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Category Symbol Source: HGNC EntrezGene Ensembl GeneCards RNA genes CroW21

GIFtS
-

SMAD4 Gene

protein-coding GIFtS: 71
GCID: GC18P048494

SMAD family member 4

(Previous names: MAD, mothers against decapentaplegic homolog 4 (Drosophila),...)
(Previous symbol: MADH4)

Explore 114 diseases affiliated with
SMAD4 via

MalaCards

our new
Human Malady Compendium

(According to ¹HGNC, ²Entrez Gene,
³UniProtKB/Swiss-Prot, ⁴UniProtKB/TrEMBL, ⁵OMIM, ⁶GeneLoc, ⁷Ensembl, ⁸DME, ⁹miRBase, and/or ¹⁰fRNAdb)
About This Section

Aliases
SMAD Family Member 4¹ ² ³		Deleted In Pancreatic Carcinoma Locus 4²
DPC4¹ ² ³ ⁵		Mothers Against Decapentaplegic Homolog 4²
MADH4¹ ² ³ ⁵		Mothers Against Decapentaplegic, Drosophila, Homolog Of, 4²
Deletion Target In Pancreatic Carcinoma 4² ³		SMAD, Mothers Against DPP Homolog 4²
MAD Homolog 4² ³		Smad4³
JIP² ⁵		HSMAD4¹
MAD, Mothers Against Decapentaplegic Homolog 4 (Drosophila)¹		Mothers Against DPP Homolog 4³
SMAD, Mothers Against DPP Homolog 4 (Drosophila)¹		SMAD 4³
MYHRS²

External Ids:	HGNC: 6770¹	Entrez Gene: 4089²	Ensembl: ENSG00000141646⁷	OMIM: 600993⁵	UniProtKB: Q13485³

Export aliases for SMAD4 gene to outside databases

Previous GC identifers: GC18P046809 GC18P046810 GC18P048556 GC18P045411

(According to Entrez Gene, Tocris Bioscience, Wikipedia's Gene Wiki, PharmGKB,
UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL)
About This Section

Entrez Gene summary for SMAD4:

This gene encodes a member of the Smad family of signal transduction proteins. Smad proteins are phosphorylated and

activated by transmembrane serine-threonine receptor kinases in response to TGF-beta signaling. The product of this

gene forms homomeric complexes and heteromeric complexes with other activated Smad proteins, which then accumulate in

the nucleus and regulate the transcription of target genes. This protein binds to DNA and recognizes an 8-bp

palindromic sequence (GTCTAGAC) called the Smad-binding element (SBE). The Smad proteins are subject to complex

regulation by post-translational modifications. Mutations or deletions in this gene have been shown to result in

pancreatic cancer, juvenile polyposis syndrome, and hereditary hemorrhagic telangiectasia syndrome. (provided by

RefSeq, Oct 2009)

UniProtKB/Swiss-Prot: SMAD4_HUMAN, Q13485

Function: Common SMAD (co-SMAD) is the coactivator and mediator of signal transduction by TGF-beta (transforming growth

factor). Component of the heterotrimeric SMAD2/SMAD3-SMAD4 complex that forms in the nucleus and is required for the

TGF-mediated signaling. Promotes binding of the SMAD2/SMAD4/FAST-1 complex to DNA and provides an activation function

required for SMAD1 or SMAD2 to stimulate transcription. Component of the multimeric SMAD3/SMAD4/JUN/FOS complex which

forms at the AP1 promoter site; required for syngernistic transcriptional activity in response to TGF-beta. May act as

a tumor suppressor. Positively regulates PDPK1 kinase activity by stimulating its dissociation from the 14-3-3 protein

YWHAQ which acts as a negative regulator

Gene Wiki entry for SMAD4 (Mothers against decapentaplegic homolog 4)

(According to GeneLoc and/or HGNC, and/or
Entrez Gene (NCBI build 37),
and/or miRBase,
Genomic Views according to UCSC (hg19) and Ensembl (release 69), Regulatory elements and Epigenetics data according to QIAGEN, SABiosciences, and/or SwitchGear Genomics)
About This Section

RefSeq DNA sequence:

NC_000018.9 NC_018929.1 NT_010966.14

Regulatory elements:

SABiosciences Regulatory transcription factor binding sites in the SMAD4 gene promoter:
AP-1 ATF-2 Nkx2-5 c-Jun
Other transcription factors

SwitchGear Promoter luciferase reporter plasmid: SMAD4 promoter sequence

Search SABiosciences Chromatin IP Primers for SMAD4

Epigenetics:

QIAGEN PyroMark CpG Assay predesigned Pyrosequencing DNA Methylation assays in human, mouse, rat SMAD4

Genomic Location:
Genomic View: UCSC Golden Path with GeneCards custom track

Entrez Gene cytogenetic band: 18q21.1 Ensembl cytogenetic band: 18q21.2 HGNC cytogenetic band: 18q21.1

SMAD4 Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
SMAD4 gene location

GeneLoc information about chromosome 18 GeneLoc Exon Structure

GeneLoc location for GC18P048494: view genomic region (about GC identifiers)

Start:	48,494,410 bp from pter	End:	48,611,415 bp from pter
Size:	117,006 bases	Orientation:	plus strand

(According to ¹UniProtKB, HORDE, neXtProt, Ensembl, and/or Reactome, Modification sites according to ²PhosphoSitePlus, Specific Peptides from DME, Protein expression images according to data from SPIRE MOPED and PaxDb, RefSeq according to NCBI, PDB rendering according to OCA and/or Proteopedia, Recombinant Proteins from EMD Millipore, R&D Systems, GenScript, Enzo Life Sciences, OriGene, Novus Biologicals, Sino Biological, ProSpec, and/or Uscn,
Biochemical Assays by EMD Millipore, R&D Systems, OriGene, GenScript, Cell Signaling Technology, Enzo Life Sciences, and/or Uscn, Ontologies according to Gene Ontology Consortium 01 Mar 2013 and Entrez Gene, Antibodies by EMD Millipore, R&D Systems, GenScript, Cell Signaling Technology, OriGene, Novus Biologicals, Thermo Fisher Scientific, Abcam, and/or Uscn)
About This Section

