Related to EMT.
    action: KEEP_AS_NON_CORE
    reason: Valid downstream effect related to TGF-beta-induced EMT. Non-core pleiotropic process.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:16200078
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:16200078). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:17292623
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:17292623). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0070410
    label: co-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:8774881
  review:
    summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric
      complex (PMID:8774881).
    action: ACCEPT
    reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
- term:
    id: GO:0070410
    label: co-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:9111321
  review:
    summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric
      complex (PMID:9111321).
    action: ACCEPT
    reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
- term:
    id: GO:0070410
    label: co-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:9311995
  review:
    summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric
      complex (PMID:9311995).
    action: ACCEPT
    reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
- term:
    id: GO:0070410
    label: co-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:9732876
  review:
    summary: co-SMAD binding confirmed. SMAD3 directly binds SMAD4 via MH2 domain to form the transcriptionally active heteromeric
      complex (PMID:9732876).
    action: ACCEPT
    reason: Core molecular function. SMAD3-SMAD4 interaction is the central event in TGF-beta signal transduction.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smads 2 and 3 interacted with Smad4 after TbetaR activation in transfected COS cells
- term:
    id: GO:0070412
    label: R-SMAD binding
  evidence_type: IPI
  original_reference_id: PMID:9311995
  review:
    summary: R-SMAD binding confirmed. SMAD3 binds other R-SMADs (SMAD2) and itself (PMID:9311995).
    action: ACCEPT
    reason: Well-supported interaction. SMAD3 interacts with SMAD2 as part of the TGF-beta signaling pathway.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: we observed TbetaR-activation-dependent interaction between Smad2 and Smad3
- term:
    id: GO:0000976
    label: transcription cis-regulatory region binding
  evidence_type: IDA
  original_reference_id: PMID:9732876
  review:
    summary: SMAD3 directly binds to TPA-responsive elements (TREs/AP-1 sites) in promoter cis-regulatory regions. Zhang et
      al. (1998) showed that Smad3 interacts directly with the TRE and can activate TGF-beta-inducible transcription from
      the TRE (PMID:9732876).
    action: ACCEPT
    reason: Core molecular function. SMAD3 directly binds DNA at cis-regulatory regions including SMAD binding elements (SBEs)
      and TPA-responsive elements (TREs). This is supported by direct assay evidence from Zhang et al. (1998) in Nature.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription
        from the TRE in the absence of c-Jun and c-Fos.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IDA
  original_reference_id: PMID:9732876
  review:
    summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:9732876).
    action: ACCEPT
    reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
- term:
    id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:9732876
  review:
    summary: TGF-beta receptor signaling pathway confirmed (PMID:9732876).
    action: ACCEPT
    reason: Core biological process. SMAD3 is phosphorylated by TGFBR1 and is essential for canonical TGF-beta signaling.
- term:
    id: GO:0045893
    label: positive regulation of DNA-templated transcription
  evidence_type: IDA
  original_reference_id: PMID:9732876
  review:
    summary: SMAD3 activates DNA-templated transcription. Zhang et al. (1998) showed that Smad3 and Smad4 cooperate with c-Jun/c-Fos
      to activate transcription in response to TGF-beta through TPA-responsive elements (PMID:9732876).
    action: ACCEPT
    reason: Core biological process. SMAD3 is a transcriptional activator of TGF-beta target genes. Direct assay evidence
      demonstrates SMAD3 activates transcription both independently and in cooperation with AP-1 factors.
    supported_by:
    - reference_id: PMID:9732876
      supporting_text: Smad3 and Smad4 also act together with c-Jun and c-Fos to activate transcription in response to TGF-beta,
        through a TGF-beta-inducible association of c-Jun with Smad3 and an interaction of Smad3 and c-Fos.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IDA
  original_reference_id: PMID:9111321
  qualifier: contributes_to
  review:
    summary: DNA-binding transcription factor activity confirmed by experimental evidence (PMID:9111321).
    action: ACCEPT
    reason: Core molecular function of SMAD3. Directly binds DNA and regulates transcription.
- term:
    id: GO:0005160
    label: transforming growth factor beta receptor binding
  evidence_type: IPI
  original_reference_id: PMID:9311995
  review:
    summary: SMAD3 binds the TGF-beta type I receptor. Nakao et al. (1997) showed that Smad3 interacted with the kinase-deficient
      TGF-beta type I receptor after it was phosphorylated by TbetaR-II kinase (PMID:9311995).
    action: ACCEPT
    reason: Core molecular function. SMAD3 physically interacts with the activated TGF-beta type I receptor (ALK5/TGFBR1)
      as an essential step in signal transduction. This interaction is required for SMAD3 phosphorylation and pathway activation.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was
        phosphorylated by TbetaR-II kinase.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:9311995
  review:
    summary: Nucleus localization confirmed (PMID:9311995). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IDA
  original_reference_id: PMID:9311995
  review:
    summary: Cytoplasm localization confirmed (PMID:9311995). SMAD3 resides in cytoplasm before activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
- term:
    id: GO:0031053
    label: primary miRNA processing
  evidence_type: TAS
  original_reference_id: PMID:19018011
  review:
    summary: SMAD3 has been reported to participate in primary miRNA processing through noncanonical roles. Davis et al. showed
      that SMADs directly interact with DROSHA and facilitate maturation of specific miRNAs (PMID:18548003, PMID:19018011).
    action: KEEP_AS_NON_CORE
    reason: Valid but represents a noncanonical function of SMAD3. While SMAD proteins can interact with the DROSHA/DGCR8
      microprocessor complex to promote processing of specific pri-miRNAs (e.g., miR-21), this is a secondary function distinct
      from the core TGF-beta signal transduction and transcriptional regulation roles.
