Просмотр кода страницы RHEB

{{GNF_Protein_box
 | Name = Гомолог Ras, витаминизирующий мозг
 | image = Protein_RHEB_PDB_1xtq.png
 | image_source = [[Protein_Data_Bank|PDB]] представлен на примере 1xtq.
 | PDB = {{PDB2|1XTQ}}, {{PDB2|1XTR}}, {{PDB2|1XTS}}, {{PDB2|3SEA}}, {{PDB2|3T5G}}
 | HGNCid = 10011
 | MGIid = 97912
 | Symbol = RHEB
 | AltSymbols =; RHEB2
 | IUPHAR = 
 | ChEMBL = 
 | OMIM = 601293
 | ECnumber = 
 | Homologene = 123916
 | GeneAtlas_image1 = PBB_GE_RHEB_201452_at_tn.png
 | GeneAtlas_image2 = PBB_GE_RHEB_201453_x_at_tn.png
 | GeneAtlas_image3 = PBB_GE_RHEB_213404_s_at_tn.png
 | Protein_domain_image = 
 | Function = {{GNF_GO|id=GO:0003924 |text = GTPase activity}} {{GNF_GO|id=GO:0005515 |text = protein binding}} {{GNF_GO|id=GO:0005525 |text = GTP binding}} {{GNF_GO|id=GO:0019003 |text = GDP binding}} {{GNF_GO|id=GO:0019901 |text = protein kinase binding}} {{GNF_GO|id=GO:0046872 |text = metal ion binding}}
 | Component = {{GNF_GO|id=GO:0005681 |text = spliceosomal complex}} {{GNF_GO|id=GO:0005829 |text = cytosol}} {{GNF_GO|id=GO:0005886 |text = plasma membrane}} {{GNF_GO|id=GO:0016020 |text = membrane}} {{GNF_GO|id=GO:0030425 |text = dendrite}} {{GNF_GO|id=GO:0043025 |text = neuronal cell body}} {{GNF_GO|id=GO:0070062 |text = extracellular vesicular exosome}}
 | Process = {{GNF_GO|id=GO:0006184 |text = GTP catabolic process}} {{GNF_GO|id=GO:0007050 |text = cell cycle arrest}} {{GNF_GO|id=GO:0007165 |text = signal transduction}} {{GNF_GO|id=GO:0007264 |text = small GTPase mediated signal transduction}} {{GNF_GO|id=GO:0008286 |text = insulin receptor signaling pathway}} {{GNF_GO|id=GO:0032008 |text = positive regulation of TOR signaling}}
 | Hs_EntrezGene = 6009
 | Hs_Ensembl = ENSG00000106615
 | Hs_RefseqmRNA = NM_005614
 | Hs_RefseqProtein = NP_005605
 | Hs_GenLoc_db = hg19
 | Hs_GenLoc_chr = 7
 | Hs_GenLoc_start = 151163098
 | Hs_GenLoc_end = 151217206
 | Hs_Uniprot = Q15382
 | Mm_EntrezGene = 19744
 | Mm_Ensembl = ENSMUSG00000028945
 | Mm_RefseqmRNA = NM_053075
 | Mm_RefseqProtein = NP_444305
 | Mm_GenLoc_db = mm10
 | Mm_GenLoc_chr = 5
 | Mm_GenLoc_start = 24802823
 | Mm_GenLoc_end = 24842624
 | Mm_Uniprot = Q921J2
 | path = PBB/6009
}}

''' ГТФ-связывающий белок Rheb ''' также известный как  ''' гомолог Ras, витаминизирующий мозг ''' (Rheb) — [[белок]], кодируемый  у человека [[ген]]ом '' Rheb ''. <ref name="pmid8661031">{{cite journal | author = Mizuki N, Kimura M, Ohno S, Miyata S, Sato M, Ando H, Ishihara M, Goto K, Watanabe S, Yamazaki M, Ono A, Taguchi S, Okumura K, Nogami M, Taguchi T, Ando A, Inoko H | title = Isolation of cDNA and genomic clones of a human Ras-related GTP-binding protein gene and its chromosomal localization to the long arm of chromosome 7, 7q36 | journal = Genomics | volume = 34 | issue = 1 | pages = 114–8 |date=Dec 1996 | pmid = 8661031 | pmc =  | doi =10.1006/geno.1996.0248  }}</ref>


== Функция ==
Rheb является недавно обнаруженным членом {{нп5|Надсемейство Ras|надсемейства Ras||Ras superfamily}}, который может быть вовлечён в пластичность нейронов. Эта функция является новой и обычно не связанной с белками Ras.

