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RAF1 (v-raf-1 murine leukemia viral oncogene homolog 1)

Written2007-03Max Cayo, David Yu Greentblatt, Muthusamy Kunnimalaiyaan, Herbert Chen
Endocrine Surgery Research Laboratories, Department of Surgery, Paul P. Carbone Comprehensive Cancer Center, University of Wisconsin, Madison, Wisconsin, USA

(Note : for Links provided by Atlas : click)

Identity

Alias_namesserine/threonine kinase
Alias_symbol (synonym)Raf-1
c-Raf
CRAF
Other alias
HGNC (Hugo) RAF1
LocusID (NCBI) 5894
Atlas_Id 42032
Location 3p25.2  [Link to chromosome band 3p25]
Location_base_pair Starts at 12583601 and ends at 12664201 bp from pter ( according to hg19-Feb_2009)  [Mapping RAF1.png]
Fusion genes
(updated 2016)		AGGF1 (5q13.3) / RAF1 (3p25.2)CLCN6 (1p36.22) / RAF1 (3p25.2)CMTM8 (3p22.3) / RAF1 (3p25.2)
ESRP1 (8q22.1) / RAF1 (3p25.2)FYCO1 (3p21.31) / RAF1 (3p25.2)HACL1 (3p25.1) / RAF1 (3p25.2)
LMNA (1q22) / RAF1 (3p25.2)MBNL1 (3q25.1) / RAF1 (3p25.2)MPRIP (17p11.2) / RAF1 (3p25.2)
PAPD7 (5p15.31) / RAF1 (3p25.2)QKI (6q26) / RAF1 (3p25.2)RAF1 (3p25.2) / ACTR2 (2p14)
RAF1 (3p25.2) / AGGF1 (5q13.3)RAF1 (3p25.2) / DAZL (3p24.3)RAF1 (3p25.2) / ESRP1 (8q22.1)
RAF1 (3p25.2) / MSS51 (10q22.2)RAF1 (3p25.2) / RSRC2 (12q24.31)RAF1 (3p25.2) / SRGAP3 (3p25.3)
RAF1 (3p25.2) / TMEM40 (3p25.2)RAF1 (3p25.2) / XPC (3p25.1)SRGAP3 (3p25.3) / RAF1 (3p25.2)
TRAK1 (3p22.1) / RAF1 (3p25.2)

DNA/RNA

Note History and Nomenclature:
c-Raf-1 was the first successfully cloned functional human homolog of the v-Raf gene, and thus the gene product of c-Raf-1 has historically been referred to in the literature simply as Raf-1. Subsequently, B-Raf and A-Raf-1 paralogues ( BRAF, located in Xq13 and ARAF, located in Xp11) were discovered. A suitable nomenclature is as follows: A-RAF, B-RAF, and C-RAF for the functional human proteins and A-RAF, B-RAF, and C-RAF for the corresponding genes; a-raf, b-raf, and c-raf for the murine proteins and A-Raf, B-Raf, and C-Raf for the corresponding genes. Raf-1 (or RAF-1) is generally taken to mean C-RAF-1 but could apply to A-RAF-1 equally. Here, RAF-1 will be taken to mean C-RAF-1 (RAF-1 = C-RAF-1, etc.).
Description C-RAF (RAF-1, C-RAF-1) encompasses 80,570 bp of DNA; 17 Exons.
Transcription RAF-1 transcribed mRNA contains 3212-3216 nucleotides.