- term:
    id: GO:0042060
    label: wound healing
  evidence_type: TAS
  original_reference_id: PMID:19018011
  review:
    summary: SMAD3 participates in wound healing through TGF-beta signaling. TGF-beta/SMAD3 signaling is a major regulator
      of wound healing and fibrosis responses (PMID:19018011).
    action: KEEP_AS_NON_CORE
    reason: Valid downstream biological outcome of TGF-beta/SMAD3 signaling. Wound healing is a well-known physiological context
      in which TGF-beta/SMAD3 signaling plays a role, but represents a tissue-level process rather than a core molecular or
      signaling function.
- term:
    id: GO:0045893
    label: positive regulation of DNA-templated transcription
  evidence_type: IDA
  original_reference_id: PMID:9111321
  review:
    summary: SMAD3 positively regulates DNA-templated transcription. Shi et al. (1997) demonstrated heteromeric and homomeric
      interactions of Smad3 and Smad4/DPC4 that correlate with signaling activity and functional cooperativity in transcriptional
      activation (PMID:9111321).
    action: ACCEPT
    reason: Core biological process. Positive regulation of transcription is a primary function of SMAD3. This IDA evidence
      is consistent with other accepted annotations for this GO term.
- term:
    id: GO:0045893
    label: positive regulation of DNA-templated transcription
  evidence_type: IDA
  original_reference_id: PMID:9311995
  review:
    summary: SMAD3 activates transcription in TGF-beta signaling. Nakao et al. (1997) showed that Smads 2, 3 and 4 showed
      a synergistic effect in a transcriptional reporter assay using the TGF-beta-inducible PAI-1 promoter (PMID:9311995).
    action: ACCEPT
    reason: Core biological process. Transcriptional activation is a primary function of SMAD3. Direct assay evidence from
      Nakao et al. showing synergistic transcriptional activation using a TGF-beta responsive promoter.
    supported_by:
    - reference_id: PMID:9311995
      supporting_text: Smads 2, 3 and 4 accumulated in the nucleus upon TGF-beta1 treatment in Mv1Lu cells, and showed a synergistic
        effect in a transcriptional reporter assay using the TGF-beta-inducible plasminogen activator inhibitor-1 promoter.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19122240
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:19122240). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0031625
    label: ubiquitin protein ligase binding
  evidence_type: IPI
  original_reference_id: PMID:19122240
  review:
    summary: Ubiquitin protein ligase binding confirmed. SMAD3 interacts with E3 ligases including SMURF2, NEDD4L, STUB1/CHIP
      (PMID:19122240).
    action: ACCEPT
    reason: Well-supported. SMAD3 is a substrate of multiple E3 ubiquitin ligases that regulate its stability and function.
- term:
    id: GO:0006955
    label: immune response
  evidence_type: IMP
  original_reference_id: PMID:16886151
  review:
    summary: SMAD3 participates in immune regulation as a downstream effector of TGF-beta signaling. TGF-beta is a key immunomodulatory
      cytokine, and SMAD3-dependent signaling regulates T cell differentiation, tolerance, and inflammatory responses (PMID:16886151).
    action: KEEP_AS_NON_CORE
    reason: Valid but represents a downstream pleiotropic effect of TGF-beta/SMAD3 signaling rather than a core molecular
      function. TGF-beta/SMAD3 signaling is critical for immune homeostasis, particularly T regulatory cell differentiation
      and suppression of inflammatory responses, but this is a pathway-level outcome.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:11274402
  review:
    summary: Protein binding annotation from high-throughput or targeted interaction study (PMID:11274402). SMAD3 interacts
      with numerous proteins as a signaling hub.
    action: REMOVE
    reason: '''Protein binding'' (GO:0005515) is uninformative for a transcription factor and signaling hub like SMAD3. More
      specific binding terms (co-SMAD binding, R-SMAD binding, TF binding, ubiquitin ligase binding) are preferred and already
      present in the annotation set.'
- term:
    id: GO:0000122
    label: negative regulation of transcription by RNA polymerase II
  evidence_type: IMP
  original_reference_id: PMID:14555988
  review:
    summary: SMAD3 represses transcription of specific target genes in the TGF-beta pathway. Differential regulation through
      SMAD2, SMAD3 and SMAD4 has been demonstrated, with SMAD3 contributing to transcriptional repression at specific promoters
      (PMID:14555988).
    action: ACCEPT
    reason: Core transcriptional function. SMAD3 acts as both a transcriptional activator and repressor depending on the target
      gene and cofactor context. At repressive loci, SMAD3 recruits co-repressors such as Ski/SnoN/TGIF to silence gene expression.
      This is consistent with other annotations for this GO term already accepted with IDA evidence.
- term:
    id: GO:0000122
    label: negative regulation of transcription by RNA polymerase II
  evidence_type: IDA
  original_reference_id: PMID:8774881
  review:
    summary: Negative regulation of transcription by Pol II confirmed (PMID:8774881). SMAD3 can repress transcription.
    action: ACCEPT
    reason: Valid. SMAD3 acts as transcriptional repressor at certain loci with Ski/SnoN/TGIF cofactors.
- term:
    id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  evidence_type: IDA
  original_reference_id: PMID:8774881
  review:
    summary: Positive regulation of Pol II transcription confirmed (PMID:8774881).
    action: ACCEPT
    reason: Core biological process. SMAD3 activates transcription of TGF-beta responsive genes.
- term:
    id: GO:0001666
    label: response to hypoxia
  evidence_type: IMP
  original_reference_id: PMID:12411310
  review:
    summary: Cellular response to hypoxia involves signaling via Smad proteins including SMAD3 (PMID:12411310). Hypoxia activates
      TGF-beta/SMAD signaling, with SMAD3 acting as a mediator of hypoxia-induced gene expression changes.
    action: KEEP_AS_NON_CORE
    reason: Valid annotation supported by IMP evidence, but represents a stimulus-response context for SMAD3 signaling rather
      than a core function. SMAD3 mediates responses to hypoxia through its role in TGF-beta signaling, but hypoxia response
      is not a primary function of SMAD3.
