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Category Symbol Source: HGNC EntrezGene Ensembl GeneCards RNA genes CroW21

GIFtS
-

GREM1 Gene

protein-coding GIFtS: 59
GCID: GC15P033010

gremlin 1, DAN family BMP antagonist

(Previous names: cysteine knot superfamily 1, BMP antagonist 1, gremlin...)
(Previous symbol: CKTSF1B1)

Explore 35 diseases affiliated with
GREM1 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
Gremlin 1, DAN Family BMP Antagonist¹ ²		DAN Domain Family Member 2² ³
CKTSF1B1¹ ² ³ ⁵		Gremlin¹
Cysteine Knot Superfamily 1, BMP Antagonist 1¹ ² ³		Gremlin 1, Cysteine Knot Superfamily, Homolog (Xenopus Laevis)¹
DAND2¹ ² ³		GREMLIN²
DRM¹ ² ³		Gremlin 1, Cysteine Knot Superfamily, Homolog²
Gremlin 1¹ ²		Gremlin 1-Like Protein²
Cell Proliferation-Inducing Gene 2 Protein² ³		Gremlin-1¹
Down-Regulated In Mos-Transformed Cells Protein² ³		Increased In High Glucose-2²
Increased In High Glucose Protein 2² ³		Proliferation-Inducing Gene 2²
IHG-2² ³

External Ids:	HGNC: 2001¹	Entrez Gene: 26585²	Ensembl: ENSG00000166923⁷	OMIM: 603054⁵	UniProtKB: O60565³

Export aliases for GREM1 gene to outside databases

Previous GC identifers: GC15P030728 GC15P030797 GC15P009872

(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 GREM1:

This gene encodes a member of the BMP (bone morphogenic protein) antagonist family. Like BMPs, BMP antagonists contain

cystine knots and typically form homo- and heterodimers. The CAN (cerberus and dan) subfamily of BMP antagonists, to

which this gene belongs, is characterized by a C-terminal cystine knot with an eight-membered ring. The antagonistic

effect of the secreted glycosylated protein encoded by this gene is likely due to its direct binding to BMP proteins.

As an antagonist of BMP, this gene may play a role in regulating organogenesis, body patterning, and tissue

differentiation. In mouse, this protein has been shown to relay the sonic hedgehog (SHH) signal from the polarizing

region to the apical ectodermal ridge during limb bud outgrowth. Alternatively spliced transcript variants encoding

different isoforms have been found for this gene. (provided by RefSeq, Jul 2010)

UniProtKB/Swiss-Prot: GREM1_HUMAN, O60565

Function: Cytokine that may play an important role during carcinogenesis and metanephric kidney organogenesis, as a BMP

antagonist required for early limb outgrowth and patterning in maintaining the FGF4-SHH feedback loop. Down-regulates

the BMP4 signaling in a dose-dependent manner. Acts as inhibitor of monocyte chemotaxis (By similarity)

(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_000015.9 NC_018926.1 NT_010194.17

Regulatory elements:

SABiosciences Regulatory transcription factor binding sites in the GREM1 gene promoter:
GATA-3 p53 AP-2alpha isoform 3 AP-2alpha isoform 2 STAT3 AP-2alpha isoform 4 AP-2alpha ATF AP-2alphaA
Other transcription factors

SwitchGear Promoter luciferase reporter plasmid: GREM1 promoter sequence

Search SABiosciences Chromatin IP Primers for GREM1

Epigenetics:

QIAGEN PyroMark CpG Assay predesigned Pyrosequencing DNA Methylation assays in human, mouse, rat GREM1

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

Entrez Gene cytogenetic band: 15q13.3 Ensembl cytogenetic band: 15q13.3 HGNC cytogenetic band: 15q13.3

GREM1 Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
GREM1 gene location

GeneLoc information about chromosome 15 GeneLoc Exon Structure

GeneLoc location for GC15P033010: view genomic region (about GC identifiers)

Start:	33,010,175 bp from pter	End:	33,026,870 bp from pter
Size:	16,696 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: GREM1_HUMAN, O60565 (See protein sequence)

Recommended Name: Gremlin-1 precursor

Size: 184 amino acids; 20697 Da

Subunit: Interacts with SLIT1 and SLIT2 in a glycosylation-dependent manner (By similarity)