Protein

Description The RAF proteins share three conserved domains: two (CR1 and CR2) in the N terminus and a third (CR3-encoding for the serine/threonine kinase domain) in the C terminus. The RAF proteins exhibit complex regulation involving numerous phosphorylation sites throughout the proteins. Despite constitutional similarity, the Raf isoforms have been shown to carry out non-redundant functions, implying that they are distinct.
RAF-1 (C-RAF-1): 72-74 kDa.
Note: A-RAF: about 68 kDa.
Note: B-RAF (which undergoes alternate splicing): ranges from 75 to 100 kDa.
Expression C-RAF (RAF-1) and A-RAF mRNA is expressed ubiquitously. A-RAF mRNA is highly expressed in urogenital organs. B-RAF is expressed in a wide range of tissues, but most substantially in neuronal tissues.
Localisation Cytosolic.
Function RAF proteins are part of the conserved MAPK (mitogen-activated protein kinase)/ERK (extracellular signal-regulated kinase) signaling cascade between the cell surface and the nucleus. RAF is regulated by the upstream RAS family of small G proteins. RAS is predominantly located on the inner leaflet of the plasma membrane and is functionally activated by GTP-binding. Binding of various extracellular ligands such as growth factors and hormones activates RAS and subsequently RAF proteins. RAS binds directly to the N-terminal regulatory domain or RAF (the RAS binding domain (RBD)). RAS interacts secondarily with the cysteine-rich domain (CRD) on CR1 of RAF. RAS-RAF binding can be affected by 14-3-3 proteins and other scaffold/adaptor proteins kinase suppressor of RAS (KSR), the multidomain protein connector-enhancer of KSR (CNK), and the leucine-rich-repeat protein suppressor of RAS mutations-8 (SUR8), which cause formation of various homo- and heterodimers and subsequently affect signal transduction. RAF activation leads to activation of the protein kinases MEK1 and MEK2 and subsequently the MAPK proteins ERK1 and ERK2. The downstream effects of MEK1/2-ERK1/2 activation are varied, complex, and depend on the cellular context. Resultant effects include activation of transcription factors involved in tumorigenesis, cell growth, survival, differentiation, metabolism, and cytoskeletal rearrangements. RAF-1 (C-RAF-1), A-RAF, and B-RAF are all capable of activating the MEK1/2-ERK1/2 signaling pathway.
RAF-1 is capable of activating the NF-kB transcription factor through an unknown mechanism that does not seem to involve direct phosphorylation of NF-kB and is independent of MEK1/2-ERK1/2 signaling.
RAF-1 is known to directly affect cell survival through phosphorylation of BAG1 (BCL2-associated athanogene-1), an anti-apoptotic protein that binds to BCL2, a second anti-apoptotic factor, also the prototype for a family of mammalian genes involved in mitochondrial outer membrane permeability (MOMP), thus restoring its function. BCL2 also targets RAF-1 to the mitochondrial membrane, where it is able to more readily phosphorylate substrates. The RAF-1/BAG1/BCL2 interaction allows RAF-1 to phosphorylate the pro-apoptotic protein BAD at the mitochondrial membrane, promoting cell survival.
Other known substrates of RAF-1 include the phosphatase CDC25C, the apoptosis signal-regulating kinase-1 (ASK1), and the tumor-suppressor protein retinoblastoma (Rb).
RAF-1 is tightly regulated by the AKT/PKB pathway through phosphorylation at S259.

Mutations

Somatic It has been widely established that RAF-1 over activity, typically via ras-activating mutations, is central to tumorigenesis and cell proliferation in numerous cancers (about 30% of all human cancers). However, it has come to the fore that oncogenesis may be due to ras/RAF-1 dysregulation (either increased or decreased expression) rather than increases in ras/RAF-1 activity exclusively.

Implicated in

Note
  
Entity Medullary Thyroid Cancer (MTC)
Disease A neuroendocrine tumor derived from parafollicular C cells of the thyroid gland, MTC is the third most common form of thyroid cancer, accounting for 3-5% of all cases. MTC cells secrete hormones and tumor markers such as calcitonin, chromogranin A (CgA), and carcinoembryonic antigen (CEA).
Symptoms are related to either direct invasion or metastasis (neck mass, dyspnea, dysphagia, voice changes, pain) or tumor secretion of bioactive amines and peptides (diarrhea, flushing).
Prognosis Currently, surgery is the only potentially curative therapy for patients with MTC. The recommended operation is total thyroidectomy with lymph node dissection. However, 50% of patients treated with surgery suffer persistent or recurrent disease.
Oncogenesis 20% of patients with medullary thyroid cancer have an autosomal dominant inherited form of the disease, which is the result of well-characterized point mutations in the RET proto-oncogene. RAF-1 is conserved but not expressed at baseline in MTC. Pre-clinical studies have shown that activation of RAF-1 in MTC (TT) cells by means of RAF-1 gene transfection or RAF-1 activating small molecules (ZM336372) results in tumor cell growth inhibition in vitro and in vivo.
  
  
Entity Carcinoid Tumors
Disease Carcinoids are tumors that arise from the diffuse neuroendocrine cell system of the gut, lungs, and other organs. The incidence is 1-5 per 100,000 individuals. Carcinoids frequently metastasize to the liver and are the second most common source of isolated liver metastases. Carcinoids secrete various bioactive hormones such as 5-HT (5-hydroxy tryptophan, also known as serotonin) and chromogranin A.
Prognosis Patients with hepatic metastases suffer debilitating symptoms such as abdominal pain, flushing, bronchoconstriction, and diarrhea. Palliative treatment for these hormone-induced symptoms includes somatostatin analogs (such as octeotride). Conventional anticancer treatments such as chemotherapy and external beam radiation is largely ineffective for carcinoid tumors.
Oncogenesis RAF-1 activation is detrimental to tumorigenesis in carcinoid cells. Marked reduction in neuroendocrine phenotypic markers such as human achaete-scute complex like-1 (ASCL-1) and bioactive hormones 5-HT, chromogranin A, and synaptophysin has been noted upon RAF-1 activation using an estrogen-inducible RAF-1 construct in human GI (BON) and pulmonary carcinoid cell lines (NCI-H727) .
Treatment of GI carcinoid cells with RAF-1 activator ZM336372 led to a decrease in bioactive hormone levels, a suppression of cellular proliferation, an increase in cell cycle inhibitors p21 and p18, as well as a decrease in the neuroendocrine phenotypic marker ASCL-1. ZM336372 treatments also led to progressive phosphorylation (activation) of MEK1/2, ERK1/2, and RAF-1.
  