- term:
    id: GO:0032909
    label: regulation of transforming growth factor beta2 production
  evidence_type: IMP
  original_reference_id: PMID:12411310
  review:
    summary: SMAD3 participates in regulation of TGF-beta2 production (PMID:12411310). This represents a feedback mechanism
      within the TGF-beta signaling pathway, where SMAD3-dependent transcription influences production of TGF-beta2 ligand.
    action: KEEP_AS_NON_CORE
    reason: Valid annotation representing a downstream transcriptional outcome of SMAD3 signaling. Regulation of TGF-beta2
      production is a specific transcriptional target/feedback mechanism rather than a core function. SMAD3 regulates many
      target genes; this is one pathway-specific output.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:12446380
  review:
    summary: Nucleus localization confirmed (PMID:12446380). SMAD3 translocates to nucleus upon TGF-beta activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 accumulates in nucleus after TGF-beta-induced phosphorylation.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IDA
  original_reference_id: PMID:12446380
  review:
    summary: Cytoplasm localization confirmed (PMID:12446380). SMAD3 resides in cytoplasm before activation.
    action: ACCEPT
    reason: Core cellular localization. SMAD3 shuttles between cytoplasm and nucleus.
- term:
    id: GO:0030308
    label: negative regulation of cell growth
  evidence_type: IDA
  original_reference_id: PMID:8774881
  review:
    summary: SMAD3 mediates TGF-beta-induced growth inhibition. Zhang et al. (1996) demonstrated that hMAD-3 (SMAD3) and hMAD-4
      (SMAD4) synergize to induce TGF-beta-like responses including growth arrest, and that C-terminally truncated forms act
      as dominant-negative inhibitors of normal TGF-beta response (PMID:8774881).
    action: KEEP_AS_NON_CORE
    reason: Well-supported by early functional studies. TGF-beta-induced growth inhibition is a major biological outcome of
      SMAD3 signaling, but is a downstream cellular response rather than a core molecular or signaling function. SMAD3 mediates
      this through transcriptional regulation of cell cycle inhibitors like p15INK4b and p21.
    supported_by:
    - reference_id: PMID:8774881
      supporting_text: hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses. When truncated
        at their carboxy termini, hMAD-3 and -4 act as dominant-negative inhibitors of the normal TGF-beta response.
- term:
    id: GO:0042803
    label: protein homodimerization activity
  evidence_type: IPI
  original_reference_id: PMID:8774881
  review:
    summary: SMAD3 forms homomeric complexes. Zhang et al. (1996) showed that hMAD-3 (SMAD3) associates with itself, and Nakao
      et al. (1997) demonstrated TGF-beta receptor activation-dependent interaction between SMAD2 and SMAD3, as well as SMAD3
      self-association (PMID:8774881, PMID:9311995).
    action: ACCEPT
    reason: Valid molecular function. SMAD3 forms homotrimers via its MH2 domain, a key step in the signaling pathway. Crystal
      structures confirm the MH2 domain mediates trimerization. While SMAD3 primarily signals as a heterotrimer with SMAD4,
      homodimerization/trimerization is a well-documented intrinsic activity.
    supported_by:
    - reference_id: PMID:8774881
      supporting_text: hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses.
- term:
    id: GO:0043235
    label: receptor complex
  evidence_type: IMP
  original_reference_id: PMID:8774881
  review:
    summary: SMAD3 transiently associates with the TGF-beta receptor complex. Zhang et al. (1996) showed that hMAD-3 (SMAD3)
      was phosphorylated and associated with the ligand-bound receptor complex (PMID:8774881). Nakao et al. (1997) confirmed
      that SMAD3 interacts with the activated TGF-beta type I receptor (PMID:9311995).
    action: KEEP_AS_NON_CORE
    reason: Valid but SMAD3 is not a permanent constituent of the receptor complex. SMAD3 transiently associates with the
      activated TGF-beta type I receptor for phosphorylation and then dissociates. The primary localization of SMAD3 is cytoplasm
      (basal) and nucleus (activated). This CC annotation is technically correct but could be misleading about stable residency.
    supported_by:
    - reference_id: PMID:8774881
      supporting_text: hMAD-3 but not hMAD-4 was phosphorylated and associated with the ligand-bound receptor complex.
- term:
    id: GO:0043565
    label: sequence-specific DNA binding
  evidence_type: IDA
  original_reference_id: PMID:10823886
  review:
    summary: 'SMAD3 directly binds DNA in a sequence-specific manner. The MH1 domain of SMAD3 recognizes the SMAD binding
      element (SBE: GTCT/AGAC) and CAGA-box variants. Structural studies confirm zinc-dependent DNA binding by the MH1 domain
      (PMID:10823886, PMID:12686552).'
    action: ACCEPT
    reason: Core molecular function. Sequence-specific DNA binding through the MH1 domain is a fundamental activity of SMAD3.
      This is well-established by crystal structures, EMSA, and ChIP experiments. Consistent with the IEA annotation for this
      term already accepted.
references:
- id: GO_REF:0000024
  title: Manual transfer of experimentally-verified manual GO annotation data to orthologs by curator judgment of sequence
    similarity
  findings: []
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
  findings: []
- id: GO_REF:0000043
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot keyword mapping
  findings: []
- id: GO_REF:0000052
  title: Gene Ontology annotation based on curation of immunofluorescence data
  findings: []
- id: GO_REF:0000107
  title: Automatic transfer of experimentally verified manual GO annotation data to orthologs using Ensembl Compara
  findings: []
- id: GO_REF:0000113
  title: Gene Ontology annotation of human sequence-specific DNA binding transcription factors (DbTFs) based on the TFClass
    database
  findings: []
- id: GO_REF:0000117
  title: Electronic Gene Ontology annotations created by ARBA machine learning models
  findings: []
- id: GO_REF:0000120
  title: Combined Automated Annotation using Multiple IEA Methods
  findings: []
- id: PMID:10681527
  title: Identification and characterization of a PDZ protein that interacts with activin type II receptors.
