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Category Symbol Source: HGNC EntrezGene Ensembl GeneCards RNA genes CroW21

GIFtS
-

TWSG1 Gene

protein-coding GIFtS: 50
GCID: GC18P009324

twisted gastrulation homolog 1 (Drosophila)

Explore 1 disease affiliated with
TWSG1 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
Twisted Gastrulation Homolog 1 (Drosophila)¹ ²
TSG¹ ² ³ ⁵
Twisted Gastrulation Protein Homolog 1²

External Ids:	HGNC: 12429¹	Entrez Gene: 57045²	Ensembl: ENSG00000128791⁷	OMIM: 605049⁵	UniProtKB: Q9GZX9³

Export aliases for TWSG1 gene to outside databases

Previous GC identifers: GC00U990679 GC18P009391

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

UniProtKB/Swiss-Prot: TWSG1_HUMAN, Q9GZX9

Function: May be involved in dorsoventral axis formation. Seems to antagonize BMP signaling by forming ternary

complexes with CHRD and BMPs, thereby preventing BMPs from binding to their receptors. In addition to the anti-BMP

function, also has pro-BMP activity, partly mediated by cleavage and degradation of CHRD, which releases BMPs from

ternary complexes. May be an important modulator of BMP-regulated cartilage development and chondrocyte

differentiation. May play a role in thymocyte development (By similarity)

Gene Wiki entry for TWSG1

(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_010859.14

Regulatory elements:

SABiosciences Regulatory transcription factor binding sites in the TWSG1 gene promoter:
TBP STAT5B ISGF-3 STAT5A AP-2gamma HNF-1A HNF-1 TFIID AP-2beta AP-2alphaA
Other transcription factors

SwitchGear Promoter luciferase reporter plasmid: TWSG1 promoter sequence

Search SABiosciences Chromatin IP Primers for TWSG1

Epigenetics:

QIAGEN PyroMark CpG Assay predesigned Pyrosequencing DNA Methylation assays in human, mouse, rat TWSG1

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

Entrez Gene cytogenetic band: 18p11.3 Ensembl cytogenetic band: 18p11.22 HGNC cytogenetic band: 18p11.3

TWSG1 Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
TWSG1 gene location

GeneLoc information about chromosome 18 GeneLoc Exon Structure

GeneLoc location for GC18P009324: view genomic region (about GC identifiers)

Start:	9,334,765 bp from pter	End:	9,402,418 bp from pter
Size:	67,654 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: TWSG1_HUMAN, Q9GZX9 (See protein sequence)

Recommended Name: Twisted gastrulation protein homolog 1 precursor

Size: 223 amino acids; 25017 Da

Subunit: Interacts with CHRD and BMP4. This interaction enhances CHRD/BMP4 complex formation. Interacts with BMP7 (By

similarity)

Subcellular location: Secreted (By similarity)

Developmental stage: Expressed in brain throughout development

Sequence caution: Sequence=BAB55246.1; Type=Erroneous translation; Note=Wrong choice of CDS;

Secondary accessions: B2RE08 D3DUH9 Q8NBI7 Q96K46

Alternative splicing: 2 isoforms: Q9GZX9-1 Q9GZX9-2 (No experimental confirmation available. May be produced at very low levels due to a premature stop codon in the mRNA, leading to nonsense-mediated mRNA decay)

Explore the universe of human proteins at neXtProt for TWSG1: NX_Q9GZX9

Post-translational modifications:

View modification sites using PhosphoSitePlus²

View neXtProt modification sites for NX_Q9GZX9

TWSG1 Protein expression data from MOPED and PaxDb: About this image

REFSEQ proteins: NP_065699.1

ENSEMBL proteins:

ENSP00000463331 ENSP00000262120 ENSP00000464458

Human Recombinant Protein Products for TWSG1:

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	Uscn Proteins for TWSG1

Gene Ontology (GO): 1 cellular component term (GO ID links to tree view): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005615	extracellular space	IEA	--

TWSG1 for ontologies About GeneDecksing

TWSG1 Antibody Products:

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

TWSG1 for domains About GeneDecksing

1 InterPro domain/family:

IPR006761 Tsg

Graphical View of Domain Structure for InterPro Entry Q9GZX9

ProtoNet protein and cluster: Q9GZX9

UniProtKB/Swiss-Prot: TWSG1_HUMAN, Q9GZX9

Domain: The N-terminal domain is sufficient to interact with BMP4 (By similarity)