  
Entity Small Cell Lung Cancer (SCLC)
Disease SCLC tends to present with metastatic and regional spread. Carcinoids rarely metastasize, arise from major bronchi, and express neuron-specific enolase, chromogranin, and synaptophysin. Neuroendocrine carcinoids or atypical carcinoids have a more aggressive course.
Oncogenesis Human small-cell lung cancer (SCLC) cell lines rarely harbor ras-activating mutations. In one cell line of SCLC, DMS53, it was shown that by RAF-1 induction using an estrogen-inducible RAF-1 construct SCLC cells underwent differentiation and G1-specific growth arrest in conjunction with MEK/ERK1/2 pathway activation.
  
  
Entity Non-Small Cell Lung Cancer (NSCLC).
Disease Adenocarcinoma is the most common type of NSCLC accounting for about 40% of cases. Lesions are generally located peripherally and develop systemic metastases despite small primary tumors. 25% of NSCLC are squamous cell carcinomas which often remain localized.
Oncogenesis RAF-1 is over-expressed due to oncogenic ras mutations in about 35% of NSCLC.
The majority of NSCLC exhibits EGFR over-expression leading to upregulation of RAF-1 activity. NSCLC has been shown to be mediated by a TGF-a/EGFR-mediated autocrine loop activated by signaling involving RAF-1 and PI3K-Akt.
  
  
Entity Pheochromocytoma.
Disease Pheochromocytomas are neuroectodermal in origin and arise from the chromaffin cells of the adrenal medulla. 10% of tumors are bilateral. Typical symptoms such as hypertension, headaches, diaphoresis, palpitations, diarrhea, and skin rashes, are related to tumor production of catecholamines, especially in patients with metastases. Pheochromocytoma is potentially fatal, but relatively uncommon (2-8 cases per million people annually). Curative therapy is surgery, usually accomplished by laparoscopic adrenalectomy.
Oncogenesis Activation of MEK1/2-ERK1/2 is necessary for differentiation of pheochromocytoma (PC12) cells and leads to decreased cell proliferation. RAF-1 activation in pheochromocytoma cells using ZM336372 led to cellular differentiation, growth arrest, and a decrease in the neuroendocrine marker chromogranin A.
  
  
Entity Non-Neuroendocrine Cancers with ras-activating Mutations.
Oncogenesis About 30% of all human cancers express ras-activating mutations. More than 85% of pancreatic adenocarcinomas, and 50% of colonic adenocarcinomas harbor K-ras mutations. K-ras is an upstream effector of RAF-1 in the RAF-1/MEK/ERK1/2 signaling pathway. Ras mutations have also been linked to tumorigenesis of cholangiocarcinoma, adenocarcinoma of the lung, squamous cell cancer, gastric adenocarcinoma, small bowel adenocarcinoma, and malignant melanoma.
  
  
Entity Colorectal Cancer.
Oncogenesis RAF-1 is over-activated due to oncogenic ras mutations in about 50% of colon cancers. These mutations are associated with poor prognosis, and are necessary for maintenance of the malignant phenotype.
RAF-1 inhibition in response to interaction with RAF kinase inhibitor protein (RKIP) (up-regulated in conjunction with the nuclear factor kappa B signaling pathway) has been linked with overall and disease-free survival in patients with colorectal cancers. RKIP has been identified as potentially useful for identifying early-stage CRC patients at risk for relapse.
  
  
Entity Pancreatic Carcinoma.
Oncogenesis RAF-1 is overactivated due to oncogenic ras mutations in about 90% of pancreatic carcinomas (Panc-1 and Mia-PaCa2). It has been shown that malignancy of these cells is reduced using k-ras RNAi. Pharmacological inhibition of the RAF/MEK/ERK pathway in pancreatic cancer cell lines (via MEK inhibition) results in reduction in cellular proliferation and an increase in cell cycle arrest.
  