  findings: []
- id: PMID:10823886
  title: Transforming growth factor beta -inducible independent binding of SMAD to the Smad7 promoter.
  findings: []
- id: PMID:11274402
  title: Inactivation of menin, a Smad3-interacting protein, blocks transforming growth factor type beta signaling.
  findings: []
- id: PMID:11278251
  title: Smurf1 interacts with transforming growth factor-beta type I receptor through Smad7 and induces receptor degradation.
  findings: []
- id: PMID:11278756
  title: Ski-interacting protein interacts with Smad proteins to augment transforming growth factor-beta-dependent transcription.
  findings: []
- id: PMID:11387212
  title: The adaptor molecule Disabled-2 links the transforming growth factor beta receptors to the Smad pathway.
  findings: []
- id: PMID:12154125
  title: Smad3 allostery links TGF-beta receptor kinase activation to transcriptional control.
  findings: []
- id: PMID:12411310
  title: Cellular response to hypoxia involves signaling via Smad proteins.
  findings: []
- id: PMID:12446380
  title: TGFbeta-Smad signalling in postoperative human lens epithelial cells.
  findings: []
- id: PMID:12650946
  title: Identification of three novel Smad binding proteins involved in cell polarity.
  findings: []
- id: PMID:12686552
  title: Features of a Smad3 MH1-DNA complex. Roles of water and zinc in DNA binding.
  findings: []
- id: PMID:12857746
  title: Direct interaction of Ski with either Smad3 or Smad4 is necessary and sufficient for Ski-mediated repression of transforming
    growth factor-beta signaling.
  findings: []
- id: PMID:12874272
  title: Requirement of the co-repressor homeodomain-interacting protein kinase 2 for ski-mediated inhibition of bone morphogenetic
    protein-induced transcriptional activation.
  findings: []
- id: PMID:12902338
  title: Identification of glucocorticoid receptor domains involved in transrepression of transforming growth factor-beta
    action.
  findings: []
- id: PMID:14525983
  title: DACH1 inhibits transforming growth factor-beta signaling through binding Smad4.
  findings: []
- id: PMID:14555988
  title: Differential regulation of TGF-beta signaling through Smad2, Smad3 and Smad4.
  findings: []
- id: PMID:14612439
  title: A novel E1A-like inhibitor of differentiation (EID) family member, EID-2, suppresses transforming growth factor (TGF)-beta
    signaling by blocking TGF-beta-induced formation of Smad3-Smad4 complexes.
  findings: []
- id: PMID:15051726
  title: Atrophin-1-interacting protein 4/human Itch is a ubiquitin E3 ligase for human enhancer of filamentation 1 in transforming
    growth factor-beta signaling pathways.
  findings: []
- id: PMID:15084259
  title: Integration of Smad and forkhead pathways in the control of neuroepithelial and glioblastoma cell proliferation.
  findings: []
- id: PMID:15107418
  title: Activin receptor-like kinase-7 induces apoptosis through activation of MAPKs in a Smad3-dependent mechanism in hepatoma
    cells.
  findings: []
- id: PMID:15150278
  title: Nodal and ALK7 inhibit proliferation and induce apoptosis in human trophoblast cells.
  findings: []
- id: PMID:15231748
  title: Functional proteomics mapping of a human signaling pathway.
  findings: []
- id: PMID:15334054
  title: Hepatitis C viral proteins interact with Smad3 and differentially regulate TGF-beta/Smad3-mediated transcriptional
    activation.
  findings: []
- id: PMID:15350224
  title: Structural basis of heteromeric smad protein assembly in TGF-beta signaling.
  findings: []
- id: PMID:15527767
  title: 'Proteomics-based identification of proteins interacting with Smad3: SREBP-2 forms a complex with Smad3 and inhibits
    its transcriptional activity.'
  findings: []
- id: PMID:15588252
  title: The Smad3 linker region contains a transcriptional activation domain.
  findings: []
- id: PMID:15647271
  title: The integral inner nuclear membrane protein MAN1 physically interacts with the R-Smad proteins to repress signaling
    by the transforming growth factor-{beta} superfamily of cytokines.
  findings: []
- id: PMID:15781469
  title: CHIP controls the sensitivity of transforming growth factor-beta signaling by modulating the basal level of Smad3
    through ubiquitin-mediated degradation.
  findings: []
- id: PMID:15799969
  title: Nuclear targeting of transforming growth factor-beta-activated Smad complexes.
  findings: []
- id: PMID:15897867
  title: Oligomerization of Evi-1 regulated by the PR domain contributes to recruitment of corepressor CtBP.
  findings: []
- id: PMID:16007207
  title: Hepatitis C virus core variants isolated from liver tumor but not from adjacent non-tumor tissue interact with Smad3
    and inhibit the TGF-beta pathway.
  findings: []
- id: PMID:16200078
  title: Cloning and functional characterization of a new Ski homolog, Fussel-18, specifically expressed in neuronal tissues.
  findings: []
- id: PMID:16751101
  title: PPM1A functions as a Smad phosphatase to terminate TGFbeta signaling.
  findings: []
- id: PMID:16777850
  title: The novel PIAS-like protein hZimp10 enhances Smad transcriptional activity.
  findings: []
- id: PMID:16886151
  title: Association of polymorphisms of IGF1R and genes in the transforming growth factor- beta /bone morphogenetic protein
    pathway with bacteremia in sickle cell anemia.
  findings: []
- id: PMID:17099224
  title: Potentiation of Smad-mediated transcriptional activation by the RNA-binding protein RBPMS.
  findings: []
- id: PMID:17251190
  title: The MH1 domain of Smad3 interacts with Pax6 and represses autoregulation of the Pax6 P1 promoter.
  findings: []
- id: PMID:17292623
  title: Fussel-15, a novel Ski/Sno homolog protein, antagonizes BMP signaling.
  findings: []
- id: PMID:17469184
  title: Involvement of the constitutive complex formation of c-Ski/SnoN with Smads in the impaired negative feedback regulation
    of transforming growth factor beta signaling in scleroderma fibroblasts.
