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    Smad3 SMAD family member 3 [ Rattus norvegicus (Norway rat) ]

    Gene ID: 25631, updated on 9-Dec-2024

    Summary

    Official Symbol
    Smad3provided by RGD
    Official Full Name
    SMAD family member 3provided by RGD
    Primary source
    RGD:3032
    See related
    EnsemblRapid:ENSRNOG00000008620 AllianceGenome:RGD:3032
    Gene type
    protein coding
    RefSeq status
    VALIDATED
    Organism
    Rattus norvegicus
    Lineage
    Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae; Murinae; Rattus
    Also known as
    mad3; Madh3; Smad 3
    Summary
    Enables several functions, including DNA-binding transcription factor activity; RNA polymerase II transcription regulatory region sequence-specific DNA binding activity; and beta-catenin binding activity. Involved in several processes, including cell surface receptor protein serine/threonine kinase signaling pathway; positive regulation of hydrolase activity; and regulation of gene expression. Located in nucleus. Part of transcription regulator complex. Used to study pre-malignant neoplasm. Human ortholog(s) of this gene implicated in Loeys-Dietz syndrome 3; Lynch syndrome; and pancreatic cancer. Orthologous to human SMAD3 (SMAD family member 3). [provided by Alliance of Genome Resources, Dec 2024]
    Expression
    Biased expression in Muscle (RPKM 301.2), Kidney (RPKM 125.7) and 9 other tissues See more
    Orthologs
    NEW
    Try the new Gene table
    Try the new Transcript table

    Genomic context

    See Smad3 in Genome Data Viewer
    Location:
    8q24
    Exon count:
    12
    Annotation release Status Assembly Chr Location
    RS_2024_02 current GRCr8 (GCF_036323735.1) 8 NC_086026.1 (73022204..73132324, complement)
    RS_2023_06 previous assembly mRatBN7.2 (GCF_015227675.2) 8 NC_051343.1 (64126829..64236960, complement)
    106 previous assembly Rnor_6.0 (GCF_000001895.5) 8 NC_005107.4 (68569530..68678349, complement)

    Chromosome 8 - NC_086026.1Genomic Context describing neighboring genes Neighboring gene IQ motif containing H Neighboring gene uncharacterized LOC134480189 Neighboring gene alpha- and gamma-adaptin binding protein Neighboring gene SMAD family member 6 Neighboring gene lactase-like Neighboring gene zwilch kinetochore protein

    Genomic regions, transcripts, and products

    Bibliography

    GeneRIFs: Gene References Into Functions

    What's a GeneRIF?

    Pathways from PubChem

    Interactions

    Products Interactant Other Gene Complex Source Pubs Description

    General gene information

    Markers

    Gene Ontology Provided by RGD

    Function Evidence Code Pubs
    enables DEAD/H-box RNA helicase binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription activator activity, RNA polymerase II-specific ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription factor activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    contributes_to DNA-binding transcription factor activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription factor activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription factor activity, RNA polymerase II-specific IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables DNA-binding transcription factor activity, RNA polymerase II-specific ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables DNA-binding transcription repressor activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables I-SMAD binding IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables R-SMAD binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables RNA polymerase II cis-regulatory region sequence-specific DNA binding IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables RNA polymerase II cis-regulatory region sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables RNA polymerase II transcription regulatory region sequence-specific DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables RNA polymerase II transcription regulatory region sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables RNA polymerase II-specific DNA-binding transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables bHLH transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables beta-catenin binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables chromatin DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables chromatin binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables cis-regulatory region sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables cis-regulatory region sequence-specific DNA binding ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    enables co-SMAD binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables collagen binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables double-stranded DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables enzyme binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables identical protein binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables metal ion binding IEA
    Inferred from Electronic Annotation
    more info
     
    enables nuclear glucocorticoid receptor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables nuclear mineralocorticoid receptor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables nuclear receptor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables phosphatase binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables promoter-specific chromatin binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables protein binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables protein homodimerization activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables protein kinase binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables sequence-specific DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables sterol response element binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables transcription cis-regulatory region binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables transcription coactivator binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables transcription corepressor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables transforming growth factor beta receptor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables ubiquitin binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables ubiquitin protein ligase binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables zinc ion binding ISO
    Inferred from Sequence Orthology
    more info
     