  
Entity Hepatocellular Carcinoma (HCC)
Oncogenesis RAF-1 is over-activated in about 50% of biopsies while the RAF-1 protein is over-expressed in nearly 100% of all HCC's. Angiogenesis and other functions essential to tumorigenesis in HCC have been reported to depend on the RAF/MEK/ERK signaling pathway. RAF-1 inhibitor Sorafenib has been reported (in-vitro and in-vivo) to inhibit RAF-1 activity, leading to decreased MEK/ERK activity, reduced cellular proliferation, and apoptosis in several HCC cell lines including HepG2 and PLC/PRF/5.
  
  
Entity Prostate Cancer.
Oncogenesis RAF kinase inhibitor protein (RKIP) coding mRNAs have been observed to activate interferon-inducible 2',5'-oligoadenylate synthetases (OAS). OAS activity is characteristically increased (via these mRNAs) in prostate cancer cell lines PC3, LNCaP and DU145. RKIP expression is detectable in primary prostate cancer sections but not in metastases. This suggests RKIP's characterization as an anti-metastasis gene using the RAF/MEK/ERK signaling pathway is appropriate.
RAF-1 inhibition using systemically delivered novel cationic cardiolipin liposomes (NeoPhectin-AT) containing a small interfering RNA (siRNA) against RAF-1 causes tumor growth inhibition in a xenograft model of human prostate cancer.
RAF/MEK/ERK signaling pathway activation via a biologically active peptide called a prosaptide (TX14A) stimulates cell proliferation/survival, migration, and invasion in human prostate cancer cells.
NSC 95397 and NSC 672121, cdc25 inhibitors, were shown to activate the RAF/MEK/ERK pathway in prostate cancer cells.
RAF-1 activation in LNCaP prostate cancer cells using an estrogen-inducible construct led to growth inhibition.
  
  
Entity Breast Cancer.
Oncogenesis Growth hormone releasing hormone (GHRH) has been shown to regulate breast cancer cell proliferation and differentiation. In MDA-231 breast cancer cells, exogenous GHRH stimulated dose-dependent proliferation. RAF-1 inhibition using the agent PD98059 caused prevention of MAPK phosphorylation by GHRH as well as reduced cellular proliferation.
Proliferative effects of steroid hormone estradiol on MCF-7 breast cancer cells have been linked with increased expression of RAF-1, possibly due to direct activation of RAF-1 by estradiol.
RAF kinase inhibitor protein (RKIP) is associated with metastasis suppression. RKIP expression is lost in lymph node metastases. This suggests RKIP is a metastasis inhibitor gene and that RAF-1 expression enables metastasis.
The PTK inhibitor AG 879 inhibits proliferation of human breast cancer cells through inhibition of MAP kinase activation through inhibition of expression of the RAF-1 gene.
RAF-1 down-regulation is associated with paclitaxel drug resistance in human breast cancer cell line MCF-7/Adr.
  
  
Entity Renal Cell Carcinoma.
Oncogenesis RAF-1 is overactivated in conjunction with loss of function of the VHL ( von Hippel-Lindau) tumor-suppressor gene.
  
  
Entity Glioma .
Oncogenesis RAF-1 inhibitor AAL881 inhibited growth of glioma cell xenografts.
  
  
Entity Cervical Cancer.
Oncogenesis Low RAF-1 kinase activity is significantly associated with paclitaxel sensitivity in cervical cancers.
  
  
Entity Ovarian Cancer.
Oncogenesis RAF-1 dysregulation is associated with poor prognosis and possibly carcinogenesis. RAF-1 inhibition using RNAi reduces cellular proliferatin and inhibits ovarian tumor cell growth in vitro and in vivo. Similar results were observed using antisense oligonucleotide (ASO) therapy (ISIS 5132 and ISIS 13650).
RAF-1 inhibition by the Akt pathway sensitizes human ovarian cancer cells to the drug paclitaxel.
  
  
Entity Gastric Cancer.
Oncogenesis RAF-1 inactivation using RNAi in gastric cancer cell line SGC7901 led to dramatic reductions in angiogenesis, increased apoptosis, and decreased cellular proliferation.
  
  
Entity Bladder Cancer.
Oncogenesis RAF-1 gene amplification was detected in 4% of bladder cancer samples. Deletions at the RAF-1 locus were detected in 2.2% of these samples. Both amplifications and deletions were heavily correlated with high tumor grade (P < 0.00001), advanced stage (P < 0.0001), and poor survival (P<0.05).
  
  
Entity Lymphoma.
Oncogenesis RAF-1 is typically over-expressed in thymic lymphomas from TCR transgenic mice.
  