  findings: []
- id: PMID:17785517
  title: Msk is required for nuclear import of TGF-{beta}/BMP-activated Smads.
  findings: []
- id: PMID:18505915
  title: Transforming growth factor-beta1, transforming growth factor-beta2, and transforming growth factor-beta3 enhance
    ovarian cancer metastatic potential by inducing a Smad3-dependent epithelial-to-mesenchymal transition.
  findings: []
- id: PMID:18548003
  title: SMAD proteins control DROSHA-mediated microRNA maturation.
  findings: []
- id: PMID:18568018
  title: TAZ controls Smad nucleocytoplasmic shuttling and regulates human embryonic stem-cell self-renewal.
  findings: []
- id: PMID:18729074
  title: Identification of novel Smad2 and Smad3 associated proteins in response to TGF-beta1.
  findings: []
- id: PMID:18794808
  title: Ligand-dependent ubiquitination of Smad3 is regulated by casein kinase 1 gamma 2, an inhibitor of TGF-beta signaling.
  findings: []
- id: PMID:18832382
  title: HMGA2 and Smads co-regulate SNAIL1 expression during induction of epithelial-to-mesenchymal transition.
  findings: []
- id: PMID:19018011
  title: 'Holding their own: the noncanonical roles of Smad proteins.'
  findings: []
- id: PMID:19032343
  title: Ski co-repressor complexes maintain the basal repressed state of the TGF-beta target gene, SMAD7, via HDAC3 and PRMT5.
  findings: []
- id: PMID:19049980
  title: SKI and MEL1 cooperate to inhibit transforming growth factor-beta signal in gastric cancer cells.
  findings: []
- id: PMID:19122240
  title: Pin1 down-regulates transforming growth factor-beta (TGF-beta) signaling by inducing degradation of Smad proteins.
  findings: []
- id: PMID:19135894
  title: FAM/USP9x, a deubiquitinating enzyme essential for TGFbeta signaling, controls Smad4 monoubiquitination.
  findings: []
- id: PMID:19289081
  title: Nuclear export of Smad2 and Smad3 by RanBP3 facilitates termination of TGF-beta signaling.
  findings: []
- id: PMID:19816956
  title: MiR-21 regulates adipogenic differentiation through the modulation of TGF-beta signaling in mesenchymal stem cells
    derived from human adipose tissue.
  findings: []
- id: PMID:20061380
  title: MTMR4 attenuates transforming growth factor beta (TGFbeta) signaling by dephosphorylating R-Smads in endosomes.
  findings: []
- id: PMID:20129061
  title: TMEPAI, a transmembrane TGF-beta-inducible protein, sequesters Smad proteins from active participation in TGF-beta
    signaling.
  findings: []
- id: PMID:20211142
  title: An atlas of combinatorial transcriptional regulation in mouse and man.
  findings: []
- id: PMID:20935647
  title: IL-37 is a fundamental inhibitor of innate immunity.
  findings: []
- id: PMID:21145499
  title: The Crumbs complex couples cell density sensing to Hippo-dependent control of the TGF-Ξ²-SMAD pathway.
  findings: []
- id: PMID:21258410
  title: Selective targeting of activating and inhibitory Smads by distinct WWP2 ubiquitin ligase isoforms differentially
    modulates TGFΞ² signalling and EMT.
  findings: []
- id: PMID:21297662
  title: Homeodomain protein DLX4 counteracts key transcriptional control mechanisms of the TGF-Ξ² cytostatic program and blocks
    the antiproliferative effect of TGF-Ξ².
  findings: []
- id: PMID:21307346
  title: Response gene to complement 32 interacts with Smad3 to promote epithelial-mesenchymal transition of human renal tubular
    cells.
  findings: []
- id: PMID:21532621
  title: Nodal enhances the activity of FoxO3a and its synergistic interaction with Smads to regulate cyclin G2 transcription
    in ovarian cancer cells.
  findings: []
- id: PMID:21597466
  title: Antagonistic regulation of EMT by TIF1Ξ³ and Smad4 in mammary epithelial cells.
  findings: []
- id: PMID:21828274
  title: HEB and E2A function as SMAD/FOXH1 cofactors.
  findings: []
- id: PMID:21947082
  title: USP15 is a deubiquitylating enzyme for receptor-activated SMADs.
  findings: []
- id: PMID:21988832
  title: Toward an understanding of the protein interaction network of the human liver.
  findings: []
- id: PMID:22045334
  title: Ablation of Smurf2 reveals an inhibition in TGF-Ξ² signalling through multiple mono-ubiquitination of Smad3.
  findings: []
- id: PMID:22155034
  title: Bone morphogenetic protein-3b (BMP-3b) inhibits osteoblast differentiation via Smad2/3 pathway by counteracting Smad1/5/8
    signaling.
  findings: []
- id: PMID:22442258
  title: Intercellular variation in signaling through the TGF-Ξ² pathway and its relation to cell density and cell cycle phase.
  findings: []
- id: PMID:22538441
  title: CSMD1 exhibits antitumor activity in A375 melanoma cells through activation of the Smad pathway.
  findings: []
- id: PMID:23723426
  title: Kindlin-2 mediates activation of TGF-Ξ²/Smad signaling and renal fibrosis.
  findings: []
- id: PMID:24378993
  title: Crosstalk between TGF-Ξ²/Smad3 and BMP/BMPR2 signaling pathways via miR-17-92 cluster in carotid artery restenosis.
  findings: []
- id: PMID:24613385
  title: Hsp70 and Hsp90 oppositely regulate TGF-Ξ² signaling through CHIP/Stub1.
  findings: []
- id: PMID:24627487
  title: C18 ORF1, a novel negative regulator of transforming growth factor-Ξ² signaling.
  findings: []
- id: PMID:24964035
  title: Pin1 promotes degradation of Smad proteins and their interaction with phosphorylated tau in Alzheimer's disease.