    Process Evidence Code Pubs
    involved_in JNK cascade ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in MAPK cascade ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in SMAD protein signal transduction IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in SMAD protein signal transduction IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of SMAD protein signal transduction ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within SMAD protein signal transduction ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in SMAD protein signal transduction ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in T cell activation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in activin receptor signaling pathway IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in activin receptor signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in adrenal gland development IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in anatomical structure morphogenesis IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    acts_upstream_of_or_within apoptotic process ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in apoptotic signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cell differentiation IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    acts_upstream_of_or_within cell population proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cell-cell junction organization ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to glucose stimulus IEP
    Inferred from Expression Pattern
    more info
    PubMed 
    involved_in cellular response to transforming growth factor beta stimulus IEP
    Inferred from Expression Pattern
    more info
    PubMed 
    involved_in cellular response to transforming growth factor beta stimulus ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to virus ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within developmental growth ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within embryonic cranial skeleton morphogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within embryonic foregut morphogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within embryonic pattern specification ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within endoderm development ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in extrinsic apoptotic signaling pathway IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in extrinsic apoptotic signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within gastrulation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within heart looping ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in immune response ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within immune system development ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within in utero embryonic development ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within lens fiber cell differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within liver development ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within mesoderm formation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of apoptotic process IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of cardiac muscle hypertrophy in response to stress IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in negative regulation of cell growth ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of cell population proliferation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of cell population proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of cytosolic calcium ion concentration ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of fat cell differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of negative regulation of gene expression ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of inflammatory response ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of lung blood pressure IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in negative regulation of miRNA transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of ossification ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within negative regulation of osteoblast differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of osteoblast proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of protein catabolic process IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of protein catabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in negative regulation of transcription by RNA polymerase II IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within negative regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of wound healing ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in nodal signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within osteoblast development ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in osteoblast differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within paraxial mesoderm morphogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within pericardium development ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within positive regulation of DNA-templated transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of DNA-templated transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of SMAD protein signal transduction ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of alkaline phosphatase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of bone mineralization IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of canonical Wnt signaling pathway IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of cell migration IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of chondrocyte differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of epithelial to mesenchymal transition ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of positive regulation of extracellular matrix assembly ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of extracellular matrix assembly ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of focal adhesion assembly IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of gene expression ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of interleukin-1 beta production IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of positive regulation of miRNA transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of miRNA transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of nitric oxide biosynthetic process IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in positive regulation of nitric oxide biosynthetic process ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of positive chemotaxis IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of stress fiber assembly IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of transcription by RNA polymerase II IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in positive regulation of transcription by RNA polymerase II IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of transcription by RNA polymerase II IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within positive regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of transforming growth factor beta3 production IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in protein stabilization IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of DNA-templated transcription IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in regulation of DNA-templated transcription ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within regulation of epithelial cell proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of immune response ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of mitochondrial membrane potential IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within regulation of striated muscle tissue development ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within regulation of transforming growth factor beta receptor signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of transforming growth factor beta2 production ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in release of cytochrome c from mitochondria IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of response to angiotensin ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in response to hypoxia ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within response to transforming growth factor beta ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in signal transduction involved in regulation of gene expression IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within skeletal system development ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within somitogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within thyroid gland development ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in transdifferentiation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in transforming growth factor beta receptor signaling pathway IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in transforming growth factor beta receptor signaling pathway IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within transforming growth factor beta receptor signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in transforming growth factor beta receptor signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in ureteric bud development ISO
    Inferred from Sequence Orthology
    more info
     