Subcellular location: Secreted (Probable)

Secondary accessions: Q52LV3 Q8N914 Q8N936

Alternative splicing: 2 isoforms: O60565-1 O60565-2 (No experimental confirmation available)

Explore the universe of human proteins at neXtProt for GREM1: NX_O60565

Post-translational modifications:

View modification sites using PhosphoSitePlus²

View neXtProt modification sites for NX_O60565

GREM1 Protein expression data from MOPED and PaxDb: About this image

REFSEQ proteins (3 alternative transcripts):

NP_001178251.1 NP_001178252.1 NP_037504.1

ENSEMBL proteins:

ENSP00000300177 ENSP00000453387 ENSP00000453141 ENSP00000323101

Human Recombinant Protein Products:

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	Novus Biologicals GREM1 Proteins Novus Biologicals GREM1 Lysate
	Browse Sino Biological Recombinant Proteins
	Browse ProSpec Recombinant Proteins
	Uscn Proteins for GREM1

Gene Ontology (GO): 2 cellular component terms (GO ID links to tree view): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005615	extracellular space	NAS	9660951
GO:0009986	cell surface	IEA	--

GREM1 for ontologies About GeneDecksing

GREM1 Antibody Products:

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	Browse Enzo Life Sciences for kits & assays
	Uscn ELISAs and CLIAs for GREM1

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

GREM1 for domains About GeneDecksing

3 InterPro domains/families:

IPR006207 Cys_knot_C

IPR004133 DAN

IPR017159 Gremlin_precursor

Graphical View of Domain Structure for InterPro Entry O60565

ProtoNet protein and cluster: O60565

1 Blocks protein family: IPB006207 Cystine knot

UniProtKB/Swiss-Prot: GREM1_HUMAN, O60565

Similarity: Belongs to the DAN family

Similarity: Contains 1 CTCK (C-terminal cystine knot-like) 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: GREM1_HUMAN, O60565

Function: Cytokine that may play an important role during carcinogenesis and metanephric kidney organogenesis, as a BMP

antagonist required for early limb outgrowth and patterning in maintaining the FGF4-SHH feedback loop. Down-regulates

the BMP4 signaling in a dose-dependent manner. Acts as inhibitor of monocyte chemotaxis (By similarity)

Induction: By high glucose through TGFB1-mediated pathways in mesangial cell. Down-regulated in tumor cell lines

Summary:

During embryonic development, GREM1 as signaling molecule is secreted from the following cells: Loose Mesenchyme Cells in Metanephric Mesenchyme, Metanephric Mesenchyme Cells in Metanephric Mesenchyme

It affects the following cells: Ureteric Bud Tip Cells in Ureteric Bud

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		QIAGEN Custom miScript Target Protector blocks miRNA-binding site of human, mouse, rat GREM1
		8/45 QIAGEN miScript miRNA Assays for microRNAs that regulate GREM1 (see all 45):
		hsa-miR-323-3p hsa-miR-642a hsa-miR-199a-3p hsa-miR-128 hsa-miR-218 hsa-miR-23a hsa-miR-3174 hsa-miR-181b

		Browse SwitchGear 3'UTR luciferase reporter plasmids

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 GREM1 (see all 7) OriGene shRNA RFP: GREM1 OriGene siRNA: GREM1
		QIAGEN FlexiTube/FlexiPlate siRNA for gene silencing in human, mouse, rat GREM1

Gene Editing
Products:

DNA2.0 Custom Protein Engineering Service for GREM1

Clone Products:		Browse Clones for the Expression of Recombinant Proteins Available from EMD Millipore
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		GenScript: all cDNA clones in your preferred vector (see all 3): GREM1 (NM_013372)
		Sino Biological Human cDNA Clone for GREM1
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		Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat GREM1

Cell Line Products:		GenScript Custom overexpressing Cell Line Services for GREM1
		LifeMap BioReagents: cell line associated with GREM1: PureStem Z11, Meso Progenitor

In Situ Assay
Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for GREM1

Gene Ontology (GO): 5/7 molecular function terms (GO ID links to tree view) (see all 7): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0005125	cytokine activity	IEA	--
GO:0005515	protein binding	IPI	16545136
GO:0016015	morphogen activity	ISS	9660951
GO:0030296	protein tyrosine kinase activator activity	IEA	--
GO:0036122	BMP binding	ISS	9660951