Этот ген является членом небольшого [[ГТФ]] надсемейства и кодирует липидный якорь, [[белок]] клеточной мембраны с пятью повторами RAS, связанными с ГТФ-связывающей областью. Этот белок играет важную роль в регуляции роста и клеточного цикла, усиленную  его ролью в сигнальном пути [[инсулин]]а/{{нп5|mammalian target of rapamycin|||Mechanistic target of rapamycin}}/[[S6K]]. Белок обладает активностью ГТФ и является челноком между границей [[Гуанозиндифосфат|GDP]] и границей [[Гуанозинтрифосфат|GTP]].Для этой деятельности требуется фарнезилирование белка. Для трёх псевдогенов были созданы карты, двух на хромосоме 10 и одного на хромосоме 22. <ref name="entrez">{{cite web | title = Entrez Gene: RHEB Ras homolog enriched in brain| url = http://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=6009| accessdate = }}</ref>

== Взаимодействия ==
Rheb,как было выявлено, [[Белок-белковые взаимодействия|взаимодействует]] с:
[[C-Raf]],<ref name=pmid15854902>{{cite journal |last=Long |first=Xiaomeng |authorlink= |coauthors=Lin Yenshou, Ortiz-Vega Sara, Yonezawa Kazuyoshi, Avruch Joseph |date=Apr 2005 |title=Rheb binds and regulates the mTOR kinase |journal=Curr. Biol. |volume=15 |issue=8 |pages=702–13 |publisher= |location = England| issn = 0960-9822| pmid = 15854902 |doi = 10.1016/j.cub.2005.02.053 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid15150271>{{cite journal |last=Karbowniczek |first=Magdalena |authorlink= |author2=Cash Timothy |author3=Cheung Mitchell |author4=Robertson Gavin P |author5=Astrinidis Aristotelis |author6=Henske Elizabeth Petri  |date=Jul 2004 |title=Regulation of B-Raf kinase activity by tuberin and Rheb is mammalian target of rapamycin (mTOR)-independent |journal=J. Biol. Chem. |volume=279 |issue=29 |pages=29930–7 |publisher= |location = United States| issn = 0021-9258| pmid = 15150271 |doi = 10.1074/jbc.M402591200 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid9001246>{{cite journal |last=Yee |first=W M |authorlink= |author2=Worley P F  |date=Feb 1997 |title=Rheb interacts with Raf-1 kinase and may function to integrate growth factor- and protein kinase A-dependent signals |journal=Mol. Cell. Biol. |volume=17 |issue=2 |pages=921–33 |publisher= |location = UNITED STATES| issn = 0270-7306| pmid = 9001246 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = |pmc=231818 }}</ref> {{нп5|Mammalian target of rapamycin|||Mammalian target of rapamycin}},<ref name=pmid15854902/><ref name=pmid15878852>{{cite journal |last=Long |first=Xiaomeng |authorlink= |coauthors=Ortiz-Vega Sara, Lin Yenshou, Avruch Joseph |date=Jun 2005 |title=Rheb binding to mammalian target of rapamycin (mTOR) is regulated by amino acid sufficiency |journal=J. Biol. Chem. |volume=280 |issue=25 |pages=23433–6 |publisher= |location = United States| issn = 0021-9258| pmid = 15878852 |doi = 10.1074/jbc.C500169200 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid15772076>{{cite journal |last=Smith |first=Ewan M |authorlink= |author2=Finn Stephen G |author3=Tee Andrew R |author4=Browne Gareth J |author5=Proud Christopher G  |date=May 2005 |title=The tuberous sclerosis protein TSC2 is not required for the regulation of the mammalian target of rapamycin by amino acids and certain cellular stresses |journal=J. Biol. Chem. |volume=280 |issue=19 |pages=18717–27 |publisher= |location = United States| issn = 0021-9258| pmid = 15772076 |doi = 10.1074/jbc.M414499200 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid16915281>{{cite journal |last=Bernardi |first=Rosa |authorlink= |coauthors=Guernah Ilhem, Jin David, Grisendi Silvia, Alimonti Andrea, Teruya-Feldstein Julie, [[Carlos Cordon-Cardo|Cordon-Cardo Carlos]], Simon M Celeste, Rafii Shahin, Pandolfi Pier Paolo |date=Aug 2006 |title=PML inhibits HIF-1alpha translation and neoangiogenesis through repression of mTOR |journal=[[Nature (journal)|Nature]] |volume=442 |issue=7104 |pages=779–85 |publisher= |location = England| issn = | pmid = 16915281 |doi = 10.1038/nature05029 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref> [[TSC2]],<ref name=pmid15854902/><ref name=pmid12842888>{{cite journal |last=Castro |first=Ariel F |authorlink= |author2=Rebhun John F |author3=Clark Geoffrey J |author4=Quilliam Lawrence A  |date=Aug 2003 |title=Rheb binds tuberous sclerosis complex 2 (TSC2) and promotes S6 kinase activation in a rapamycin- and farnesylation-dependent manner |journal=J. Biol. Chem. |volume=278 |issue=35 |pages=32493–6 |publisher= |location = United States| issn = 0021-9258| pmid = 12842888 |doi = 10.1074/jbc.C300226200 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid12869586>{{cite journal |last=Inoki |first=Ken |authorlink= |author2=Li Yong |author3=Xu Tian |author4=Guan Kun-Liang  |date=Aug 2003 |title=Rheb GTPase is a direct target of TSC2 GAP activity and regulates mTOR signaling |journal=Genes Dev. |volume=17 |issue=15 |pages=1829–34 |publisher= |location = United States| issn = 0890-9369| pmid = 12869586 |doi = 10.1101/gad.1110003 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = |pmc=196227 }}</ref><ref name=pmid12820960>{{cite journal |last=Garami |first=Attila |authorlink= |author2=Zwartkruis Fried J T |author3=Nobukuni Takahiro |author4=Joaquin Manel |author5=Roccio Marta |author6=Stocker Hugo |author7=Kozma Sara C |author8=Hafen Ernst |author9=Bos Johannes L |author10=Thomas George  |date=Jun 2003 |title=Insulin activation of Rheb, a mediator of mTOR/S6K/4E-BP signaling, is inhibited by TSC1 and 2 |journal=Mol. Cell |volume=11 |issue=6 |pages=1457–66 |publisher= |location = United States| issn = 1097-2765| pmid = 12820960 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = |doi=10.1016/S1097-2765(03)00220-X }}</ref><ref name=pmid12771962>{{cite journal |last=Zhang |first=Yong |authorlink= |author2=Gao Xinsheng |author3=Saucedo Leslie J |author4=Ru Binggen |author5=Edgar Bruce A |author6=Pan Duojia  |date=Jun 2003 |title=Rheb is a direct target of the tuberous sclerosis tumour suppressor proteins |journal=Nat. Cell Biol. |volume=5 |issue=6 |pages=578–81 |publisher= |location = England| issn = 1465-7392| pmid = 12771962 |doi = 10.1038/ncb999 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref><ref name=pmid17077083>{{cite journal |last=Cao |first=Yongheng |authorlink= |author2=Kamioka Yuji |author3=Yokoi Norihide |author4=Kobayashi Toshiyuki |author5=Hino Okio |author6=Onodera Masafumi |author7=Mochizuki Naoki |author8=Nakae Jun  |date=Dec 2006 |title=Interaction of FoxO1 and TSC2 induces insulin resistance through activation of the mammalian target of rapamycin/p70 S6K pathway |journal=J. Biol. Chem. |volume=281 |issue=52 |pages=40242–51 |publisher= |location = United States| issn = 0021-9258| pmid = 17077083 |doi = 10.1074/jbc.M608116200 | bibcode = | oclc =| id = | url = | language = | format = | accessdate = | laysummary = | laysource = | laydate = | quote = }}</ref> [[ATM (белок)|ATM]],<ref name=pmid15854902/> [[KIAA1303]]<ref name=pmid15854902/>  [[ATR (биология)|ATR]].<ref name=pmid15854902/>