UniProtKB/Swiss-Prot: SMAD4_HUMAN, Q13485 (See protein sequence)

Recommended Name: Mothers against decapentaplegic homolog 4

Size: 552 amino acids; 60439 Da

Subunit: Interacts with CITED2 (By similarity). Monomer; in the absence of TGF-beta activation. Heterodimer; on

TGF-beta activation. Composed of two molecules of a C-terminally phosphorylated R-SMAD molecule, SMAD2 or SMAD3, and

one molecule of SMAD4 to form the transcriptional active SMAD2/SMAD3-SMAD4 complex. Found in a ternary complex

composed of SMAD4, STK11/LKB1 and STK11IP. Interacts with ATF2, COPS5, DACH1, MSG1, SKI, STK11/LKB1, STK11IP and

TRIM33. Interacts with ZNF423; the interaction takes place in response to BMP2 leading to activation of transcription

of BMP target genes. Interacts with ZNF521; the interaction takes place in response to BMP2 leading to activation of

transcription of BMP target genes. Interacts with USP9X. Interacts (via the MH1 and MH2 domains) with RBPMS. Interacts

with WWTR1 (via coiled-coil domain). Component of the multimeric complex SMAD3/SMAD4/JUN/FOS which forms at the AP1

promoter site; required for syngernistic transcriptional activity in response to TGF-beta. Interacts with CITED1.

Interacts with PDPK1 (via PH domain) (By similarity). Interacts with VPS39; this interaction affects heterodimer

formation with SMAD3, but not with SMAD2, and leads to inhibition of SMAD3-dependent transcription activation.

Interactions with VPS39 and SMAD2 may be mutually exclusive

Subcellular location: Cytoplasm. Nucleus. Note=Cytoplasmic in the absence of ligand. Migrates to the nucleus when

complexed with R-SMAD. PDPK1 prevents its nuclear translocation in response to TGF-beta

6 PDB 3D structures from and Proteopedia for SMAD4:

1DD1 (3D)

1G88 (3D)

1MR1 (3D)

1U7F (3D)

1U7V (3D)

1YGS (3D)

Secondary accessions: A8K405

Explore the universe of human proteins at neXtProt for SMAD4: NX_Q13485

Post-translational modifications:

Phosphorylated by PDPK1¹

Monoubiquitinated on Lys-519 by E3 ubiquitin-protein ligase TRIM33. Monoubiquitination hampers its ability to form a

stable complex with activated SMAD2/3 resulting in inhibition of TGF-beta/BMP signaling cascade. Deubiqitination by

USP9X restores its competence to mediate TGF-beta signaling¹

View modification sites using PhosphoSitePlus²

View neXtProt modification sites for NX_Q13485

SMAD4 Protein expression data from MOPED and PaxDb: About this image

REFSEQ proteins: NP_005350.1

ENSEMBL proteins:

ENSP00000465878 ENSP00000381452 ENSP00000466934 ENSP00000464772 ENSP00000466941

ENSP00000465874 ENSP00000464901 ENSP00000468611 ENSP00000466118 ENSP00000341551

ENSP00000409551

Reactome Protein details: Q13485
Human Recombinant Protein Products:

	Browse Purified and Recombinant Proteins at EMD Millipore
	Browse R&D Systems for human recombinant proteins
	Browse recombinant and purified proteins available from Enzo Life Sciences
	OriGene Purified Protein: SMAD4 OriGene Protein Over-expression Lysate: SMAD4 OriGene Custom Protein Services for SMAD4
	GenScript Custom Purified and Recombinant Proteins Services for SMAD4
	Novus Biologicals SMAD4 Proteins Novus Biologicals SMAD4 Lysate
	Browse Sino Biological Recombinant Proteins
	ProSpec Recombinant Protein for SMAD4
	Uscn Proteins for SMAD4

Gene Ontology (GO): 5/10 cellular component terms (GO ID links to tree view) (see all 10): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005634	nucleus	IDA	--
GO:0005654	nucleoplasm	TAS	--
GO:0005667	transcription factor complex	IPI	12374795
GO:0005730	NOT nucleolus	IDA	--
GO:0005737	cytoplasm	IDA	--

SMAD4 for ontologies About GeneDecksing

SMAD4 Antibody Products:

	EMD Millipore Mono- and Polyclonal Antibodies for the study of SMAD4
	R&D Systems Antibodies for SMAD4
	Cell Signaling Technology (CST) Antibodies for SMAD4
	OriGene Antibodies (see all 4): SMAD4 OriGene Custom Antibody Services for SMAD4
	GenScript Custom Superior Antibodies Services for SMAD4
	Novus Biologicals SMAD4 Antibodies
	Abcam antibodies for SMAD4
	Uscn Antibodies for SMAD4
	ThermoFisher Antibody for SMAD4

Assay Products for SMAD4:

	Browse Kits and Assays available from EMD Millipore
	OriGene Custom Immunoassay Development Browse OriGene Fluorogenic Cell Assay Kits
	R&D Systems Transcription Factor Binding & Immunoprecipitation Assays for SMAD4
	GenScript Custom Assay Services for SMAD4
	Browse Enzo Life Sciences for kits & assays
	Uscn ELISAs and CLIAs for SMAD4

(According to InterPro, ProtoNet, UniProtKB, and/or BLOCKS, Sets of similar genes according to GeneDecks)
About This Section

SMAD4 for domains About GeneDecksing

5/6 InterPro domains/families (see all 6):

IPR013019 MAD_homology_MH1

IPR008984 SMAD_FHA_domain

IPR001132 SMAD_dom_Dwarfin-type

IPR017855 SMAD_dom-like

IPR013790 Dwarfin

Graphical View of Domain Structure for InterPro Entry Q13485

ProtoNet protein and cluster: Q13485

1 Blocks protein family: IPB001132 Dwarfin protein

UniProtKB/Swiss-Prot: SMAD4_HUMAN, Q13485

Domain: The MH1 domain is required for DNA binding

Domain: The MH2 domain is required for both homomeric and heteromeric interactions and for transcriptional regulation.