GREM1 for ontologies About GeneDecksing

Animal Models:
11 MGI mutant phenotypes (inferred from 5 alleles) (MGI details for Grem1):

cellular	craniofacial	embryogenesis	homeostasis/metabolism	limbs/digits/tail
mortality/aging	nervous system	renal/urinary system	reproductive system	respiratory system
skeleton

GREM1 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 About this table

See pathways by source

Super-pathway

contained gene-specific pathways

TGF-beta/Smad Signaling

TGF-beta/Smad Signaling

1.00

Angiogenesis

Angiogenesis

1.00

Signaling by BMP

BMP receptor signaling

0.31

Pathway sources
See GeneCards unified pathways
Show all pathways

2 Cell Signaling Technology (CST) Pathways for GREM1

	Angiogenesis
	TGF-beta/Smad Signaling

1 BioSystems Pathway for GREM1

BMP receptor signaling

GREM1 for pathways About GeneDecksing

Interactions:

Search SABiosciences Gene Network Central^TM Interacting Genes and Proteins Networks for GREM1

STRING Interaction Network Preview (showing 5 interactants - click image to see 10)

5/10 Interacting proteins for GREM1 (O60565^{1, 2, 3} ENSP00000300177⁴) via UniProtKB, MINT, STRING, and/or I2D (see all 10)

Interactant		Interaction Details
GeneCard	External ID(s)	Interaction Details
YWHAH	Q04917¹^,²^,³, ENSP00000248975⁴	EBI-944395,EBI-306940 MINT-8007394 I2D: score=4 STRING: ENSP00000248975
BMP2	P12643³, ENSP00000368104⁴	I2D: score=1 STRING: ENSP00000368104
BAMBI	ENSP00000364683⁴	STRING: ENSP00000364683
BMP4	ENSP00000245451⁴	STRING: ENSP00000245451
BMPR1A	ENSP00000224764⁴	STRING: ENSP00000224764

About this table

Gene Ontology (GO): 5/46 biological process terms (GO ID links to tree view) (see all 46): About this table

GO ID	Qualified GO term	Evidence	PubMed IDs
GO:0000902	cell morphogenesis	IDA	16545136
GO:0001525	angiogenesis	--	--
GO:0001658	branching involved in ureteric bud morphogenesis	IEA	--
GO:0002042	cell migration involved in sprouting angiogenesis	ISS	9660951
GO:0002092	positive regulation of receptor internalization	ISS	9660951

GREM1 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

GREM1 for compounds About GeneDecksing

Browse Small Molecules at EMD Millipore

Browse drugs & compounds from Enzo Life Sciences

Browse Tocris compounds for GREM1
5 Novoseek chemical compound relationships for GREM1 gene About this table

Compound	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
cystine	52.9	2	11179006 (1), 11984879 (1)
cysteine	27.5	8	15254711 (2), 15806314 (1), 16816361 (1), 19715619 (1) (see all 5)
dexamethasone	4.49	2	15268896 (1)
glucose	0	11	10744662 (3), 12386293 (1), 15957132 (1)
tyrosine	0	2	17077323 (1), 16545136 (1)

Search CenterWatch for drugs/clinical trials and news about GREM1

(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 GREM1 gene (3 alternative transcripts):

NM_001191322.1 NM_001191323.1 NM_013372.6

Unigene Cluster for GREM1:

Gremlin 1

Hs.40098 [show with all ESTs]

Unigene Representative Sequence: AY232290

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

ENST00000300177(uc010uby.2 uc001zhd.2 uc001zhe.2) ENST00000560677

ENST00000560830 ENST00000322805

miRNA Products:		OriGene 3'-UTR Clone: GREM1 Browse OriGene MicroRNA Expression Plasmids
		QIAGEN Custom miScript Target Protector blocks miRNA-binding site of human, mouse, rat GREM1
		8/45 QIAGEN miScript miRNA Assays for microRNAs that regulate GREM1 (see all 45):
		hsa-miR-323-3p hsa-miR-642a hsa-miR-199a-3p hsa-miR-128 hsa-miR-218 hsa-miR-23a hsa-miR-3174 hsa-miR-181b

