CTGF (connective tissue growth factor)

Identity

Other names	CCN2
	Fisp12
	betaIG-M2
	Hcs24
	HBGF-0.8
	ecogenin
	IGFBP8
	IGFBP-rP3
HGNC	CTGF
Location	6q23.1
Location_base_pair	Starts at 132311010 and ends at 132314211 bp from pter ( according to hg18-Mar_2006).

Note

CCN family protein 2/connective tissue growth factor (CCN2/CTGF) was initially identified in the culture supernatant of vascular endothelial cells. Subsequently, the CCN2/CTGF gene has been classified as a representative member of the CCN gene family that is an acronym of the original names of the 3 early members, Cyr61, CTGF and NOV. The mammalian genome contains 6 functional CCN family members, which were renamed based on the proposal of a unified nomenclature. CCN2/CTGF displays multiple functions via interaction with a variety of other molecules. It is important to note that CCN2/CTGF induces the development and regeneration of mesenchymal tissues including bone, cartilage and blood vessels.

DNA/RNA

	Structure of human CCN2/CTGF and its mRNA. Abbreviations S, I, V, T, and C denote the coding regions for the signal peptide, IGFBP module, VWC module, TSP1 module and CT module, respectively.
Description	Unlike the other CCN family members, the CCN2/CTGF gene is conserved among all of the vertebrates and several invertebrates. In Drosophila melanogaster, only one gene is designated as ccn, which is thought to have evolved from a prototypic CCN2/CTGF gene. Therefore, CCN2/CTGF may be regarded evolutionarily as an oldest gene in the CCN family. Consistent with such findings, human CCN2/CTGF gene is also quite compact. The total transcribed sequence is as short as 3.2 kb, and contains 4 intronic sequences of less than 400 bp.
Transcription	The mature mRNA, 2.3 kb in length, is formed by connecting 5 exons encoding a signal peptide, an IGFBP module, a VWC module, a TSP1 module and CT module in the order. The last exon yields a long 3'-untranslated region of more than1.0 kb on the mRNA, which contains the critical elements for post-transcriptional regulation. The proximal promoter area upstream of the transcription initiation site is known to contain several enhancer elements that are critical for transcriptional regulation.

Protein

	Interaction of each module in CCN2/CTGF with other growth factors.
Description	The CCN2/CTGF is composed of 4 distinct modules: the N-terminal signal peptide for secretion; insulin-like growth factor binding protein-like (IGFBP) module, von Willebrand factor type-C repeat (VWC) module, thrombospondin type 1 repeat (TSP1) module and the C-terminal cystine knot (CT) module. This is consistent with other CCN family proteins. Because of this unique structure, the 6 CCN2/CTGF-related proteins are thought to form a distinct protein class which is distinct from the IGFBP family, despite the involvement of IGFBP module. The involvement of cysteine residues that are also conserved among the CCN family members is a prominent structural characteristic. All of the 4 modules are highly interactive with other biomolecules including growth factors, cell-surface receptor molecules and extracellular matrix components. In addition, the tetramodular construction of these modules provides the structural basis for the multiple functionality of CCN2/CTGF, which is described in another section.
Expression	CCN2/CTGF is differentially expressed in certain tissues and organs, particularly in the cardiovascular, gonadal, renal and skeletal systems, during development of vertebrates. The cell population that expresses the CCN2/CTGF gene is highly restricted in each tissue; for example, this factor is produced in cartilage primarily by pre-hypertrophic and hypertrophic chondrocytes, in fact, it was previously referred to as hypertrophic chondrocyte-specific protein 24 (Hcs24).
Localisation	In addition to the tissues containing the cells which express this protein described above, CCN2/CTGF is abundantly present in platelets, although its origin is still unknown.
Function	Under the multiple interactions with specific molecular counterparts, CCN2/CTGF conducts the local information network of extracellular signaling molecules and exerts multiple functions, depending upon the microenvironmental conditions. Indeed, CCN2/CTGF promotes both proliferation and differentiation of mesenchymal stem cells, chondrocytes, osteoblasts, periodontal ligament cells, fibroblasts and vascular endothelial cells in vitro. As a result of these effects, this factor enhances wound healing and tissue regeneration of cartilage and bone.
Homology	CCN2/CTGF is structurally homologous to the other 5 CCN family members. The cysteine residues are highly conserved among the members.