Sufficient for nuclear import

Similarity: Belongs to the dwarfin/SMAD family

Similarity: Contains 1 MH1 (MAD homology 1) domain

Similarity: Contains 1 MH2 (MAD homology 2) domain

(According to ¹UniProtKB, Genatlas, LifeMap Discovery™, IUBMB, and/or ²DME, Human phenotypes from GenomeRNAi, Animal models from MGI Mar 06 2013,
bound targets from SABiosciences, miRNA Gene Targets from miRTarBase shRNA from OriGene, RNAi from EMD Millipore, siRNAs from OriGene, QIAGEN, microRNA from QIAGEN, Gene Editing from DNA2.0, Clones from EMD Millipore, OriGene, SwitchGear Genomics, GenScript, Sino Biological, DNA2.0, and Vector BioLabs, Cell Lines from GenScript, LifeMap BioReagents, In Situ Hybridization Assays from Advanced Cell Diagnostics, Ontologies according to Gene Ontology Consortium 01 Mar 2013 via Entrez Gene.)
About This Section

Function Summary:

UniProtKB/Swiss-Prot: SMAD4_HUMAN, Q13485

Function: Common SMAD (co-SMAD) is the coactivator and mediator of signal transduction by TGF-beta (transforming growth

factor). Component of the heterotrimeric SMAD2/SMAD3-SMAD4 complex that forms in the nucleus and is required for the

TGF-mediated signaling. Promotes binding of the SMAD2/SMAD4/FAST-1 complex to DNA and provides an activation function

required for SMAD1 or SMAD2 to stimulate transcription. Component of the multimeric SMAD3/SMAD4/JUN/FOS complex which

forms at the AP1 promoter site; required for syngernistic transcriptional activity in response to TGF-beta. May act as

a tumor suppressor. Positively regulates PDPK1 kinase activity by stimulating its dissociation from the 14-3-3 protein

YWHAQ which acts as a negative regulator

miRNA Products:		miRTarBase miRNAs that target SMAD4: hsa-mir-17 (MIRT005629), hsa-mir-483-3p (MIRT004118), hsa-mir-20a (MIRT005631), hsa-mir-19a (MIRT005630), hsa-mir-26a (MIRT003803), hsa-mir-92a (MIRT005632), hsa-mir-18a (MIRT004736)

		OriGene 3'-UTR Clone: SMAD4 Browse MicroRNA Expression Plasmids
		QIAGEN Custom miScript Target Protector blocks miRNA-binding site of human, mouse, rat SMAD4
		8/195 QIAGEN miScript miRNA Assays for microRNAs that regulate SMAD4 (see all 195):
		hsa-miR-411* hsa-miR-106a hsa-miR-146a hsa-miR-605 hsa-miR-519a hsa-miR-3921 hsa-miR-298 hsa-miR-130b

		SwitchGear 3'UTR luciferase reporter plasmid: SMAD4 3' UTR sequence

Inhib. RNA Products:		Browse for Gene Knock-down Tools from EMD Millipore
		OriGene 29mer shRNA kits in GFP-retroviral vector in human, mouse, rat for SMAD4 (see all 7) OriGene shRNA RFP: SMAD4 OriGene siRNA: SMAD4
		QIAGEN FlexiTube/FlexiPlate siRNA for gene silencing in human, mouse, rat SMAD4

Gene Editing
Products:

DNA2.0 Custom Protein Engineering Service for SMAD4

Clone Products:		Browse Clones for the Expression of Recombinant Proteins Available from EMD Millipore
		OriGene Myc/DDK tagged cDNA clones in CMV expression vector in human, mouse, rat for SMAD4 (see all 3) OriGene untagged cDNA clones in CMV expression vector in human, mouse, rat for SMAD4 OriGene custom cloning services � gene synthesis, subcloning, mutagenesis, variant library, vector shuttling
		GenScript: all cDNA clones in your preferred vector: SMAD4 (NM_005359)
		Sino Biological Human cDNA Clone for SMAD4
		DNA2.0 Custom Codon Optimized Gene Synthesis Service for SMAD4
		Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat SMAD4

Cell Line Products:		GenScript Custom overexpressing Cell Line Services for SMAD4
		Search LifeMap BioReagents cell lines for SMAD4

In Situ Assay
Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for SMAD4

Gene Ontology (GO): 5/16 molecular function terms (GO ID links to tree view) (see all 16): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0000987	core promoter proximal region sequence-specific DNA binding	IDA	18832382
GO:0000988	contributes to protein binding transcription factor activity	IDA	18832382
GO:0003677	contributes to DNA binding	IDA	9389648
GO:0003682	chromatin binding	IEA	--
GO:0003700	contributes to sequence-specific DNA binding transcription factor activity	IDA	17438144

SMAD4 for ontologies About GeneDecksing

6 GenomeRNAi human phenotypes for SMAD4:

Increased S DNA content, incre	Increased mitotic index	Increased sensitivity to pacli	S arrest
Synthetic lethal with gemcitab	Upregulation of Wnt/beta-caten

Animal Models:
     Mouse knock-outs for SMAD4: Smad4^tm1.1Rob Smad4^tm1.1Rdp Smad4^tm2.1Rob Smad4^tm2.2Cxd Smad4^tm1Cxd Smad4^tm1Mak
                                               Smad4^tm2Cxd Smad4^tm1Mmt
     15/19 MGI mutant phenotypes (inferred from 17 alleles) (MGI details for Smad4) (see all 19):

behavior/neurological	cardiovascular system	cellular	digestive/alimentary	embryogenesis
endocrine/exocrine gland	growth/size	hematopoietic system	homeostasis/metabolism	immune system
integument	liver/biliary system	mortality/aging	muscle	nervous system