		Browse SwitchGear 3'UTR luciferase reporter plasmids

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 GREM1 (see all 7) OriGene shRNA RFP: GREM1 OriGene siRNA: GREM1
		QIAGEN FlexiTube/FlexiPlate siRNA for gene silencing in human, mouse, rat GREM1

Clone Products:		OriGene Myc/DDK tagged cDNA clones in CMV expression vector in human, mouse, rat for GREM1 (see all 5) OriGene untagged cDNA clones in CMV expression vector in human, mouse, rat for GREM1 OriGene custom cloning services � gene synthesis, subcloning, mutagenesis, variant library, vector shuttling
		GenScript: all cDNA clones in your preferred vector (see all 3): GREM1 (NM_013372)
		DNA2.0 Custom Codon Optimized Gene Synthesis Service for GREM1
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Primer Products:		OriGene genome-wide validated SYBR primer pairs in human, mouse, rat for GREM1 Browse OriGene validated miRNA SYBR primer pairs
		SABiosciences RT² qPCR Primer Assay in human, mouse, rat GREM1
		QIAGEN QuantiTect SYBR Green Assays in human, mouse, rat GREM1
		QIAGEN QuantiFast Probe-based Assays in human, mouse, rat GREM1

Additional cDNA sequence:

AF045800.1 AF110137.2 AF154054.1 AK095741.1 AK095890.1 AK095972.1 AK312265.1 AL359059.1

AL359060.1 AY232290.1 BC069525.1 BC093778.1 BC101611.1

9 DOTS entries:

DT.455758 DT.101971550 DT.95071440 DT.121024008 DT.102832129 DT.100025856 DT.100789439 DT.121024016

DT.70100915

24/279 AceView cDNA sequences (see all 279):

AL359060 AA373673 BM994968 BF431876 BP377797 BQ010116 BM995363 AW068940

AA373975 AA788723 AA670062 AI753469 AI753593 AW069165 BU180797 AI037867

AI521673 CB160772 BQ009519 AI309543 AW069006 CB161671 AI754354 AI753621

GeneLoc Exon Structure

3 Alternative Splicing Database (ASD) splice patterns (SP) for GREM1 About this scheme

ExUns:	1a	·	1b	^	2a	·	2b	^	3a	·	3b	·	3c	·	3d
SP1:							-
SP2:									-
SP3:

ECgene alternative splicing isoforms for GREM1

(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

GREM1 expression in normal human tissues (normalized intensities)
See probesets specificity/sensitivity at GeneAnnot
About this imageBioGPS
CGAP TAG: AATCTGTAAC

About this image

GREM1 expression in embryonic tissues and stem cells

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

8 LifeMap In Vivo Development Anatomical Compartments/Cells

Tissue

Anatomical Compartment

Cell

Category (developmental path)

Kidney

Metanephric Mesenchyme

Loose Mesenchyme Cells

Kidney

Metanephric Mesenchyme

Metanephric Mesenchyme Cells

Kidney

Limb

Autopod

Prechondrocytic Mesenchymal Cells

Bone, Cartilage

Limb

Limb Bud

Limb Bud Mesenchyme Cells

Mesoderm

Bone, Cartilage

Prechondrocytic Mesenchymal Cells

Bone, Cartilage

Limb

Zeugopod

Prechondrocytic Mesenchymal Cells

Bone, Cartilage

Cartilage

Meckel's Cartilage

Cartilage

Expression:

Positive

Negative

Selective marker

Experimental details:

Curated

Microarrays

In-situ hybridization

Stem Cell Differentiation: 10/14 LifeMap Cells (see all 14)

Name

Category

PureStem™ myogenic progenitor Z11 (Embryonic Progenitor Cell)

Heart, Myocardium, Smooth Muscle

PureStem™ progenitor E68 (Embryonic Progenitor Cell)

PureStem™ mesenchymal Progenitor SK11 (Embryonic Progenitor Cell)

Adipose, Bone, Cartilage

PureStem™ mesenchymal progenitor MEL2 (Embryonic Progenitor Cell)

Adipose, Bone, Cartilage

PureStem™ mesenchymal progenitor 7SMOO32 (Embryonic Progenitor Cell)