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AAL881, a novel small molecule inhibitor of RAF and vascular endothelial growth factor receptor activities, blocks the growth of malignant glioma.
Sathornsumetee S, Hjelmeland AB, Keir ST, McLendon RE, Batt D, Ramsey T, Yusuff N, Rasheed BK, Kieran MW, Laforme A, Bigner DD, Friedman HS, Rich JN
Cancer research. 2006 ; 66 (17) : 8722-8730.
PMID 16951188
 
High-throughput tissue microarray analysis of 3p25 (RAF1) and 8p12 (FGFR1) copy number alterations in urinary bladder cancer.
Simon R, Richter J, Wagner U, Fijan A, Bruderer J, Schmid U, Ackermann D, Maurer R, Alund G, Knönagel H, Rist M, Wilber K, Anabitarte M, Hering F, Hardmeier T, Schönenberger A, Flury R, Jäger P, Fehr JL, Schraml P, Moch H, Mihatsch MJ, Gasser T, Sauter G
Cancer research. 2001 ; 61 (11) : 4514-4519.
PMID 11389083
 
Raf-1 activation suppresses neuroendocrine marker and hormone levels in human gastrointestinal carcinoid cells.
Sippel RS, Carpenter JE, Kunnimalaiyaan M, Lagerholm S, Chen H
American journal of physiology. Gastrointestinal and liver physiology. 2003 ; 285 (2) : G245-G254.
PMID 12851216
 
Autocrine/paracrine regulation of breast cancer cell proliferation by growth hormone releasing hormone via Ras, Raf, and mitogen-activated protein kinase.
Siriwardana G, Bradford A, Coy D, Zeitler P
Molecular endocrinology (Baltimore, Md.). 2006 ; 20 (9) : 2010-2019.
PMID 16613992
 
Expression of raf family proto-oncogenes in normal mouse tissues.
Storm SM, Cleveland JL, Rapp UR
Oncogene. 1990 ; 5 (3) : 345-351.
PMID 1690378
 
Raf and the road to cell survival: a tale of bad spells, ring bearers and detours.
Troppmair J, Rapp UR
Biochemical pharmacology. 2003 ; 66 (8) : 1341-1345.
PMID 14555207
 
14-3-3 proteins: active cofactors in cellular regulation by serine/threonine phosphorylation.
Tzivion G, Avruch J
The Journal of biological chemistry. 2002 ; 277 (5) : 3061-3064.
PMID 11709560
 
In-vivo activation of Raf-1 inhibits tumor growth and development in a xenograft model of human medullary thyroid cancer.
Vaccaro A, Chen H, Kunnimalaiyaan M
Anti-cancer drugs. 2006 ; 17 (7) : 849-853.
PMID 16926634
 
ZM336372, a Raf-1 activator, suppresses growth and neuroendocrine hormone levels in carcinoid tumor cells.
Van Gompel JJ, Kunnimalaiyaan M, Holen K, Chen H
Molecular cancer therapeutics. 2005 ; 4 (6) : 910-917.
PMID 15956248
 
Bcl-2 targets the protein kinase Raf-1 to mitochondria.
Wang HG, Rapp UR, Reed JC
Cell. 1996 ; 87 (4) : 629-638.
PMID 8929532
 
Bcl-2 interacting protein, BAG-1, binds to and activates the kinase Raf-1.
Wang HG, Takayama S, Rapp UR, Reed JC
Proceedings of the National Academy of Sciences of the United States of America. 1996 ; 93 (14) : 7063-7068.
PMID 8692945
 
Raf-1 physically interacts with Rb and regulates its function: a link between mitogenic signaling and cell cycle regulation.
Wang S, Ghosh RN, Chellappan SP
Molecular and cellular biology. 1998 ; 18 (12) : 7487-7498.
PMID 9819434
 
The effects of beta-estradiol on Raf activity, cell cycle progression and growth factor synthesis in the MCF-7 breast cancer cell line.
Weinstein-Oppenheimer CR, Burrows C, Steelman LS, McCubrey JA
Cancer biology & therapy. 2002 ; 1 (3) : 256-262.
PMID 12432273
 
The RAF proteins take centre stage.
Wellbrock C, Karasarides M, Marais R
Nature reviews. Molecular cell biology. 2004 ; 5 (11) : 875-885.
PMID 15520807
 
How Ras-related proteins talk to their effectors.
Wittinghofer A, Nassar N
Trends in biochemical sciences. 1996 ; 21 (12) : 488-491.
PMID 9009833
 
Craf-1 protein kinase is essential for mouse development.
Wojnowski L, Stancato LF, Zimmer AM, Hahn H, Beck TW, Larner AC, Rapp UR, Zimmer A
Mechanisms of development. 1998 ; 76 (1-2) : 141-149.
PMID 9767153
 
Phosphorylation and regulation of Raf by Akt (protein kinase B).
Zimmermann S, Moelling K
Science (New York, N.Y.). 1999 ; 286 (5445) : 1741-1744.
PMID 10576742
 