SMAD4 for phenotypes About GeneDecksing

(Pathways according to EMD Millipore, R&D Systems, Cell Signaling Technology, KEGG, PharmGKB, BioSystems, Reactome, Tocris Bioscience, GeneGo (Thomson Reuters), QIAGEN, and/or UniProtKB, Sets of similar genes according to GeneDecks, Interaction Networks according to SABiosciences, and/or STRING, Interactions according to ¹UniProtKB, ²MINT, ³I2D, and/or ⁴STRING, with links to IntAct and Ensembl, Ontologies according to Gene Ontology Consortium 01 Mar 2013 via Entrez Gene).
About This Section

Unified GeneCards pathways - 5/44 super-pathways (see all 44) About this table

See pathways by source

Super-pathway

contained gene-specific pathways

Transcriptional activity of SMAD2/SMAD3:SMAD4 heterotrimer

	Transcriptional activity of SMAD2/SMAD3:SMAD4 heterotrimer	1.00			Signaling by TGF-beta Receptor Complex	0.61
	SMAD2/SMAD3:SMAD4 heterotrimer regulates transcription	0.63			Downregulation of SMAD2/3:SMAD4 transcriptional activity	0.56

Ubiquitin-dependent degradation of the Smad complex terminates BMP2 signalling

	Ubiquitin-dependent degradation of the Smad complex terminates BMP2 signalling	1.00			BMP Signalling Pathway	0.38
	Ubiquitin-dependent degradation of the SMAD complex terminates TGF-beta signaling	0.38			ALK2 signaling events	0.31

Signaling by BMP

	Signaling by BMP	1.00			BMP receptor signaling	0.31
	BMP signalling and regulation	0.35

Wnt Signaling Pathway

	DNA damage response (only ATM dependent)	0.35			Wnt Signaling Pathway NetPath	0.22
	Wnt signaling pathway	0.30

Development_TGF-beta receptor signaling

Development_TGF-beta receptor signaling

1.00

Development TGF-beta receptor signaling

0.96

Pathway sources
See GeneCards unified pathways
Show all pathways

5/8 EMD Millipore Pathways for SMAD4 (see all 8)

	Development Role of Activin A in cell differentiation and proliferation
	Selected targets of Oct-3/4
	Cell cycle Regulation of G1/S transition (part 1)
	Signal transduction JNK pathway
	Signal transduction Activin A signaling regulation

2 R&D Systems Pathways for SMAD4

	Th17 Differentiation
	TGF-beta Signaling Pathways

1 Cell Signaling Technology (CST) Pathway for SMAD4

TGF-beta/Smad Signaling

5/7 GeneGo (Thomson Reuters) Pathways for SMAD4 (see all 7)

	Signal transduction Activin A signaling regulation
	Signal transduction PKA signaling
	Cell cycle Regulation of G1/S transition (part 1)
	Development Role of Activin A in cell differentiation and proliferation
	Cytokine production by Th17 cells in CF (Mouse model)

5/24 BioSystems Pathways for SMAD4 (see all 24)

	Heart Development
	Integrated Breast Cancer Pathway
	TGF Beta Signaling Pathway
	BMP signalling and regulation
	DNA damage response (only ATM dependent)

5/15 Reactome Pathways for SMAD4 (see all 15)

	Developmental Biology
	TGF-beta receptor signaling activates SMADs
	Generic Transcription Pathway
	Transcriptional activity of SMAD2/SMAD3:SMAD4 heterotrimer
	Degradation of SMAD2/3:SMAD4 complex

5/8 Kegg Pathways (Kegg details for SMAD4) (see all 8):

	Cell cycle
	Wnt signaling pathway
	TGF-beta signaling pathway
	Adherens junction
	Pathways in cancer

SMAD4 for pathways About GeneDecksing

Interactions:

SABiosciences Gene Network Central^TM Interacting Genes and Proteins Network for SMAD4

STRING Interaction Network Preview (showing 5 interactants - click image to see 25)

5/399 Interacting proteins for SMAD4 (Q13485^{1, 2, 3} ENSP00000341551⁴) via UniProtKB, MINT, STRING, and/or I2D (see all 399)

Interactant		Interaction Details
GeneCard	External ID(s)	Interaction Details
SMAD2	Q15796¹^,²^,³, ENSP00000262160⁴	EBI-347263,EBI-1040141 MINT-6799119 MINT-15900 MINT-61945 I2D: score=10 STRING: ENSP00000262160
TRIM33	Q9UPN9¹^,²^,³, ENSP00000351250⁴	EBI-347263,EBI-2214398 MINT-8200333 MINT-8200350 I2D: score=4 STRING: ENSP00000351250
UBE2I	P63279¹^,²^,³, ENSP00000324897⁴	EBI-347263,EBI-80168 MINT-61950 I2D: score=7 STRING: ENSP00000324897
SKI	P12755¹^,²^,³, ENSP00000367797⁴	EBI-347263,EBI-347281 MINT-61954 I2D: score=6 STRING: ENSP00000367797
SMAD1	Q15797²^,³, ENSP00000305769⁴	MINT-62001 MINT-4298076 I2D: score=7 STRING: ENSP00000305769

About this table

Gene Ontology (GO): 5/47 biological process terms (GO ID links to tree view) (see all 47): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0000122	negative regulation of transcription from RNA polymerase II promoter	TAS	--
GO:0001658	branching involved in ureteric bud morphogenesis	IEA	--
GO:0001666	response to hypoxia	IMP	12411310
GO:0001701	in utero embryonic development	IEA	--
GO:0001702	gastrulation with mouth forming second	IEA	--

SMAD4 for ontologies About GeneDecksing

(Chemical Compounds according to UniProtKB, Enzo Life Sciences, EMD Millipore, Tocris Bioscience HMDB, BitterDB, and/or Novoseek, and Drugs according to DrugBank, Enzo Life Sciences, and/or PharmGKB, with drugs/clinical trials/news search links to CenterWatch)
About This Section