Adipose, Bone, Cartilage

PureStem™ mesenchymal progenitor 4D20.8 (Embryonic Progenitor Cell)

Lateral Plate Mesoderm, Neural Crest

PureStem™ progenitor E44 (Embryonic Progenitor Cell)

PureStem™ progenitor EN2 (Embryonic Progenitor Cell)

PureStem™ progenitor F15 (Embryonic Progenitor Cell)

PureStem™ progenitor SM22 (Embryonic Progenitor Cell)

See GREM1 Protein Expression from SPIRE MOPED and PaxDB
Genevestigator expression for GREM1

SOURCE GeneReport for Unigene cluster: Hs.40098

UniProtKB/Swiss-Prot: GREM1_HUMAN, O60565

Tissue specificity: Highly expressed in small intestine, fetal brain and colon. Expression is restricted to intestinal

subepithelial myofibroblasts (ISEMFs) at the crypt base. In subjects with HMPS1, by contrast, GREM1 is expressed, not

only in basal ISEMFs, but also at very high levels in epithelial cells (predominantly colonocytes), with expression

extending most of the way up the sides of the crypt. Weakly expressed in brain, ovary, prostate, pancreas and skeletal

muscle. In brain found in the region localized around the internal capsule in the large subcortical nuclei, including

caudate, putamen, substantia nigra, thalamus and subthalamus. Predominantly expressed in normal cells including

neurons, astrocytes and fibroblasts

SABiosciences Expression via Pathway-Focused PCR Arrays including GREM1:

Lung Cancer in human mouse rat

Fibrosis in human mouse rat

Hedgehog Signaling Pathway in human mouse rat

Common Cytokines in human mouse rat

Inflammatory Response & Autoimmunity 384HT in human mouse rat

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		QIAGEN QuantiFast Probe-based Assays in human, mouse, rat GREM1

In Situ
Assay Products:

Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for GREM1

(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 GREM1 gene from 7/15 species (see all 15) About this table

Organism	Taxonomic classification	Gene	Description	Human Similarity	Orthology Type	Details
mouse (Mus musculus)	Mammalia	Grem1^{1 , 5}	gremlin 1^{1, 5}	89.31(n)¹ 97.83(a)¹		2 (57.43 cM)⁵ 23892¹ NM_011824.4¹ NP_035954.1¹ 113746164⁵
chicken (Gallus gallus)	Aves	GREM1¹	gremlin 1	78.02(n) 84.62(a)		395826 NM_204978.1 NP_990309.1
lizard (Anolis carolinensis)	Reptilia	-- --	--	78(a) 46(a)	many → 1 many → 1	1(32878860-32879405) 4(144540986-144541540)
African clawed frog (Xenopus laevis)	Amphibia	Cktsf1b1²	gremlin	75.47(n)		AF045798.1
zebrafish (Danio rerio)	Actinopterygii	LOC100005728¹	gremlin-1-like	70.88(n) 72.89(a)		100005728 XM_001344664.2 XP_001344700.2
mosquito (Anopheles gambiae)	Insecta	AgaP_AGAP007994¹	AGAP007994-PA	64.31(n) 61.95(a)		3291626 XM_555475.3 XP_555475.3
worm (Caenorhabditis elegans)	Secernentea	F35B12.10⁶	--	12(a)	1 → many	V(11625952-11629304)

ENSEMBL Gene Tree for GREM1 (if available)
TreeFam Gene Tree for GREM1 (if available)