Mutations

Germinal

Until today, no association between mutations in the CCN2/CTGF gene and specific genetic diseases has been described. In mouse models, deletion of both CCN2/CTGF alleles results in severe skeletal malformation, leading to respiratory failure upon delivery. Histological and cell biological analysis revealed that the endochondral ossification process was specifically affected by the CCN2/CTGF deletion, particularly at the final stage that is supported by blood vessel invasion. However, no apparent phenotypic complication has been observed in CCN2/CTGF (-/+) heterozygous mice.

Somatic

At present, nearly 30 SNPs have been described in the NCBI database. Among the SNPs, 35.7% and 39.3% of the total cases have been reported to exist outside of the transcribed area and in the areas of untranslated regions (UTRs) and introns, respectively. Mutation/variation in the open reading frame (ORF) was found in 25% of the total. Missense mutation occurred at a frequency of 42.7 %, whereas the other cases were silent. Excluding the UTRs, the IGFBP-encoding 2nd exon has been a hot spot of mutation and variation (57.1% of the one within ORF). In a single case, an African population was analyzed to compute the actual frequency. This demonstrated that the frequency of a mutation in the 3rd exon, which caused a missense change of amino acid 118 from asparagine to serine by an A to G transition, was 1.3%. Including these cases, no association of a SNP in the CCN2/CTGF gene with any particular human disorder has been described until now.

Implicated in

Entity	Cancers
Oncogenesis	Although the CCN2/CTGF expression is observed in a number of different types of malignant neoplasm, the role of CCN2/CTGF in oncogenesis and its association with malignant phenotypes are quite controversial. According to a previous study, a positive correlation was observed between the level of CCN2/CTGF expression and the degree of malignancy in breast cancer and colorectal cancer cases. These findings are consistent with other recent reports describing the contribution of CCN/CTGF in developing bone metastasis of breast cancers. The ability to promote metastasis can be partially ascribed to the angiogenic activity of CCN2/CTGF. However, over expression of CCN2/CTGF in cells of the same origin is reported to induce apoptosis. Furthermore, in chondrosarcoma cases, patients with higher CCN2/CTGF expression in tumors survived longer than those with lower CCN2/CTGF expression. These findings are consistent with the observation that overexpression of CCN2/CTGF results in benign conversion of the phenotype in oral squamous carcinoma cells and induces cell cycle arrest in fibroblasts. The observations above suggest that CCN2/CTGF produced by tumor cells may exert paracrine angiogenic and autocrine/intracrine anti-proliferative effects in solid tumors.
Entity	Fibrotic disorders.
Disease	Since its initial discovery, CCN2/CTGF has been widely known as a profibrotic factor that is involved in a variety of fibrotic disorders. CCN2/CTGF is associated with systemic sclerosis, keloids, pulmonary fibrosis, diabetic renal fibrosis, liver cirrhosis, pancreatic fibrosis, atherosclerosis, myocardial fibrosis, biliary atresia and cataracts. Since CCN2/CTGF has a positive role in wound healing and mesenchymal tissue regeneration, the fibrotic changes observed in those diseases may be regarded as a result of dysregulated regeneration of corresponding tissues.
Prognosis	Fibrotic changes are usually irreversible; however, antibody-mediated molecular therapeutics against CCN2/CTGF is currently being developed to prevent the development of fibrotic lesions.