Similarity: Belongs to the twisted gastrulation protein family

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

Molecular Function:

UniProtKB/Swiss-Prot Summary: TWSG1_HUMAN, Q9GZX9

Function: May be involved in dorsoventral axis formation. Seems to antagonize BMP signaling by forming ternary

complexes with CHRD and BMPs, thereby preventing BMPs from binding to their receptors. In addition to the anti-BMP

function, also has pro-BMP activity, partly mediated by cleavage and degradation of CHRD, which releases BMPs from

ternary complexes. May be an important modulator of BMP-regulated cartilage development and chondrocyte

differentiation. May play a role in thymocyte development (By similarity)

Gene Ontology (GO): 1 molecular function term (GO ID links to tree view): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005515	protein binding	--	--

TWSG1 for ontologies About GeneDecksing

Phenotypes:
3 GenomeRNAi human phenotypes for TWSG1:

G0/1 arrest

Increased G1 DNA content

Increased HPV18 LCR reporter a

15/19 MGI mutant phenotypes (inferred from 6 alleles) (MGI details for Twsg1) (see all 19):

behavior/neurological	cardiovascular system	cellular	craniofacial	digestive/alimentary
embryogenesis	endocrine/exocrine gland	growth/size	hearing/vestibular/ear	hematopoietic system
homeostasis/metabolism	immune system	limbs/digits/tail	mortality/aging	nervous system

TWSG1 for phenotypes About GeneDecksing

Animal Models:
Mouse knock-outs for TWSG1: Twsg1^tm1Nosa Twsg1^tm1Dgen Twsg1^tm1.1Mboc

inGenious Targeting Laboratory - Customizable classic, inducible, and humanized mouse model solutions for TWSG1

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		hsa-miR-3671 hsa-miR-136 hsa-miR-548am hsa-miR-7-2* hsa-miR-606 hsa-miR-548aa hsa-miR-20b* hsa-miR-664

		SwitchGear 3'UTR luciferase reporter plasmid: TWSG1 3' UTR sequence

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In Situ Assay
Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for TWSG1

(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

Interactions:

SABiosciences Gene Network Central^TM Interacting Genes and Proteins Network for TWSG1

STRING Interaction Network Preview (showing 3 interactants - click image to see more details)

5 Interacting proteins for TWSG1 (Q9GZX9³ ENSP00000262120⁴) via UniProtKB, MINT, STRING, and/or I2D

Interactant		Interaction Details
GeneCard	External ID(s)	Interaction Details
CHRD	Q9H2X0³, ENSP00000204604⁴	I2D: score=3 STRING: ENSP00000204604
BMP4	P12644³, ENSP00000245451⁴	I2D: score=2 STRING: ENSP00000245451
BMP1	P13497³, ENSP00000305714⁴	I2D: score=1 STRING: ENSP00000305714
EEF2	P13639³	I2D: score=5
SIRT4	Q9Y6E7³	I2D: score=3

About this table

Gene Ontology (GO): 5/13 biological process terms (GO ID links to tree view) (see all 13): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0001503	ossification	IEA	--
GO:0001707	mesoderm formation	IEA	--
GO:0007435	salivary gland morphogenesis	IEA	--
GO:0009790	embryo development	--	--
GO:0009888	tissue development	--	--

TWSG1 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

Browse Small Molecules at EMD Millipore

Browse drugs & compounds from Enzo Life Sciences

Browse Tocris compounds for TWSG1
Search CenterWatch for drugs/clinical trials and news about TWSG1

(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,
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About This Section

REFSEQ mRNAs for TWSG1 gene:

NM_020648.5

Unigene Cluster for TWSG1:

Twisted gastrulation homolog 1 (Drosophila)

Hs.514685 [show with all ESTs]

Unigene Representative Sequence: NM_020648

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

ENST00000583147 ENST00000262120(uc002knz.3 uc002koa.3) ENST00000581641

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		hsa-miR-3671 hsa-miR-136 hsa-miR-548am hsa-miR-7-2* hsa-miR-606 hsa-miR-548aa hsa-miR-20b* hsa-miR-664