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

Paralogs for GREM1 gene

NBL1² GREM2²

2 SIMAP similar genes for GREM1 using alignment to 4 protein entries: GREM1_HUMAN (see all proteins):

GREM2 NBL1

GREM1 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)
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SNP ID	Valid	Clinical significance	Chr 15 pos	Sequence	#	AA Chg	Type	More	#	Allele freq	Pop	Total sample	More
Genomic Data					Transcription Related Data				Allele Frequencies
				--	--	--		--	--
rs77865017^1,2		--	33008231(+)	`ACAGAG/TAGGGG`	3	--	us2k¹		1		EA	120
rs138291687^1,2		--	33008262(+)	`GTGGGA/GGCTCT`	3	--	us2k¹		0	--	--	--	--
rs143874276^1,2		--	33008348(+)	`ACAGTG/TTCAGG`	3	--	us2k¹		0	--	--	--	--
rs11633862^1,2	C,F,A,H,	--	33008359(+)	`GCTCCC/AAAAAA`	3	--	us2k¹ trp³		3		NA	8
rs66751912^1,2	C	--	33008359(-)	`TTTTT-/TGGAGC`	3	--	us2k¹		1		NA	2
rs36079334^1,2	C,	--	33008407(+)	`TTGTAT/-TTTGA`	3	--	us2k¹		1		NA	2
rs66623203^1,2	C	--	33008409(+)	`TATTTT/-GATTT`	3	--	us2k¹		1		NA	2
rs73376923^1,2	C,	--	33008596(+)	`GGTCAT/CCGCCT`	3	--	us2k¹		2		WA	120
rs147247707^1,2		--	33008682(+)	`ATGCTC/TCCATT`	3	--	us2k¹		0	--	--	--	--
rs3812934^1,2	C,F,A,H,	--	33008870(-)	`GAAATA/GGATAT`	3	--	us2k¹ tfbs³		21		NS EA NA WA CSA	2346

HapMap Linkage Disequilibrium report for GREM1 (33010175 - 33026870 bp)
Structural Variations (Copy Number Variations, Insertions/Deletions, Inversions)
Database of Genomic Variants (DGV): 3 variations for GREM1
3 CNVs: 0185 7058 5324
Human Gene Mutation Database (HGMD): GREM1

SABiosciences Cancer Mutation PCR Assays

QIAGEN SeqTarget long-range PCR primers in human, mouse, rat for resequencing GREM1

DNA2.0 Custom Variant and Variant Library Synthesis for GREM1

(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.)
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GREM1 for disorders About GeneDecksing

OMIM gene information: 603054 OMIM disorders: --

UniProtKB/Swiss-Prot: GREM1_HUMAN, O60565

Defects in GREM1 are the cause of polyposis syndrome, mixed hereditary type 1 (HMPS1) [MIM:601228]. A disease

characterized by apparent autosomal dominant inheritance of multiple types of colorectal polyp, with colorectal

carcinoma occurring in a high proportion of affected individuals. Patients can develop polyps of multiple and mixed

morphologies including serrated lesions, Peutz-Jeghers polyps, juvenile polyps, conventional adenomas and colorectal

carcinoma in the absence of any identifiable extra-colonic features. Note=HMPS1 is caused by a duplication spanning

the 3' end of the SCG5 gene and a region upstream of the GREM1 locus. This duplication is associated with increased

allele-specific GREM1 expression that may cause reduced bone morphogenetic protein (BMP) pathway activity. This

mechanism also underlies tumorigenesis in juvenile polyposis of the large bowel (PubMed:22561515)

20/35 diseases for GREM1 (see all 35): About MalaCards

asphyxiating thoracic dystrophy radioulnar synostosis clear cell renal cell carcinoma idiopathic interstitial pneumonia

proliferative vitreoretinopathy synostosis idiopathic pulmonary fibrosis norrie disease

cleft lip renal cell carcinoma cleft palate vitreoretinopathy

pulmonary fibrosis hearing loss renal hypoplasia malignant mesothelioma

pulmonary sarcoidosis sarcoidosis glaucoma nephropathy

6 Novoseek disease relationships for GREM1 gene About this table

Disease	-log (P-Val)	Hits	PubMed IDs for Articles with Shared Sentences (# sentences)
diabetic nephropathies	61.5	32	15957132 (4), 17980714 (4), 20150533 (3), 19303394 (3) (see all 11)
renal disease	28.6	5	12885271 (2), 16582577 (1), 17980714 (1), 18455002 (1)
gastric cancer	17	2	19121317 (1)
tumors	0	7	9660951 (1), 19121317 (1), 17077323 (1), 15334466 (1) (see all 6)
carcinoma	0	3	16545136 (2), 17003113 (1)
cancer	0	4	16545136 (2), 16234815 (1), 17003113 (1)

Human Genome Epidemiology (HuGE) Navigator: GREM1 (12 documents)

Export disorders for GREM1 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)
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PubMed articles for GREM1 gene, integrated from 9 sources (see all 120):
(articles sorted by number of sources associating them with GREM1)