  findings: []
- id: PMID:25060702
  title: A Smad3 and TTF-1/NKX2-1 complex regulates Smad4-independent gene expression.
  findings: []
- id: PMID:25416956
  title: A proteome-scale map of the human interactome network.
  findings: []
- id: PMID:25502805
  title: A massively parallel pipeline to clone DNA variants and examine molecular phenotypes of human disease mutations.
  findings: []
- id: PMID:25556234
  title: New host factors important for respiratory syncytial virus (RSV) replication revealed by a novel microfluidics screen
    for interactors of matrix (M) protein.
  findings: []
- id: PMID:25605017
  title: Reduction of miR-29c enhances pancreatic cancer cell migration and stem cell-like phenotype.
  findings: []
- id: PMID:25609649
  title: Proteomic analyses reveal distinct chromatin-associated and soluble transcription factor complexes.
  findings: []
- id: PMID:25670079
  title: 14-3-3ΞΆ turns TGF-Ξ²'s function from tumor suppressor to metastasis promoter in breast cancer by contextual changes
    of Smad partners from p53 to Gli2.
  findings: []
- id: PMID:25893292
  title: Syntenin regulates TGF-Ξ²1-induced Smad activation and the epithelial-to-mesenchymal transition by inhibiting caveolin-mediated
    TGF-Ξ² type I receptor internalization.
  findings: []
- id: PMID:25910212
  title: Widespread macromolecular interaction perturbations in human genetic disorders.
  findings: []
- id: PMID:26311719
  title: MicroRNA-143 Activation Regulates Smooth Muscle and Endothelial Cell Crosstalk in Pulmonary Arterial Hypertension.
  findings: []
- id: PMID:26680585
  title: ShcA Protects against Epithelial-Mesenchymal Transition through Compartmentalized Inhibition of TGF-Ξ²-Induced Smad
    Activation.
  findings: []
- id: PMID:27038547
  title: MiRNA-199a-5p influences pulmonary artery hypertension via downregulating Smad3.
  findings: []
- id: PMID:27060871
  title: Suppression of innate inflammation and immunity by interleukin-37.
  findings: []
- id: PMID:27107012
  title: Pooled-matrix protein interaction screens using Barcode Fusion Genetics.
  findings: []
- id: PMID:28467929
  title: PPARΞ³ Links BMP2 and TGFΞ²1 Pathways in Vascular Smooth Muscle Cells, Regulating Cell Proliferation and Glucose Metabolism.
  findings: []
- id: PMID:28471448
  title: MiR-1 suppresses tumor cell proliferation in colorectal cancer by inhibition of Smad3-mediated tumor glycolysis.
  findings: []
- id: PMID:29789615
  title: IL-37 isoform D downregulates pro-inflammatory cytokines expression in a Smad3-dependent manner.
  findings: []
- id: PMID:29892012
  title: An interactome perturbation framework prioritizes damaging missense mutations for developmental disorders.
  findings: []
- id: PMID:29899023
  title: NEDD9 targets COL3A1 to promote endothelial fibrosis and pulmonary arterial hypertension.
  findings: []
- id: PMID:29997244
  title: 'LuTHy: a double-readout bioluminescence-based two-hybrid technology for quantitative mapping of protein-protein
    interactions in mammalian cells.'
  findings: []
- id: PMID:30729664
  title: Novel role of the clustered miR-23b-3p and miR-27b-3p in enhanced expression of fibrosis-associated genes by targeting
    TGFBR3 in atrial fibroblasts.
  findings: []
- id: PMID:31023188
  title: LRP1 Deficiency in Vascular SMC Leads to Pulmonary Arterial Hypertension That Is Reversed by PPARΞ³ Activation.
  findings: []
- id: PMID:31515488
  title: Extensive disruption of protein interactions by genetic variants across the allele frequency spectrum in human populations.
  findings: []
- id: PMID:31582430
  title: Structural basis for distinct roles of SMAD2 and SMAD3 in FOXH1 pioneer-directed TGF-Ξ² signaling.
  findings: []
- id: PMID:32141990
  title: miR-4286/TGF-Ξ²1/Smad3-Negative Feedback Loop Ameliorated Vascular Endothelial Cell Damage by Attenuating Apoptosis
    and Inflammatory Response.
  findings: []
- id: PMID:32296183
  title: A reference map of the human binary protein interactome.
  findings: []
- id: PMID:32814053
  title: Interactome Mapping Provides a Network of Neurodegenerative Disease Proteins and Uncovers Widespread Protein Aggregation
    in Affected Brains.
  findings: []
- id: PMID:33961781
  title: Dual proteome-scale networks reveal cell-specific remodeling of the human interactome.
  findings: []
- id: PMID:34432647
  title: GDF-8 stimulates trophoblast cell invasion by inducing ALK5-SMAD2/3-mediated MMP2 expression.
  findings: []
- id: PMID:35359452
  title: The Interplay Between TGF-Ξ² Signaling and Cell Metabolism.
  findings: []
- id: PMID:35512704
  title: Systematic discovery of mutation-directed neo-protein-protein interactions in cancer.
  findings: []
- id: PMID:39243984
  title: Gain-of-function variants in SMAD4 compromise respiratory epithelial function.
  findings: []
- id: PMID:8774881
  title: Receptor-associated Mad homologues synergize as effectors of the TGF-beta response.
  findings: []
- id: PMID:9111321
  title: Heteromeric and homomeric interactions correlate with signaling activity and functional cooperativity of Smad3 and
    Smad4/DPC4.
  findings: []
- id: PMID:9311995
  title: TGF-beta receptor-mediated signalling through Smad2, Smad3 and Smad4.
  findings: []
- id: PMID:9732876
  title: Smad3 and Smad4 cooperate with c-Jun/c-Fos to mediate TGF-beta-induced transcription.
  findings: []
- id: PMID:9865696
  title: SARA, a FYVE domain protein that recruits Smad2 to the TGFbeta receptor.
  findings: []
- id: PMID:9892009
  title: Roles of pathway-specific and inhibitory Smads in activin receptor signaling.