SMAD4 for compounds About GeneDecksing

Browse Small Molecules at EMD Millipore

Browse drugs & compounds from Enzo Life Sciences

Browse Tocris compounds for SMAD4
10/30 Novoseek chemical compound relationships for SMAD4 gene (see all 30) About this table

Compound	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
tgf beta1	66.2	23	11784716 (4), 11196469 (2), 12935295 (2), 17469181 (2) (see all 10)
threonine	43.3	4	9312063 (1), 14656470 (1), 11889196 (1), 15702493 (1)
serine	38	5	9312063 (1), 14656470 (1), 11889196 (1), 12759229 (1) (see all 5)
crcs	34.8	2	18471510 (2)
leptomycin b	30.2	5	11074002 (2), 18519565 (1)
propoxur	24.3	1	12010891 (1)
pd 98,059	21.8	1	11371552 (1)
doxazosin	9.66	3	12771931 (2), 16007219 (1)
paraffin	6.79	8	11866987 (2), 12479060 (2), 11075857 (1), 12010891 (1)
mg 132	6.25	3	17200344 (2), 11818334 (1)

Search CenterWatch for drugs/clinical trials and news about SMAD4

(Secondary structures according to fRNAdb,
GenBank/EMBL/DDBJ Accessions according to
Unigene (Build 235 Homo sapiens; Mar 10 2013) or GenBank,
RefSeq according to Entrez Gene,
DOTS (version 10), and/or AceView, transcript ids from Ensembl with links to UCSC,
exon structure from GeneLoc, alternative splicing isoforms according to ASD and/or ECgene,
RNAi Products from EMD Millipore,
siRNAs from OriGene, QIAGEN, shRNA from OriGene, microRNA from QIAGEN,
Tagged/untagged cDNA clones from OriGene, SwitchGear Genomics, GenScript, DNA2.0, Vector BioLabs, Primers from OriGene, SABiosciences, and/or QIAGEN )
About This Section

REFSEQ mRNAs for SMAD4 gene:

NM_005359.5

Unigene Cluster for SMAD4:

SMAD family member 4

Hs.75862 [show with all ESTs]

Unigene Representative Sequence: NM_005359

17 Ensembl transcripts including schematic representations, and UCSC links where relevant:

ENST00000588860 ENST00000398417(uc002lfb.4) ENST00000589076 ENST00000590061

ENST00000591914 ENST00000592911 ENST00000589941 ENST00000588745 ENST00000592186

ENST00000589706 ENST00000585448 ENST00000591126 ENST00000590499 ENST00000593223

ENST00000586253 ENST00000342988(uc010xdp.2) ENST00000452201(uc010xdo.1)

miRNA Products:		OriGene 3'-UTR Clone: SMAD4 Browse OriGene MicroRNA Expression Plasmids
		QIAGEN Custom miScript Target Protector blocks miRNA-binding site of human, mouse, rat SMAD4
		8/195 QIAGEN miScript miRNA Assays for microRNAs that regulate SMAD4 (see all 195):
		hsa-miR-411* hsa-miR-106a hsa-miR-146a hsa-miR-605 hsa-miR-519a hsa-miR-3921 hsa-miR-298 hsa-miR-130b

		SwitchGear 3'UTR luciferase reporter plasmid: SMAD4 3' UTR sequence

Inhib. RNA Products:		Browse for Gene Knock-down Tools from EMD Millipore
		OriGene 29mer shRNA kits in GFP-retroviral vector in human, mouse, rat for SMAD4 (see all 7) OriGene shRNA RFP: SMAD4 OriGene siRNA: SMAD4
		QIAGEN FlexiTube/FlexiPlate siRNA for gene silencing in human, mouse, rat SMAD4

Clone Products:		OriGene Myc/DDK tagged cDNA clones in CMV expression vector in human, mouse, rat for SMAD4 (see all 3) OriGene untagged cDNA clones in CMV expression vector in human, mouse, rat for SMAD4 OriGene custom cloning services � gene synthesis, subcloning, mutagenesis, variant library, vector shuttling
		GenScript: all cDNA clones in your preferred vector: SMAD4 (NM_005359)
		DNA2.0 Custom Codon Optimized Gene Synthesis Service for SMAD4
		Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat SMAD4

Primer Products:		OriGene genome-wide validated SYBR primer pairs in human, mouse, rat for SMAD4 Browse OriGene validated miRNA SYBR primer pairs
		SABiosciences RT² qPCR Primer Assay in human, mouse, rat SMAD4
		QIAGEN QuantiTect SYBR Green Assays in human, mouse, rat SMAD4
		QIAGEN QuantiFast Probe-based Assays in human, mouse, rat SMAD4

Additional cDNA sequence:

AK290770.1 AL832789.1 BC002379.2 BX647129.1 U44378.1

17 DOTS entries:

DT.317208 DT.121090072 DT.105413 DT.100691422 DT.40284956 DT.100001068 DT.97802720 DT.121090073

DT.92438228 DT.92438229 DT.95078262 DT.100818887 DT.121087023 DT.40264776 DT.75168357 DT.91684967

DT.95370406

24/272 AceView cDNA sequences (see all 272):

BE926578 AI439735 AA815191 CB155810 BU783336 BI495903 BC002379 BQ270199

BF842398 BQ186401 CA489596 AA456439 AW169700 AI985283 AA810812 BU684427

CD657319 F08858 AW087656 NM_005359 CR616451 AU119862 BU620105 AA954065

GeneLoc Exon Structure

5/10 Alternative Splicing Database (ASD) splice patterns (SP) for SMAD4 (see all 10) About this scheme

ExUns:	1	^	2	^	3a	·	3b	^	4	^	5	^	6a	·	6b	·	6c	^	7a	·	7b	·	7c	^	8a	·	8b	^	9a	·	9b	^	10	^	11	^	12	^	13	^	14a	·	14b	^	15	^	16	^	17
SP1:																											-		-												-						-
SP2:																																									-						-
SP3:			-		-		-		-		-
SP4:
SP5:

ECgene alternative splicing isoforms for SMAD4

(RNA expression data according to H-InvDB, NONCODE, miRBase, and RNAdb, Expression images according to data from BioGPS, Illumina Human BodyMap, and CGAP SAGE, Sets of similar genes according to GeneDecks, in vivo and in vitro expression data from LifeMap Discovery™, plus additional links to Genevestigator, and/or SOURCE, and/or BioGPS, and/or UniProtKB,
PCR Arrays from SABiosciences, Primers from OriGene, SABiosciences, and/or QIAGEN, In Situ Hybridization Assays from Advanced Cell Diagnostics)
About This Section

SMAD4 expression in normal human tissues (normalized intensities)
See probesets specificity/sensitivity at GeneAnnot
About this imageBioGPS
CGAP TAG: TATGTTATGA

About this image

SMAD4 expression in embryonic tissues and stem cells

Expression by the Database of Embryonic development, Stem cell research, and Regenerative medicine About this table

6 LifeMap In Vivo Development Anatomical Compartments/Cells

Tissue

Anatomical Compartment

Cell

Category (developmental path)

Extraembryonic Mesoderm

Extraembryonic Mural Cells

Extraembryonic Tissues

Head Mesenchyme

Branchial Arch 3,4,6

Cardiac Neural Crest Cells

Neural Crest

Heart

Outflow Tract

Neural Crest Outflow Tract Cells

Neural Crest

Lateral Plate Mesoderm

Splanchnic Mesoderm

Mural Cells

Pericytes, Smooth Muscle

Primitive Streak

Primitive Streak Cells

Early Embryo

Brain

Medulla Oblongata

Brain

Expression:

Positive

Negative

Selective marker

Experimental details:

Curated

Microarrays

In-situ hybridization

See SMAD4 Protein Expression from SPIRE MOPED and PaxDB
Genevestigator expression for SMAD4

SOURCE GeneReport for Unigene cluster: Hs.75862

SABiosciences Expression via Pathway-Focused PCR Arrays including SMAD4 (see all 13):

Osteogenesis in human mouse rat

Oncogenes & Tumor Suppressor Genes in human mouse rat

Fibrosis in human mouse rat

TGFB/BMP Signaling Pathway in human mouse rat

Stem Cell Signaling in human mouse rat

Primer Products:		OriGene genome-wide validated SYBR primer pairs in human, mouse, rat for SMAD4 Browse OriGene validated miRNA SYBR primer pairs
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In Situ
Assay Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for SMAD4

(Orthologs according to ^1,2HomoloGene (²older version, for species not in ¹newer version), ³euGenes, ⁴SGD , ⁵MGI Mar 06 2013, with possible further links to Flybase and/or WormBase, and/or ⁶Ensembl pan taxonomic compara , Gene Trees according to Ensembl and TreeFam)
About This Section

This gene was present in the common ancestor of animals.

Orthologs for SMAD4 gene from 5/22 species (see all 22) About this table

Organism	Taxonomic classification	Gene	Description	Human Similarity	Orthology Type	Details
lizard (Anolis carolinensis)	Reptilia	SMAD4⁶	--	95(a)	1 ↔ 1	GL343471.1(494708-516203)
African clawed frog (Xenopus laevis)	Amphibia	Xl.885²	Xenopus laevis mRNA for Xsmad4a, complete cds	78.03(n)		AB022721.1
zebrafish (Danio rerio)	Actinopterygii	madh4²	MAD homolog 4 (Drosophila)	77.62(n)		30640 AF229175.1
fruit fly (Drosophila melanogaster)	Insecta	Med¹	Medea	61.24(n) 70.6(a)		43725 NM_170559.2 NP_733438.1
worm (Caenorhabditis elegans)	Secernentea	sma-4⁶ daf-3⁶	abnormal DAuer Formation family member (daf-3)	33(a) 15(a)	possible ortholog 1 ↔ 1	III(5816159-5820159) X(817926-825474)

ENSEMBL Gene Tree for SMAD4 (if available)
TreeFam Gene Tree for SMAD4 (if available)

(Paralogs according to ¹HomoloGene,
²Ensembl, and ³SIMAP, Pseudogenes according to ⁴Pseudogene.org Build 68)
About This Section

Paralogs for SMAD4 gene

SMAD2² SMAD3² SMAD1² SMAD5² SMAD6² SMAD9² SMAD7²

6 SIMAP similar genes for SMAD4 using alignment to 11 protein entries: SMAD4_HUMAN (see all proteins):

SMAD1 DKFZp686J10186 SMAD5 SMAD3 SMAD7 SMAD2

SMAD4 for paralogs About GeneDecksing

(SNPs/Variants according to the ¹NCBI SNP Database, ²Ensembl, ³PupaSUITE, UniProtKB, and DNA2.0, Linkage Disequilibrium by HapMap, Structural Variations(CNVs/InDels/Inversions) from the Database of Genomic Variants, Mutations from the Human Gene Mutation Database (HGMD) and the Locus Specific Mutation Databases (LSDB), Blood group antigen gene mutations by BGMUT, Resequencing Primers from QIAGEN, Cancer Mutation PCR Arrays and Assays and Copy Number PCR Arrays from SABiosciences)
About This Section