    Component Evidence Code Pubs
    part_of SMAD protein complex ISO
    Inferred from Sequence Orthology
    more info
     
    part_of SMAD protein complex ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in chromatin ISO
    Inferred from Sequence Orthology
    more info
     
    located_in cytoplasm IEA
    Inferred from Electronic Annotation
    more info
     
    located_in cytoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    located_in cytosol ISO
    Inferred from Sequence Orthology
    more info
     
    part_of heteromeric SMAD protein complex IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    part_of heteromeric SMAD protein complex ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nuclear inner membrane ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleoplasm IEA
    Inferred from Electronic Annotation
    more info
     
    located_in nucleoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleus IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in nucleus IEA
    Inferred from Electronic Annotation
    more info
     
    located_in nucleus ISO
    Inferred from Sequence Orthology
    more info
     
    located_in plasma membrane ISO
    Inferred from Sequence Orthology
    more info
     
    part_of receptor complex ISO
    Inferred from Sequence Orthology
    more info
     
    part_of transcription regulator complex IDA
    Inferred from Direct Assay
    more info
    PubMed 
    part_of transcription regulator complex IEA
    Inferred from Electronic Annotation
    more info
     
    part_of transcription regulator complex ISO
    Inferred from Sequence Orthology
    more info
     

    General protein information

    Preferred Names
    mothers against decapentaplegic homolog 3
    Names
    MAD (mothers against decapentaplegic, Drosophila) homolog 3
    MAD homolog 3
    mothers against DPP homolog 3

    NCBI Reference Sequences (RefSeq)

    NEW Try the new Transcript table

    RefSeqs maintained independently of Annotated Genomes

    These reference sequences exist independently of genome builds. Explain

    These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

    mRNA and Protein(s)

    1. NM_013095.3NP_037227.1  mothers against decapentaplegic homolog 3

      See identical proteins and their annotated locations for NP_037227.1

      Status: VALIDATED

      Source sequence(s)
      BC064437, JAXUCZ010000008
      UniProtKB/Swiss-Prot
      P84025
      UniProtKB/TrEMBL
      A0A8I5ZS20, A6J5A4
      Related
      ENSRNOP00000037346.3, ENSRNOT00000039730.6
      Conserved Domains (2) summary
      cd10491
      Location:8132
      MH1_SMAD_2_3; N-terminal Mad Homology 1 (MH1) domain in SMAD2 and SMAD3
      cd10985
      Location:224414
      MH2_SMAD_2_3; C-terminal Mad Homology 2 (MH2) domain in SMAD2 and SMAD3

    RefSeqs of Annotated Genomes: GCF_036323735.1-RS_2024_02

    The following sections contain reference sequences that belong to a specific genome build. Explain

    Reference GRCr8

    Genomic

    1. NC_086026.1 Reference GRCr8

      Range
      73022204..73132324 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_039080897.2XP_038936825.1  mothers against decapentaplegic homolog 3 isoform X2

      UniProtKB/TrEMBL
      A0A8I5ZS20
      Conserved Domains (2) summary
      cd10985
      Location:119309
      MH2_SMAD_2_3; C-terminal Mad Homology 2 (MH2) domain in SMAD2 and SMAD3
      cl00055
      Location:127
      MH1; N-terminal Mad Homology 1 (MH1) domain
    2. XM_039080896.2XP_038936824.1  mothers against decapentaplegic homolog 3 isoform X1

      UniProtKB/TrEMBL
      A0A8I5ZS20
      Related
      ENSRNOP00000081624.2, ENSRNOT00000099327.2
      Conserved Domains (2) summary
      cd10985
      Location:202392
      MH2_SMAD_2_3; C-terminal Mad Homology 2 (MH2) domain in SMAD2 and SMAD3
      cl00055
      Location:48110
      MH1; N-terminal Mad Homology 1 (MH1) domain
    3. XM_008766216.4XP_008764438.1  mothers against decapentaplegic homolog 3 isoform X3

      See identical proteins and their annotated locations for XP_008764438.1

      Conserved Domains (1) summary
      cd10985
      Location:29219
      MH2_SMAD_2_3; C-terminal Mad Homology 2 (MH2) domain in SMAD2 and SMAD3