Citation

This paper should be referenced as such :
Chen, H ; Kunnimalaiyaan, M ; Greentblatt, DY ; Cayo, M
RAF1 (v-raf-1 murine leukemia viral oncogene homolog 1)
Atlas Genet Cytogenet Oncol Haematol. 2007;11(3):239-244.
Free journal version : [ pdf ]   [ DOI ]
On line version : http://AtlasGeneticsOncology.org/Genes/RAF1ID42032ch3p25.html

Other Leukemias implicated (Data extracted from papers in the Atlas) [ 2 ]
  Mycosis fungoides/Sezary's syndrome
Primary cutaneous CD30+ anaplastic large cell lymphoma

Other Solid tumors implicated (Data extracted from papers in the Atlas) [ 3 ]
  Neuro-Endocrine/Endocrine System: Carcinoid tumors
Thyroid: Papillary Carcinoma with inv(7)(q21q34) AKAP9/BRAF
Prostate tumors: an overview

Other Cancer prone implicated (Data extracted from papers in the Atlas) [ 2 ]
  LEOPARD syndrome Noonan syndrome

External links

Nomenclature
HGNC (Hugo)RAF1   9829
LRG (Locus Reference Genomic)LRG_413
Cards
AtlasRAF1ID42032ch3p25
Entrez_Gene (NCBI)RAF1  5894  Raf-1 proto-oncogene, serine/threonine kinase
AliasesCMD1NN; CRAF; NS5; Raf-1; 
c-Raf
GeneCards (Weizmann)RAF1
Ensembl hg19 (Hinxton)ENSG00000132155 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000132155 [Gene_View]  chr3:12583601-12664201 [Contig_View]  RAF1 [Vega]
ICGC DataPortalENSG00000132155
TCGA cBioPortalRAF1
AceView (NCBI)RAF1
Genatlas (Paris)RAF1
WikiGenes5894
SOURCE (Princeton)RAF1
Genetics Home Reference (NIH)RAF1
Genomic and cartography
GoldenPath hg38 (UCSC)RAF1  -     chr3:12583601-12664201 -  3p25.2   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)RAF1  -     3p25.2   [Description]    (hg19-Feb_2009)
EnsemblRAF1 - 3p25.2 [CytoView hg19]  RAF1 - 3p25.2 [CytoView hg38]
Mapping of homologs : NCBIRAF1 [Mapview hg19]  RAF1 [Mapview hg38]
OMIM164760   611553   611554   615916   
Gene and transcription
Genbank (Entrez)AK226028 AK303561 AK303920 AK312248 BC018119
RefSeq transcript (Entrez)NM_002880
RefSeq genomic (Entrez)
Consensus coding sequences : CCDS (NCBI)RAF1
Cluster EST : UnigeneHs.159130 [ NCBI ]
CGAP (NCI)Hs.159130
Alternative Splicing GalleryENSG00000132155
Gene ExpressionRAF1 [ NCBI-GEO ]   RAF1 [ EBI - ARRAY_EXPRESS ]   RAF1 [ SEEK ]   RAF1 [ MEM ]
Gene Expression Viewer (FireBrowse)RAF1 [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)5894
GTEX Portal (Tissue expression)RAF1
Protein : pattern, domain, 3D structure
UniProt/SwissProtP04049   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtP04049  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProP04049
Splice isoforms : SwissVarP04049
Catalytic activity : Enzyme2.7.11.1 [ Enzyme-Expasy ]   2.7.11.12.7.11.1 [ IntEnz-EBI ]   2.7.11.1 [ BRENDA ]   2.7.11.1 [ KEGG ]   
PhosPhoSitePlusP04049
Domaine pattern : Prosite (Expaxy)PROTEIN_KINASE_ATP (PS00107)    PROTEIN_KINASE_DOM (PS50011)    PROTEIN_KINASE_ST (PS00108)    RBD (PS50898)    ZF_DAG_PE_1 (PS00479)    ZF_DAG_PE_2 (PS50081)   
Domains : Interpro (EBI)DAG/PE-bd    Kinase-like_dom    PE/DAG-bd    Prot_kinase_dom    Protein_kinase_ATP_BS    RBD_dom    Ser-Thr/Tyr_kinase_cat_dom    Ser/Thr_kinase_AS    Ubiquitin-rel_dom   
Domain families : Pfam (Sanger)C1_1 (PF00130)    Pkinase_Tyr (PF07714)    RBD (PF02196)   
Domain families : Pfam (NCBI)pfam00130    pfam07714    pfam02196   
Domain families : Smart (EMBL)C1 (SM00109)  RBD (SM00455)  S_TKc (SM00220)  
Conserved Domain (NCBI)RAF1
DMDM Disease mutations5894
Blocks (Seattle)RAF1
PDB (SRS)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
PDB (PDBSum)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
PDB (IMB)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
PDB (RSDB)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
Structural Biology KnowledgeBase1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
SCOP (Structural Classification of Proteins)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