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

The Xenopus dorsalizing factor Gremlin identifies a novel family of secreted proteins that antagonize BMP activities. (PubMed id 9660951)^{1, 2, 3, 9} Hsu D.R.... Harland R.M. (1998)
IHG-2, a mesangial cell gene induced by high glucose, is human gremlin. Regulation by extracellular glucose concentration, cyclic mechanical strain, and transforming growth factor-beta1. (PubMed id 10744662)^{1, 2, 9} McMahon R.... Brady H.R. (2000)
DRM/GREMLIN (CKTSF1B1) maps to human chromosome 15 and is highly expressed in adult and fetal brain. (PubMed id 10894942)^{1, 2, 9} Topol L.Z.... Blair D.G. (2000)
Complete sequencing and characterization of 21,243 full-length human cDNAs. (PubMed id 14702039)^{1, 2} Ota T.... Sugano S. (2004)
Signal peptide prediction based on analysis of experimentally verified cleavage sites. (PubMed id 15340161)^{1, 2} Zhang Z. and Henzel W.J. (2004)
The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). (PubMed id 15489334)^{1, 2} Gerhard D.S....Malek J. (2004)
Suppression subtractive hybridization identifies high glucose levels as a stimulus for expression of connective tissue growth factor and other genes in human mesangial cells. (PubMed id 10026205)^{1, 3} Murphy M....Brady H.R. (1999)
Elevated levels of gremlin-1 in eutopic endometrium and peripheral serum in patients with endometriosis. (PubMed id 18314105)^{1, 9} Sha G....Lang J. (2009)
Expression of gremlin, a bone morphogenetic protein antagonist, in glomerular crescents of pauci-immune glomerulonephritis. (PubMed id 17403698)^{1, 9} Mezzano S....Egido J. (2007)
Evaluation of gremlin 1 (GREM1) as a candidate suscep tibility gene for albuminuria-related traits in Mexican Americans with type 2 d iabetes mellitus. (PubMed id 19577778)^{1, 9} Thameem F....Abboud H.E. (2009)

(in PubMed, OMIM, and NCBI Bookshelf)
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		AND	OR
Aliases
Disorders
Free Text

Query String		PubMed OMIM NCBI Bookshelf
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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: 26585	HGNC: 2001	AceView: GREM1	Ensembl:ENSG00000166923	euGenes: HUgn26585
ECgene: GREM1	H-InvDB: GREM1

(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 GREM1	Pharmacogenomics, SNPs, Pathways

Intellectual Property for GREM1 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 GREM1 gene:
Search GeneIP for patents involving GREM1

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

(Antibodies, recombinant proteins, and assays from EMD Millipore, R&D Systems, OriGene, QIAGEN, GenScript, Cell Signaling Technology, SABiosciences, Novus Biologicals, Sino Biological, Enzo Life Sciences, Abcam, ProSpec, Uscn, Thermo Fisher Scientific, Gene Editing from DNA2.0, Clones from EMD Millipore, OriGene, GenScript, Sino Biological, DNA2.0, SwitchGear Genomics, Vector BioLabs, Cell lines from GenScript and LifeMap BioReagents, PCR Arrays from SABiosciences, Drugs and/or compounds from EMD Millipore, Tocris Bioscience, and/or Enzo Life Sciences),
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GREM1 Gene

gremlin 1, DAN family BMP antagonist

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GREM1 (Gene Symbol)
Gremlin 1, DAN Family BMP Antagonist
CKTSF1B1
Cysteine Knot Superfamily 1, BMP Antagonist 1
DAND2
DRM
Gremlin 1
Cell Proliferation-Inducing Gene 2 Protein
Down-Regulated In Mos-Transformed Cells Protein
Increased In High Glucose Protein 2
IHG-2
DAN Domain Family Member 2
Gremlin
Gremlin 1, Cysteine Knot Superfamily, Homolog (Xenopus Laevis)
GREMLIN
Gremlin 1, Cysteine Knot Superfamily, Homolog
Gremlin 1-Like Protein
Gremlin-1
Increased In High Glucose-2
Proliferation-Inducing Gene 2

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GREM1 Gene

gremlin 1, DAN family BMP antagonist

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