  findings: []
- id: Reactome:R-HSA-1535903
  title: Phospho R-SMAD(SMAD2/3):CO-SMAD(SMAD4):FOXO3 binds FoxO3a-binding elements
  findings: []
- id: Reactome:R-HSA-1549526
  title: Phosphorylation of SMAD2,3 by Activin:Activin Receptor
  findings: []
- id: Reactome:R-HSA-170835
  title: An anchoring protein, ZFYVE9 (SARA), recruits SMAD2/3
  findings: []
- id: Reactome:R-HSA-170847
  title: Phosphorylated SMAD2 and SMAD3 form a complex with SMAD4
  findings: []
- id: Reactome:R-HSA-170850
  title: Phosphorylated SMAD2/3 dissociates from TGFBR
  findings: []
- id: Reactome:R-HSA-170868
  title: Activated type I receptor phosphorylates SMAD2/3 directly
  findings: []
- id: Reactome:R-HSA-173481
  title: SKI/SKIL binds SMAD complex, suppressing TGF-beta signaling
  findings: []
- id: Reactome:R-HSA-173488
  title: The SMAD2/3:SMAD4 complex transfers to the nucleus
  findings: []
- id: Reactome:R-HSA-173545
  title: Ubiquitin-dependent degradation of the SMAD complex terminates TGF-beta signaling
  findings: []
- id: Reactome:R-HSA-2031355
  title: WWTR1 binds SMAD2/3:SMAD4 heterotrimer
  findings: []
- id: Reactome:R-HSA-209055
  title: PPM1A dephosphorylates nuclear SMAD2/3
  findings: []
- id: Reactome:R-HSA-2106579
  title: WWTR1:SMAD translocates to the nucleus
  findings: []
- id: Reactome:R-HSA-2127257
  title: SMAD2/3:SMAD4 heterotrimer forms a complex with RBL1, E2F4/5 and DP1/2
  findings: []
- id: Reactome:R-HSA-2176475
  title: Phosphorylation of SMAD2 and SMAD3 linker regions by CDK8 or CDK9
  findings: []
- id: Reactome:R-HSA-2176491
  title: NEDD4L binds phosphorylated linker region of SMAD2/3
  findings: []
- id: Reactome:R-HSA-2176502
  title: NEDD4L ubiquitinates SMAD2/3
  findings: []
- id: Reactome:R-HSA-2176503
  title: Degradation of SMAD2/3:SMAD4 complex
  findings: []
- id: Reactome:R-HSA-2179274
  title: SMURF2 binds SMAD3 phosphorylated in the linker region
  findings: []
- id: Reactome:R-HSA-2179276
  title: SMURF2 monoubiquitinates SMAD3
  findings: []
- id: Reactome:R-HSA-2186607
  title: TGIF recruits HDAC1 to SMAD2/3:SMAD4 heterotrimer
  findings: []
- id: Reactome:R-HSA-2186643
  title: MEN1 binds SMAD2/3:SMAD4 heterotrimer
  findings: []
- id: Reactome:R-HSA-2187309
  title: SMAD2/3:SMAD4 heterotrimer binds SP1
  findings: []
- id: Reactome:R-HSA-2187325
  title: PARP1 ADP-ribosylates SMAD3 and SMAD4
  findings: []
- id: Reactome:R-HSA-2187330
  title: PARP1 binds SMAD2/3:SMAD4 heterotrimer
  findings: []
- id: Reactome:R-HSA-2187355
  title: PMEPA1 sequesters phosphorylated SMAD2/3
  findings: []
- id: Reactome:R-HSA-2187358
  title: PMEPA1 sequesters unphosphorylated SMAD2/3
  findings: []
- id: Reactome:R-HSA-2187368
  title: STUB1 (CHIP) ubiquitinates SMAD3
  findings: []
- id: Reactome:R-HSA-2187375
  title: SMAD3 binds STUB1 (CHIP)
  findings: []
- id: Reactome:R-HSA-2187382
  title: Degradation of ubiquitinated SMAD3
  findings: []
- id: Reactome:R-HSA-2187388
  title: PPM1A protein phosphatase binds phosphorylated SMAD2/3
  findings: []
- id: Reactome:R-HSA-2187395
  title: Dephosphorylated SMAD2/3 translocates to the cytosol
  findings: []
- id: Reactome:R-HSA-2187401
  title: MTMR4 dephosphorylates SMAD2/3
  findings: []
- id: Reactome:R-HSA-2187405
  title: MTMR4 binds phosphorylated SMAD2/3
  findings: []
- id: Reactome:R-HSA-3311014
  title: SMAD4 MH2 Domain Mutants do not bind phosphorylated SMAD2 and SMAD3
  findings: []
- id: Reactome:R-HSA-3315483
  title: Phosphorylated SMAD2/3 MH2 Domain Mutants do not bind SMAD4
  findings: []
- id: Reactome:R-HSA-3656523
  title: An anchoring protein ZFYVE9 (SARA) does not recruit SMAD2/3 to TGFB1:TGFBR2:p-TGFBR1 KD Mutants
  findings: []
- id: Reactome:R-HSA-6781764
  title: USP15 deubiquitinates SMAD1,SMAD2,SMAD3, SMAD7:SMURF,KEAP1
  findings: []
- id: Reactome:R-HSA-870449
  title: TRIM33 monoubiquitinates SMAD4
  findings: []
- id: Reactome:R-HSA-870538
  title: TRIM33 (Ectodermin) binds SMAD heterotrimer in the nucleus
  findings: []
- id: Reactome:R-HSA-8878143
  title: RUNX3 binds SMAD3 and SMAD4
  findings: []
- id: Reactome:R-HSA-8878178
  title: The complex of RUNX3, SMAD3 and SMAD4 binds the CDKN1A gene promoter
  findings: []
- id: Reactome:R-HSA-8952226
  title: RUNX3 binds the BCL2L11 (BIM) gene
  findings: []
- id: Reactome:R-HSA-9008692
  title: IL37(?-218) binds SMAD3
  findings: []
- id: Reactome:R-HSA-9008928
  title: IL37(?-218):p-S423,S425-SMAD3 translocates to the nucleus
  findings: []
- id: Reactome:R-HSA-9009910
  title: IL37(?-218) binds p-S423,S425-SMAD3
  findings: []
- id: Reactome:R-HSA-9617996