External links

	Nomenclature
HGNC	CTGF   2500
Entrez_Gene	CTGF  1490  connective tissue growth factor
	Cards
Atlas	CTGFID40192ch6q23
GeneCards	CTGF
Ensembl	CTGF [Search_View]   ENSG00000118523 [Gene_View]  CTGF [Vega]
Genatlas	CTGF
GeneLynx	CTGF
eGenome	CTGF
euGene	1490
	Genomic and cartography
GoldenPath	CTGF  -  6q23.1   chr6:132311010-132314211 -  6q23.2   [Description]    (hg18-Mar_2006)
Ensembl	CTGF - 6q23.2 [CytoView]
NCBI	Mapview
OMIM	Disease map [OMIM]
HomoloGene	CTGF
	Gene and transcription
Genbank	AF127918 [ ENTREZ ]
Genbank	AK092280 [ ENTREZ ]
Genbank	AK125220 [ ENTREZ ]
Genbank	AK290884 [ ENTREZ ]
Genbank	AK315207 [ ENTREZ ]
RefSeq	NM_001901 [ SRS ]    NM_001901 [ ENTREZ ]
RefSeq	AC_000049 [ SRS ]    AC_000049 [ ENTREZ ]
RefSeq	AC_000138 [ SRS ]    AC_000138 [ ENTREZ ]
RefSeq	NC_000006 [ SRS ]    NC_000006 [ ENTREZ ]
RefSeq	NT_025741 [ SRS ]    NT_025741 [ ENTREZ ]
RefSeq	NW_001838990 [ SRS ]    NW_001838990 [ ENTREZ ]
RefSeq	NW_923184 [ SRS ]    NW_923184 [ ENTREZ ]
CCDS	CTGF CCDS - NCBI
AceView	CTGF AceView - NCBI
Unigene	Hs.591346 [ SRS ]    Hs.591346 [ NCBI ]     HS591346 [ spliceNest ]
Fast-db	16513 (alternative variants)
	Protein : pattern, domain, 3D structure
SwissProt	P29279 [ SRS]    P29279 [ EXPASY ]     P29279 [ INTERPRO ]     P29279 [ UNIPROT ] P29279 [ VarSplice ]
Prosite	PS01185 CTCK_1 [ SRS ]    PS01185 CTCK_1 [ Expasy ]
Prosite	PS01225 CTCK_2 [ SRS ]    PS01225 CTCK_2 [ Expasy ]
Prosite	PS00222 IGFBP_N_1 [ SRS ]    PS00222 IGFBP_N_1 [ Expasy ]
Prosite	PS51323 IGFBP_N_2 [ SRS ]    PS51323 IGFBP_N_2 [ Expasy ]
Prosite	PS50092 TSP1 [ SRS ]    PS50092 TSP1 [ Expasy ]
Prosite	PS01208 VWFC_1 [ SRS ]    PS01208 VWFC_1 [ Expasy ]
Prosite	PS50184 VWFC_2 [ SRS ]    PS50184 VWFC_2 [ Expasy ]
Interpro	IPR006208 Cys_knot [ SRS ]    IPR006208 Cys_knot [ EBI ]
Interpro	IPR006207 Cys_knot_C [ SRS ]    IPR006207 Cys_knot_C [ EBI ]
Interpro	IPR012395 IGFBP_CNN [ SRS ]    IPR012395 IGFBP_CNN [ EBI ]
Interpro	IPR000867 IGFBP_like [ SRS ]    IPR000867 IGFBP_like [ EBI ]
Interpro	IPR000884 TSP1 [ SRS ]    IPR000884 TSP1 [ EBI ]
Interpro	IPR001007 VWF_C [ SRS ]    IPR001007 VWF_C [ EBI ]
CluSTr	P29279
Pfam	PF00007 Cys_knot [ SRS ]    PF00007 Cys_knot [ Sanger ]    pfam00007 [ NCBI-CDD ]
Pfam	PF00219 IGFBP [ SRS ]    PF00219 IGFBP [ Sanger ]    pfam00219 [ NCBI-CDD ]
Pfam	PF00090 TSP_1 [ SRS ]    PF00090 TSP_1 [ Sanger ]    pfam00090 [ NCBI-CDD ]
Pfam	PF00093 VWC [ SRS ]    PF00093 VWC [ Sanger ]    pfam00093 [ NCBI-CDD ]
Smart	SM00041 CT [EMBL]
Smart	SM00121 IB [EMBL]
Smart	SM00209 TSP1 [EMBL]
Smart	SM00214 VWC [EMBL]
Blocks	P29279
HPRD	00412
	Protein Interaction databases
DIP	P29279
IntAct	P29279
	Polymorphism : SNP, mutations, diseases
OMIM	121009    [ map ]
GENECLINICS	121009
SNP	CTGF [dbSNP-NCBI]
SNP	NM_001901 [SNP-NCI]
SNP	CTGF [GeneSNPs - Utah]  CTGF] [HGBASE - SRS]
HAPMAP	CTGF [HAPMAP]
HGMD	CTGF
Genetic Association	CTGF
CDC HuGE	CTGF
	General knowledge