CATH (Classification of proteins structures)1C1Y    1FAQ    1FAR    1GUA    1RFA    3CU8    3IQJ    3IQU    3IQV    3KUC    3KUD    3NKX    3O8I    3OMV    4FJ3    4G0N    4G3X    4IEA    4IHL   
SuperfamilyP04049
Human Protein AtlasENSG00000132155
Peptide AtlasP04049
HPRD01265
IPIIPI00021786   IPI00900312   IPI01015603   IPI00788857   IPI00925565   
Protein Interaction databases
DIP (DOE-UCLA)P04049
IntAct (EBI)P04049
FunCoupENSG00000132155
BioGRIDRAF1
STRING (EMBL)RAF1
ZODIACRAF1
Ontologies - Pathways
QuickGOP04049
Ontology : AmiGOMAPK cascade  activation of MAPKK activity  response to hypoxia  stimulatory C-type lectin receptor signaling pathway  protein kinase activity  protein serine/threonine kinase activity  MAP kinase kinase kinase activity  protein binding  ATP binding  cytoplasm  cytoplasm  mitochondrial outer membrane  Golgi apparatus  cytosol  cytosol  plasma membrane  plasma membrane  plasma membrane  protein phosphorylation  apoptotic process  signal transduction  activation of adenylate cyclase activity  multicellular organism development  heart development  cell proliferation  negative regulation of cell proliferation  negative regulation of signal transduction  kinase activity  nuclear speck  enzyme binding  cell differentiation  platelet activation  thyroid gland development  pseudopodium  small GTPase binding  negative regulation of protein complex assembly  mitogen-activated protein kinase kinase binding  positive regulation of peptidyl-serine phosphorylation  ion transmembrane transport  somatic stem cell population maintenance  regulation of Rho protein signal transduction  insulin secretion involved in cellular response to glucose stimulus  response to muscle stretch  wound healing  identical protein binding  regulation of apoptotic process  negative regulation of apoptotic process  negative regulation of cysteine-type endopeptidase activity involved in apoptotic process  intermediate filament cytoskeleton organization  regulation of cell differentiation  positive regulation of transcription from RNA polymerase II promoter  metal ion binding  protein heterodimerization activity  neurotrophin TRK receptor signaling pathway  thymus development  face development  positive regulation of ERK1 and ERK2 cascade  death-inducing signaling complex assembly  negative regulation of extrinsic apoptotic signaling pathway via death domain receptors  regulation of cell motility  
Ontology : EGO-EBIMAPK cascade  activation of MAPKK activity  response to hypoxia  stimulatory C-type lectin receptor signaling pathway  protein kinase activity  protein serine/threonine kinase activity  MAP kinase kinase kinase activity  protein binding  ATP binding  cytoplasm  cytoplasm  mitochondrial outer membrane  Golgi apparatus  cytosol  cytosol  plasma membrane  plasma membrane  plasma membrane  protein phosphorylation  apoptotic process  signal transduction  activation of adenylate cyclase activity  multicellular organism development  heart development  cell proliferation  negative regulation of cell proliferation  negative regulation of signal transduction  kinase activity  nuclear speck  enzyme binding  cell differentiation  platelet activation  thyroid gland development  pseudopodium  small GTPase binding  negative regulation of protein complex assembly  mitogen-activated protein kinase kinase binding  positive regulation of peptidyl-serine phosphorylation  ion transmembrane transport  somatic stem cell population maintenance  regulation of Rho protein signal transduction  insulin secretion involved in cellular response to glucose stimulus  response to muscle stretch  wound healing  identical protein binding  regulation of apoptotic process  negative regulation of apoptotic process  negative regulation of cysteine-type endopeptidase activity involved in apoptotic process  intermediate filament cytoskeleton organization  regulation of cell differentiation  positive regulation of transcription from RNA polymerase II promoter  metal ion binding  protein heterodimerization activity  neurotrophin TRK receptor signaling pathway  thymus development  face development  positive regulation of ERK1 and ERK2 cascade  death-inducing signaling complex assembly  negative regulation of extrinsic apoptotic signaling pathway via death domain receptors  regulation of cell motility  