  title: FOXO1,FOXO3,FOXO4 bind p-2S-SMAD2/3:SMAD4
  findings: []
- id: Reactome:R-HSA-9618004
  title: FOXO:SMAD complex binds CDKN1A gene promoter
  findings: []
- id: Reactome:R-HSA-9618021
  title: FOXG1 binds FOXO:SMAD complexes
  findings: []
- id: Reactome:R-HSA-9727922
  title: SARS-CoV-1 SUMO1-K62-p-S177-N dimer binds to SMAD3
  findings: []
- id: Reactome:R-HSA-9731111
  title: MAPK1 and MAPK3 phosphorylate SMAD2 and SMAD3
  findings: []
- id: Reactome:R-HSA-9733207
  title: p-2S-SMAD3:p-2S-SMAD3:SMAD4 forms a complex with SP1 and EP300 at the COL1A2 gene promoter
  findings: []
- id: Reactome:R-HSA-9733247
  title: YBX1 binds COL1A2 gene promoter
  findings: []
- id: Reactome:R-HSA-9736959
  title: WWTR1:p-2S-SMAD2,3:SMAD4 binds SERPINE1 gene promoter
  findings: []
- id: Reactome:R-HSA-9736970
  title: WWTR1:p-2S-SMAD2,3:SMAD4 binds SMAD7 gene promoter
  findings: []
- id: Reactome:R-HSA-9736979
  title: p-2S-SMAD2,3:SMAD4:SP1 binds CDKN2B gene promoter
  findings: []
- id: Reactome:R-HSA-9736984
  title: p-2S-SMAD2,3:SMAD4:RBL1:E2F4,5:DP1,2 binds MYC gene promoter
  findings: []
- id: Reactome:R-HSA-9736992
  title: p-2S-SMAD2,3:SMAD4 binds JUNB gene promoter
  findings: []
- id: Reactome:R-HSA-9737710
  title: SARS-CoV-1 N binds SMAD3 and EP300 at the SERPINE1 gene promoter
  findings: []
- id: Reactome:R-HSA-9823952
  title: EOMES, SMAD2,3, and GATA6 bind the CXCR4 gene
  findings: []
- id: Reactome:R-HSA-9823959
  title: SMAD2,3 and GATA6-AS1 lncRNA bind the GATA6 gene
  findings: []
- id: Reactome:R-NUL-2186736
  title: Rnf111 binds SKI/SKIL in complex with SMAD2/3:SMAD4 upon TGF-beta stimulation
  findings: []
- id: Reactome:R-NUL-2186755
  title: Ubiquitination of SKI/SKIL by Rnf111
  findings: []
- id: Reactome:R-NUL-9625758
  title: FOXO1,FOXO3 and SMAD3 bind Trim63 gene promoter
  findings: []
core_functions:
- description: 'SMAD3 functions as a sequence-specific DNA-binding transcription factor for RNA polymerase II. The MH1 domain
    directly binds SMAD binding elements (SBE: GTCT/AGAC) and CAGA-box motifs, and cooperates with coactivators (p300/CBP)
    or corepressors (Ski/SnoN/TGIF) to regulate target gene transcription.'
  molecular_function:
    id: GO:0000981
    label: DNA-binding transcription factor activity, RNA polymerase II-specific
  directly_involved_in:
  - id: GO:0045893
    label: positive regulation of DNA-templated transcription
  - id: GO:0000122
    label: negative regulation of transcription by RNA polymerase II
  - id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  locations:
  - id: GO:0005634
    label: nucleus
  supported_by:
  - reference_id: PMID:9732876
    supporting_text: Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription
      from the TRE in the absence of c-Jun and c-Fos.
  - reference_id: PMID:8774881
    supporting_text: hMAD-3 and -4 synergized to induce strong ligand-independent TGF-beta-like responses.
- description: SMAD3 binds the activated TGF-beta type I receptor (ALK5/TGFBR1) as the principal intracellular signal transducer.
    Upon receptor binding and C-terminal SSXS phosphorylation, SMAD3 forms heteromeric complexes with SMAD4 and translocates
    to the nucleus.
  molecular_function:
    id: GO:0070410
    label: co-SMAD binding
  directly_involved_in:
  - id: GO:0007179
    label: transforming growth factor beta receptor signaling pathway
  - id: GO:0060395
    label: SMAD protein signal transduction
  locations:
  - id: GO:0005737
    label: cytoplasm
  - id: GO:0005634
    label: nucleus
  supported_by:
  - reference_id: PMID:9311995
    supporting_text: Smad2 and Smad3 interacted with the kinase-deficient TGF-beta type I receptor (TbetaR)-I after it was
      phosphorylated by TbetaR-II kinase. TGF-beta1 induced phosphorylation of Smad2 and Smad3 in Mv1Lu mink lung epithelial
      cells.
- description: SMAD3 binds to cis-regulatory DNA sequences including SMAD binding elements (SBE) and TPA-responsive elements
    (TRE/AP-1 sites), acting as a direct DNA-binding component of the TGF-beta transcriptional response.
  molecular_function:
    id: GO:0000976
    label: transcription cis-regulatory region binding
  directly_involved_in:
  - id: GO:0045944
    label: positive regulation of transcription by RNA polymerase II
  locations:
  - id: GO:0005634
    label: nucleus
  supported_by:
  - reference_id: PMID:9732876
    supporting_text: Smad3 interacts directly with the TRE and that Smad3 and Smad4 can activate TGF-beta-inducible transcription
      from the TRE in the absence of c-Jun and c-Fos.