== Примечания ==
{{Примечания}}

== Литература ==
{{refbegin | 2}}
*{{cite journal  | author=Yamagata K |title=rheb, a growth factor- and synaptic activity-regulated gene, encodes a novel Ras-related protein |journal=J. Biol. Chem. |volume=269 |issue= 23 |pages= 16333–9 |year= 1994 |pmid= 8206940 |doi=  | author-separator=,  | author2=Sanders LK  | author3=Kaufmann WE  | display-authors=3  | last4=Yee  | first4=W  | last5=Barnes  | first5=CA  | last6=Nathans  | first6=D  | last7=Worley  | first7=PF  }}
*{{cite journal  | author=Gromov PS, Madsen P, Tomerup N, Celis JE |title=A novel approach for expression cloning of small GTPases: identification, tissue distribution and chromosome mapping of the human homolog of rheb |journal=FEBS Lett. |volume=377 |issue= 2 |pages= 221–6 |year= 1996 |pmid= 8543055 |doi=10.1016/0014-5793(95)01349-0  }}
*{{cite journal  | author=Bonaldo MF, Lennon G, Soares MB |title=Normalization and subtraction: two approaches to facilitate gene discovery |journal=Genome Res. |volume=6 |issue= 9 |pages= 791–806 |year= 1997 |pmid= 8889548 |doi=10.1101/gr.6.9.791  }}
*{{cite journal  | author=Yee WM, Worley PF |title=Rheb interacts with Raf-1 kinase and may function to integrate growth factor- and protein kinase A-dependent signals |journal=Mol. Cell. Biol. |volume=17 |issue= 2 |pages= 921–33 |year= 1997 |pmid= 9001246 |doi=  | pmc=231818  }}
*{{cite journal  | author=Clark GJ |title=The Ras-related protein Rheb is farnesylated and antagonizes Ras signaling and transformation |journal=J. Biol. Chem. |volume=272 |issue= 16 |pages= 10608–15 |year= 1997 |pmid= 9099708 |doi=10.1074/jbc.272.16.10608  | author-separator=,  | author2=Kinch MS  | author3=Rogers-Graham K  | display-authors=3  | last4=Sebti  | first4=SM  | last5=Hamilton  | first5=AD  | last6=Der  | first6=CJ  }}
*{{cite journal  | author=Inohara N, Ding L, Chen S, Núñez G |title=harakiri, a novel regulator of cell death, encodes a protein that activates apoptosis and interacts selectively with survival-promoting proteins Bcl-2 and Bcl-X(L) |journal=EMBO J. |volume=16 |issue= 7 |pages= 1686–94 |year= 1997 |pmid= 9130713 |doi= 10.1093/emboj/16.7.1686  | pmc=1169772 }}
*{{cite journal  | author= |title=Toward a complete human genome sequence |journal=Genome Res. |volume=8 |issue= 11 |pages= 1097–108 |year= 1999 |pmid= 9847074 |doi=  10.1101/gr.8.11.1097  | author1= <Please add first missing authors to populate metadata.>}}
*{{cite journal  | author=Kita K |title=Search for UV-responsive genes in human cells by differential mRNA display: involvement of human ras-related GTP-binding protein, Rheb, in UV susceptibility |journal=Biochem. Biophys. Res. Commun. |volume=274 |issue= 3 |pages= 859–64 |year= 2000 |pmid= 10924367 |doi= 10.1006/bbrc.2000.3220  | author-separator=,  | author2=Wu YP  | author3=Sugaya S  | display-authors=3  | last4=Moriya  | first4=Tetsuhiro  | last5=Nomura  | first5=Jun  | last6=Takahashi  | first6=Shunji  | last7=Yamamori  | first7=Hideo  | last8=Nakajima  | first8=Nobuyuki  | last9=Suzuki  | first9=Nobuo }}