Pathways : BIOCARTA [Genes]   
Pathways : KEGG   
REACTOMEP04049 [protein]
REACTOME PathwaysR-HSA-6802955 [pathway]   
NDEx NetworkRAF1
Atlas of Cancer Signalling NetworkRAF1
Wikipedia pathwaysRAF1
Orthology - Evolution
OrthoDB5894
GeneTree (enSembl)ENSG00000132155
Phylogenetic Trees/Animal Genes : TreeFamRAF1
HOVERGENP04049
HOGENOMP04049
Homologs : HomoloGeneRAF1
Homology/Alignments : Family Browser (UCSC)RAF1
Gene fusions - Rearrangements
Fusion : MitelmanAGGF1/RAF1 [5q13.3/3p25.2]  [t(3;5)(p25;q13)]  
Fusion : MitelmanCMTM8/RAF1 [3p22.3/3p25.2]  [t(3;3)(p22;p25)]  
Fusion : MitelmanESRP1/RAF1 [8q22.1/3p25.2]  [t(3;8)(p25;q22)]  
Fusion : MitelmanFYCO1/RAF1 [3p21.31/3p25.2]  [t(3;3)(p21;p25)]  
Fusion : MitelmanQKI/RAF1 [6q26/3p25.2]  [t(3;6)(p25;q26)]  
Fusion : MitelmanRAF1/DAZL [3p25.2/3p24.3]  [t(3;3)(p24;p25)]  
Fusion : MitelmanRAF1/TMEM40 [3p25.2/3p25.2]  [t(3;3)(p25;p25)]  
Fusion : MitelmanRAF1/XPC [3p25.2/3p25.1]  [t(3;3)(p25;p25)]  
Fusion : MitelmanSRGAP3/RAF1 [3p25.3/3p25.2]  [dup(3)(p25p25)]  
Fusion : COSMICESRP1 [8q22.1]  -  RAF1 [3p25.2]  [fusion_826]  [fusion_830]  
Fusion : COSMICHACL1 [3p25.1]  -  RAF1 [3p25.2]  [fusion_1732]  [fusion_1733]  
Fusion : COSMICRAF1 [3p25.2]  -  DAZL [3p24.3]  [fusion_675]  [fusion_676]  
Fusion : COSMICRAF1 [3p25.2]  -  ESRP1 [8q22.1]  [fusion_827]  [fusion_873]  
Fusion : COSMICSRGAP3 [3p25.3]  -  RAF1 [3p25.2]  [fusion_1195]  [fusion_660]  [fusion_661]  [fusion_662]  
Fusion: TCGAAGGF1 5q13.3 RAF1 3p25.2 PRAD THCA
Fusion: TCGACMTM8 3p22.3 RAF1 3p25.2 BRCA
Fusion: TCGARAF1 3p25.2 TMEM40 3p25.2 KIRC
Fusion: TCGARAF1 3p25.2 XPC 3p25.1 LUAD
Fusion : TICdbESRP1 [8q22.1]  -  RAF1 [3p25.2]
Fusion : TICdbSRGAP3 [3p25.3]  -  RAF1 [3p25.2]
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerRAF1 [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)RAF1
dbVarRAF1
ClinVarRAF1
1000_GenomesRAF1 
Exome Variant ServerRAF1
ExAC (Exome Aggregation Consortium)RAF1 (select the gene name)
Genetic variants : HAPMAP5894
Genomic Variants (DGV)RAF1 [DGVbeta]
DECIPHERRAF1 [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisRAF1 
Mutations
ICGC Data PortalRAF1 
TCGA Data PortalRAF1 
Broad Tumor PortalRAF1
OASIS PortalRAF1 [ Somatic mutations - Copy number]
Cancer Gene: CensusRAF1 
Somatic Mutations in Cancer : COSMICRAF1  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDRAF1
LOVD (Leiden Open Variation Database)Whole genome datasets
LOVD (Leiden Open Variation Database)LOVD - Leiden Open Variation Database
LOVD (Leiden Open Variation Database)LOVD 3.0 shared installation
BioMutasearch RAF1
DgiDB (Drug Gene Interaction Database)RAF1
DoCM (Curated mutations)RAF1 (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)RAF1 (select a term)
intoGenRAF1
NCG5 (London)RAF1
Cancer3DRAF1(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Diseases
OMIM164760    611553    611554    615916   
Orphanet206    635    1032    19659   
MedgenRAF1
Genetic Testing Registry RAF1
NextProtP04049 [Medical]
TSGene5894
GENETestsRAF1
Target ValidationRAF1
Huge Navigator RAF1 [HugePedia]
snp3D : Map Gene to Disease5894
BioCentury BCIQRAF1
ClinGenRAF1 (curated)
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD5894
Chemical/Pharm GKB GenePA34183
Clinical trialRAF1
Miscellaneous
canSAR (ICR)RAF1 (select the gene name)
Probes
Litterature
PubMed499 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
CoreMineRAF1
EVEXRAF1
GoPubMedRAF1
iHOPRAF1
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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