		SwitchGear 3'UTR luciferase reporter plasmid: TWSG1 3' UTR sequence

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Additional cDNA sequence:

AF196834.1 AF294628.1 AJ297391.1 AK027629.1 AK075550.1 AK315751.1 BC020490.1

7 DOTS entries:

DT.313720 DT.97858870 DT.92434329 DT.100788882 DT.121106537 DT.95171956 DT.95171963

24/209 AceView cDNA sequences (see all 209):

N91767 CR608086 AJ297391 AA236064 AI129471 AL711102 BP342527 BM925839

BP343234 AA235953 AI087263 CK903960 BU631678 CF135658 BM053044 AI700813

BM683178 AA905905 AI670046 BQ015603 CF529431 AA329449 AI478573 BM719230

GeneLoc Exon Structure

5/6 Alternative Splicing Database (ASD) splice patterns (SP) for TWSG1 (see all 6) About this scheme

ExUns:	1a	·	1b	^	2a	·	2b	·	2c	^	3	^	4a	·	4b	^	5	^	6	^	7	^	8a	·	8b	^	9a	·	9b
SP1:											-						-								-
SP2:											-						-		-		-				-
SP3:											-						-		-		-
SP4:									-		-		-				-		-		-
SP5:

ECgene alternative splicing isoforms for TWSG1

(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

TWSG1 expression in normal human tissues (normalized intensities)
See probesets specificity/sensitivity at GeneAnnot
About this imageBioGPS
CGAP TAG: TGATTTCTAA

About this image
See TWSG1 Protein Expression from SPIRE MOPED and PaxDB
Genevestigator expression for TWSG1

SOURCE GeneReport for Unigene cluster: Hs.514685
SABiosciences Custom PCR Arrays for TWSG1

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In Situ
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(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 TWSG1 gene from 6/19 species (see all 19) About this table

Organism	Taxonomic classification	Gene	Description	Human Similarity	Orthology Type	Details
mouse (Mus musculus)	Mammalia	Twsg1^{1 , 5}	twisted gastrulation homolog 1 (Drosophila)^{1, 5}	84.68(n)¹ 93.69(a)¹		17 (35.26 cM)⁵ 65960¹ NM_023053.3¹ NP_075540.1¹ 65923066⁵
chicken (Gallus gallus)	Aves	TWSG1¹	twisted gastrulation homolog 1 (Drosophila)	82.24(n) 93.46(a)		374024 NM_204198.1 NP_989529.1
lizard (Anolis carolinensis)	Reptilia	TWSG1⁶ --	--	87(a) 39(a)	1 ↔ 1 possible ortholog	5(810830-823490) GL343776.1(533-2765)
African clawed frog (Xenopus laevis)	Amphibia	Xl.778²	Xenopus laevis twisted gastrulation protein mRNA, complete more	80.56(n)		AF279246.1
zebrafish (Danio rerio)	Actinopterygii	tsga²	twisted gastrulation a	77.98(n)		84037 AF261692.1
fruit fly (Drosophila melanogaster)	Insecta	cv¹	crossveinless	48(n) 42(a)		44510 NM_080525.2 NP_536786.1

ENSEMBL Gene Tree for TWSG1 (if available)
TreeFam Gene Tree for TWSG1 (if available)

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

(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
				--	--	--		--	--
rs149434392^1,2	C	--	9343210(+)	`TTGTT-/ATATATATATATAT ATATATATATATATATATATAT`	1	--	int¹		0	--	--	--	--
rs6146209^1,2	C	--	9343228(+)	`tatat-/ATATATATATATATA TATATATATATATATATcttgt`	1	--	int¹		1		NA	2
rs59110912^1,2	C	--	9344530(+)	`TTTTTTTTT/- GAGAG`	1	--	int¹		1		NA	2
rs35392295^1,2	C	--	9350270(+)	`ACATCATT/-GTTTC`	1	--	int¹		2		NA CSA	4
rs67687469^1,2	C	--	9353217(+)	`TAAGG-/ATGCATTT TTCAGAATGTATCCC`	1	--	int¹		0	--	--	--	--
rs33957932^1,2	C	--	9353219(+)	`AGGAT-/GCATTTTT CAGAATGTATCCCTA`	1	--	int¹		1		NA	2
rs143374240^1,2	C	--	9369135(+)	`TAAAT-/AAATA AAATA`	1	--	int¹		0	--	--	--	--
rs35584133^1,2	C	--	9369195(+)	`TTTTGA/GGAATT`	1	--	int¹		6		NA CSA WA EA	363
rs141046705^1,2		--	9369254(+)	`ATTAAA/TGTCTA`	1	--	int¹		0	--	--	--	--
rs113363814^1,2	C	--	9369304(+)	`CTCTGC/TCACCC`	1	--	int¹		2		WA CSA	3