SNP ID	Valid	Clinical significance	Chr 18 pos	Sequence	#	AA Chg	Type	More	#	Allele freq	Pop	Total sample	More
Genomic Data					Transcription Related Data				Allele Frequencies
				--	--	--		--	--
rs80338963^1,2	C	pathogenic	45447422(+)	`GAGATC/TGCTTT`	2	R C	mis¹		0	--	--	--	--
rs9951928^1,2	C,F,	--	45410092(+)	`TGATGA/CTGTTA`	1	--	us2k¹		1		WA	118
rs36222037^1,2	C,F,	--	45411538(-)	`CCTTCA/CACAGC`	1	--	us2k¹		1		WA	118
rs78950659^1,2	F,	--	45412956(+)	`ACTGCA/GTGTAC`	1	--	int¹		1		NA	120
rs9961921^1,2	C,F,	--	45413096(+)	`GTGCAG/AAGGCT`	1	--	int¹		1		WA	118
rs78685185^1,2		--	45413122(+)	`GAAGAC/TTGATA`	1	--	int¹		0	--	--	--	--
rs112190235^1,2		--	45413198(+)	`TAGTTT/CCTTAA`	1	--	int¹		2		CSA WA	120
rs112512652^1,2	C,	--	45413313(+)	`AACATA/GTAATC`	1	--	int¹		2		NA	122
rs78985558^1,2		--	45413359(+)	`CAGAAT/GTAGCA`	1	--	int¹		1		WA	118
rs1787110^1,2	C,H	--	45413494(-)	`ATGCCT/CTGTTC`	1	--	int¹		6		NS EA NA	422

HapMap Linkage Disequilibrium report for SMAD4 (48494410 - 48611415 bp)
Structural Variations (Copy Number Variations, Insertions/Deletions, Inversions)
Database of Genomic Variants (DGV) variations for SMAD4: --
Human Gene Mutation Database (HGMD): SMAD4

Locus Specific Mutation Databases (LSDB): SMAD4

1 SABiosciences Cancer Mutation PCR Assay for SMAD4:

Cosmic Id	AA Change
14057	p.Q245*

2 SABiosciences Cancer Mutation PCR Arrays containing SMAD4:

Pancreatic Cancer

Cancer Comprehensive Panel 384HT

1 SABiosciences qBiomarker Copy Number PCR Array containing SMAD4:

Oncogenes & Tumor Suppressor Genes 384HC

QIAGEN SeqTarget long-range PCR primers in human, mouse, rat for resequencing SMAD4

DNA2.0 Custom Variant and Variant Library Synthesis for SMAD4

(in which this Gene is Involved, According to MalaCards, OMIM, UniProtKB, the University of Copenhagen DISEASES database, Novoseek, Genatlas, GeneTests, GAD, HuGE Navigator, and/or TGDB.)
About This Section

SMAD4 for disorders           About GeneDecksing

OMIM gene information: 600993
OMIM disorders: 174900  175050
UniProtKB/Swiss-Prot: SMAD4_HUMAN, Q13485

Defects in SMAD4 are a cause of pancreatic cancer (PNCA) [MIM:260350]

Defects in SMAD4 are a cause of juvenile polyposis syndrome (JPS) [MIM:174900]; also known as juvenile

intestinal polyposis (JIP). JPS is an autosomal dominant gastrointestinal hamartomatous polyposis syndrome in which

patients are at risk for developing gastrointestinal cancers. The lesions are typified by a smooth histological

appearance, predominant stroma, cystic spaces and lack of a smooth muscle core. Multiple juvenile polyps usually occur

in a number of Mendelian disorders. Sometimes, these polyps occur without associated features as in JPS; here, polyps

tend to occur in the large bowel and are associated with an increased risk of colon and other gastrointestinal cancers

Defects in SMAD4 are a cause of juvenile polyposis/hereditary hemorrhagic telangiectasia syndrome (JP/HHT)

[MIM:175050]. JP/HHT syndrome phenotype consists of the coexistence of juvenile polyposis (JIP) and hereditary

hemorrhagic telangiectasia (HHT) [MIM:187300] in a single individual. JIP and HHT are autosomal dominant disorders

with distinct and non-overlapping clinical features. The former, an inherited gastrointestinal malignancy

predisposition, is caused by mutations in SMAD4 or BMPR1A, and the latter is a vascular malformation disorder caused

by mutations in ENG or ACVRL1. All four genes encode proteins involved in the transforming-growth-factor-signaling

pathway. Although there are reports of patients and families with phenotypes of both disorders combined, the genetic

etiology of this association is unknown

Defects in SMAD4 may be a cause of colorectal cancer (CRC) [MIM:114500]

Defects in SMAD4 may be a cause of primary pulmonary hypertension (PPH1) [MIM:178600]. A rare disorder

characterized by plexiform lesions of proliferating endothelial cells in pulmonary arterioles. The lesions lead to

elevated pulmonary arterial pression, right ventricular failure, and death. The disease can occur from infancy

throughout life and it has a mean age at onset of 36 years. Penetrance is reduced. Although familial PPH1 is rare,

cases secondary to known etiologies are more common and include those associated with the appetite-suppressant drugs

Defects in SMAD4 are the cause of Myhre syndrome (MYHRS) [MIM:139210]. MYHRS is a syndrome characterized by

pre- and postnatal growth deficiency, mental retardation, generalized muscle hypertrophy and striking muscular build,

decreased joint mobility, cryptorchidism, and unusual facies. Dysmorphic facial features include microcephaly, midface

hypoplasia, prognathism, and blepharophimosis. Typical skeletal anomalies are short stature, square body shape, broad

ribs, iliac hypoplasia, brachydactyly, flattened vertebrae, and thickened calvaria. Other features, such as congenital

heart disease, may also occur

20/114 diseases for SMAD4 (see all 114): About MalaCards

hereditary hemorrhagic telangiectasia juvenile polyposis syndrome pancreatitis bannayan-riley-ruvalcaba syndrome

polyposis pancreatic carcinoma juvenile polyposis/hereditary hemorrhagic telangiectasia syndrome carcinoma

pancreatic cancer mutyh-associated polyposis peutz-jeghers syndrome familial adenomatous polyposis

arteriovenous malformation non-small cell lung carcinoma biliary tract cancer bile duct adenoma

testicular germ cell tumor vipoma clear cell adenocarcinoma bile duct cancer

5 diseases from the University of Copenhagen DISEASES database for SMAD4:

Pancreatic cancer Carcinoma Hereditary hemorrhagic telangiectasia Colorectal cancer

Adenoma

10/94 Novoseek disease relationships for SMAD4 gene (see all 94) About this table

Disease	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
juvenile polyposis syndrome	88	12	19014666 (2), 16152648 (1), 17873119 (1), 15378062 (1) (see all 8)
pancreatic cancer	81	126	11323508 (3), 12479060 (3), 11783110 (3), 15592526 (3) (see all 65)
polyps juvenile	80.8	9	19266212 (5), 15447767 (1), 19014666 (1)
pancreatic carcinoma	77.5	95	18985820 (6), 9098646 (5), 12352188 (4), 11866987 (3) (see all 37)
polyposis	77.4	50	10797267 (3), 15031030 (3), 18355998 (3), 12821112 (2) (see all 32)
allelic loss	70.3	33	17088901 (2), 17659731 (2), 19443408 (2), 11337365 (2) (see all 18)
adenocarcinoma	68	148	11751510 (6), 11431719 (5), 17587453 (5), 17854080 (5) (see all 58)
pancreatic tumor	67.3	7	16135802 (2), 10415855 (1), 9644295 (1), 11553622 (1) (see all 6)
tumors	65.6	415	15814640 (6), 12352188 (6), 15014009 (5), 10331746 (4) (see all 99)
colorectal cancer	61.6	78	11172591 (4), 8898652 (4), 11481457 (4), 16144935 (3) (see all 39)

GeneTests: SMAD4

Juvenile Polyposis Syndrome

Hereditary Hemorrhagic Telangiectasia

Genetic Association Database (GAD): SMAD4
Human Genome Epidemiology (HuGE) Navigator: SMAD4 (37 documents)

Export disorders for SMAD4 gene to outside databases

(in PubMed. Associations of this gene to articles via ¹Entrez Gene, ²UniProtKB/Swiss-Prot, ³HGNC, ⁴GAD, ⁵PharmGKB, ⁶HMDB, ⁷DrugBank, ⁸UniProtKB/TrEMBL, ⁹ Novoseek, and/or ¹⁰fRNAdb)
About This Section

PubMed articles for SMAD4 gene, integrated from 9 sources (see all 961):
(articles sorted by number of sources associating them with SMAD4)

Utopia: connect your pdf to the dynamic
world of online information

Germline SMAD4 or BMPR1A mutations and phenotype of juvenile polyposis. (PubMed id 12417513)^{1, 2, 4, 9} Sayed M.G.... Howe J.R. (2002)
DPC4, a candidate tumor suppressor gene at human chromosome 18q21.1. (PubMed id 8553070)^{1, 2, 3} Hahn S.A.... Kern S.E. (1996)
Receptor-associated Mad homologues synergize as effectors of the TGF- beta response. (PubMed id 8774881)^{1, 2, 3} Zhang Y.... Derynck R. (1996)
The prevalence of MADH4 and BMPR1A mutations in juvenile polyposis and absence of BMPR2, BMPR1B, and ACVR1 mutations. (PubMed id 15235019)^{1, 4, 9} Howe J.R....Lynch H.T. (2004)
Mutations in DPC4 (SMAD4) cause juvenile polyposis syndrome, but only account for a minority of cases. (PubMed id 9811934)^{1, 2, 9} Houlston R.... Tomlinson I. (1998)
Jab1 antagonizes TGF-beta signaling by inducing Smad4 degradation. (PubMed id 11818334)^{1, 2, 9} Wan M.... Cao X. (2002)
A combined syndrome of juvenile polyposis and hereditary haemorrhagic telangiectasia associated with mutations in MADH4 (SMAD4). (PubMed id 15031030)^{1, 2, 9} Gallione C.J.... Marchuk D.A. (2004)
Genomic sequencing of DPC4 in the analysis of familial pancreatic carcinoma. (PubMed id 9098646)^{1, 2, 9} Moskaluk C.A....Kern S.E. (1997)
Dual role of the Smad4/DPC4 tumor suppressor in TGFbeta-inducible transcriptional complexes. (PubMed id 9389648)^{1, 2, 9} Liu F.... Massague J. (1997)
Potentiation of Smad-mediated transcriptional activation by the RNA- binding protein RBPMS. (PubMed id 17099224)^{1, 2, 9} Sun Y.... Ye Q. (2006)

(in PubMed, OMIM, and NCBI Bookshelf)
About This Section

		AND	OR
Aliases
Disorders
Free Text

Query String		PubMed OMIM NCBI Bookshelf
	(Note: In FireFox, select the above section and copy using Ctrl-C)

(According to Entrez Gene, HGNC, AceView, euGenes, Ensembl, miRBase, ECgene, Kegg, and/or H-InvDB)
About This Section

Entrez Gene: 4089	HGNC: 6770	AceView: SMAD4andELAC1	Ensembl:ENSG00000141646	euGenes: HUgn4089
ECgene: SMAD4	Kegg: 4089	H-InvDB: SMAD4

(According to HUGE)
About This Section

(According to PharmGKB, ATLAS, HORDE, IMGT, LEIDEN, UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL,
Wikipedia and/or GeneReviews via UniProtKB/Swiss-Prot)
About This Section

Name	Description
PharmGKB entry for SMAD4	Pharmacogenomics, SNPs, Pathways
ATLAS Chromosomes in Cancer entry for SMAD4	Genetics and Cytogenetics in Oncology and Haematology
GeneReviews	http://www.ncbi.nlm.nih.gov/sites/GeneTests/lab/gene/SMAD4

Intellectual Property for SMAD4 gene
(Patent information from GeneIP, Licensable technologies from WIS Yeda, Salk, Tufts, IP news from LifeMap Sciences, Inc.)

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Patent Information for SMAD4 gene:
Search GeneIP for patents involving SMAD4

GeneCards and IP:
Japan Patent Office Licenses GeneCards European Patent Office Licenses GeneCards Improving the IP Search

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