Family Browser	CTGF [UCSC Family Browser]
SOURCE	NM_001901
SMD	Hs.591346
SAGE	Hs.591346
GO	cartilage condensation [Amigo]  cartilage condensation
GO	ossification [Amigo]  ossification
GO	angiogenesis [Amigo]  angiogenesis
GO	regulation of cell growth [Amigo]  regulation of cell growth
GO	integrin binding [Amigo]  integrin binding
GO	protein binding [Amigo]  protein binding
GO	insulin-like growth factor binding [Amigo]  insulin-like growth factor binding
GO	extracellular region [Amigo]  extracellular region
GO	proteinaceous extracellular matrix [Amigo]  proteinaceous extracellular matrix
GO	plasma membrane [Amigo]  plasma membrane
GO	DNA replication [Amigo]  DNA replication
GO	cell adhesion [Amigo]  cell adhesion
GO	cell-matrix adhesion [Amigo]  cell-matrix adhesion
GO	integrin-mediated signaling pathway [Amigo]  integrin-mediated signaling pathway
GO	intracellular signaling cascade [Amigo]  intracellular signaling cascade
GO	growth factor activity [Amigo]  growth factor activity
GO	heparin binding [Amigo]  heparin binding
GO	fibroblast growth factor receptor signaling pathway [Amigo]  fibroblast growth factor receptor signaling pathway
GO	epidermis development [Amigo]  epidermis development
GO	response to wounding [Amigo]  response to wounding
GO	cell migration [Amigo]  cell migration
GO	cell differentiation [Amigo]  cell differentiation
PubGene	CTGF
TreeFam	CTGF
CTD	1490 [Comparative ToxicoGenomics Database]
	Other databases
	Probes
Probe	CTGF Related clones (RZPD - Berlin)
	PubMed
PubMed	146 Pubmed reference(s) in Entrez

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A novel cis-element that enhances connective tissue growth factor gene expression in chondrocytic cells.

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Suppressive effect of overexpressed connective tissue growth factor on tumor cell growth in a human oral squamous cell carcinoma-derived cell line.

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Role of CTGF/HCS24/ecogenin in skeletal growth control.

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Pathogenic role of connective tissue growth factor (CTGF/CCN2) in osteolytic metastasis of breast cancer.

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Role of CCN2/CTGF/Hcs24 in bone growth.

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Contributor(s)

Written	07-2007	Satoshi Kubota, Masaharu Takigawa
		Department of Biochemistry and Molecular Dentistry Okayama University Graduate School of Medicine, Dentistry and Pharmaceutical Sciences Dean, Okayama University Dental School 2-5-1, Shikata-cho, Okayama, 700-8525, Japan

Citation

This paper should be referenced as such :

Kubota S, Takigawa M . CTGF (connective tissue growth factor). Atlas Genet Cytogenet Oncol Haematol. July 2007 . URL : http://AtlasGeneticsOncology.org/Genes/CTGFID40192ch6q23.html

© Atlas of Genetics and Cytogenetics in Oncology and Haematology

indexed on : Sun Nov 9 19:39:20 2008