HapMap Linkage Disequilibrium report for TWSG1 (9334765 - 9402418 bp)
Structural Variations (Copy Number Variations, Insertions/Deletions, Inversions)
Database of Genomic Variants (DGV): 5 variations for TWSG1
1 CNV: 5045
4 Indels: 11920 11921 61598 40214

SABiosciences Cancer Mutation PCR Assays

QIAGEN SeqTarget long-range PCR primers in human, mouse, rat for resequencing TWSG1

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

TWSG1 for disorders           About GeneDecksing

OMIM gene information: 605049    OMIM disorders: --

1 disease for TWSG1:    About MalaCards

holoprosencephaly

1 disease from the University of Copenhagen DISEASES database for TWSG1:

Holoprosencephaly

Human Genome Epidemiology (HuGE) Navigator: TWSG1 (1 document)

Export disorders for TWSG1 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 TWSG1 gene, integrated from 9 sources (see all 19):
(articles sorted by number of sources associating them with TWSG1)

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

Homologues of Twisted gastrulation are extracellular cofactors in antagonism of BMP signalling. (PubMed id 11260715)^{1, 2, 3} Scott I.C....Greenspan D.S. (2001)
Evolutionary conservation, developmental expression, and genomic mapping of mammalian Twisted gastrulation. (PubMed id 11420619)^{1, 2, 9} Graf D.... Merkenschlager M. (2001)
Signal sequence and keyword trap in silico for selection of full- length human cDNAs encoding secretion or membrane proteins from oligo- capped cDNA libraries. (PubMed id 16303743)^{1, 2} Otsuki T....Isogai T. (2005)
Complete sequencing and characterization of 21,243 full-length human cDNAs. (PubMed id 14702039)^{1, 2} Ota T.... Sugano S. (2004)
Minimal evidence for a direct involvement of twisted gastrulation homolog 1 (TWSG1) gene in human holoprosencephaly. (PubMed id 21227728)¹ Kauvar E.F....Muenke M. (2011)
Identification of TWSG1 as a second novel erythroid r egulator of hepcidin expression in murine and human cells. (PubMed id 19414861)¹ Tanno T....Miller J.L. (2009)
hORFeome v3.1: a resource of human open reading frames representing over 10,000 human genes. (PubMed id 17207965)¹ Lamesch P.... Vidal M. (2007)
Twisted gastrulation (Tsg) is regulated by Tob and enhances TGF-beta signaling in activated T lymphocytes. (PubMed id 17164348)¹ Tzachanis D....Boussiotis V.A. (2007)
Diversification of transcriptional modulation: large-scale identification and characterization of putative alternative promoters of human genes. (PubMed id 16344560)¹ Kimura K.... Sugano S. (2006)
DNA sequence and analysis of human chromosome 18. (PubMed id 16177791)² Nusbaum C.... Lander E.S. (2005)

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(According to Entrez Gene, HGNC, AceView, euGenes, Ensembl, miRBase, ECgene, Kegg, and/or H-InvDB)
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Entrez Gene: 57045	HGNC: 12429	AceView: TWSG1	Ensembl:ENSG00000128791	euGenes: HUgn57045
ECgene: TWSG1	H-InvDB: TWSG1

(According to HUGE)
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(According to PharmGKB, ATLAS, HORDE, IMGT, LEIDEN, UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL,
Wikipedia and/or GeneReviews via UniProtKB/Swiss-Prot)
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Name	Description
PharmGKB entry for TWSG1	Pharmacogenomics, SNPs, Pathways

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TWSG1 Gene

twisted gastrulation homolog 1 (Drosophila)

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TWSG1 Gene

twisted gastrulation homolog 1 (Drosophila)

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