*{{cite journal  | author=Hanzal-Bayer M |title=The complex of Arl2-GTP and PDEδ: from structure to function |journal=EMBO J. |volume=21 |issue= 9 |pages= 2095–106 |year= 2002 |pmid= 11980706 |doi= 10.1093/emboj/21.9.2095  | pmc=125981  | author-separator=,  | author2=Renault L  | author3=Roversi P  | display-authors=3  | last4=Wittinghofer  | first4=A  | last5=Hillig  | first5=RC }}
*{{cite journal  | author=Strausberg RL |title=Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=99 |issue= 26 |pages= 16899–903 |year= 2003 |pmid= 12477932 |doi= 10.1073/pnas.242603899  | pmc=139241  | author-separator=,  | author2=Feingold EA  | author3=Grouse LH  | display-authors=3  | last4=Derge  | first4=JG  | last5=Klausner  | first5=RD  | last6=Collins  | first6=FS  | last7=Wagner  | first7=L  | last8=Shenmen  | first8=CM  | last9=Schuler  | first9=GD }}
*{{cite journal  | author=Scherer SW |title=Human Chromosome 7: DNA Sequence and Biology |journal=Science |volume=300 |issue= 5620 |pages= 767–72 |year= 2003 |pmid= 12690205 |doi= 10.1126/science.1083423  | pmc=2882961  | author-separator=,  | author2=Cheung J  | author3=MacDonald JR  | display-authors=3  | last4=Osborne  | first4=LR  | last5=Nakabayashi  | first5=K  | last6=Herbrick  | first6=JA  | last7=Carson  | first7=AR  | last8=Parker-Katiraee  | first8=L  | last9=Skaug  | first9=J }}
*{{cite journal  | author=Zhang Y |title=Rheb is a direct target of the tuberous sclerosis tumour suppressor proteins |journal=Nat. Cell Biol. |volume=5 |issue= 6 |pages= 578–81 |year= 2003 |pmid= 12771962 |doi= 10.1038/ncb999  | author-separator=,  | author2=Gao X  | author3=Saucedo LJ  | display-authors=3  | last4=Ru  | first4=Binggen  | last5=Edgar  | first5=Bruce A.  | last6=Pan  | first6=Duojia }}
*{{cite journal  | author=Garami A |title=Insulin activation of Rheb, a mediator of mTOR/S6K/4E-BP signaling, is inhibited by TSC1 and 2 |journal=Mol. Cell |volume=11 |issue= 6 |pages= 1457–66 |year= 2003 |pmid= 12820960 |doi=10.1016/S1097-2765(03)00220-X  | author-separator=,  | author2=Zwartkruis FJ  | author3=Nobukuni T  | display-authors=3  | last4=Joaquin  | first4=Manel  | last5=Roccio  | first5=Marta  | last6=Stocker  | first6=Hugo  | last7=Kozma  | first7=Sara C  | last8=Hafen  | first8=Ernst  | last9=Bos  | first9=Johannes L  }}
*{{cite journal  | author=Castro AF, Rebhun JF, Clark GJ, Quilliam LA |title=Rheb binds tuberous sclerosis complex 2 (TSC2) and promotes S6 kinase activation in a rapamycin- and farnesylation-dependent manner |journal=J. Biol. Chem. |volume=278 |issue= 35 |pages= 32493–6 |year= 2003 |pmid= 12842888 |doi= 10.1074/jbc.C300226200 }}
*{{cite journal  | author=Hillier LW |title=The DNA sequence of human chromosome 7 |journal=Nature |volume=424 |issue= 6945 |pages= 157–64 |year= 2003 |pmid= 12853948 |doi= 10.1038/nature01782  | author-separator=,  | author2=Fulton RS  | author3=Fulton LA  | display-authors=3  | last4=Graves  | first4=Tina A.  | last5=Pepin  | first5=Kymberlie H.  | last6=Wagner-Mcpherson  | first6=Caryn  | last7=Layman  | first7=Dan  | last8=Maas  | first8=Jason  | last9=Jaeger  | first9=Sara }}
*{{cite journal  | author=Tabancay AP |title=Identification of dominant negative mutants of Rheb GTPase and their use to implicate the involvement of human Rheb in the activation of p70S6K |journal=J. Biol. Chem. |volume=278 |issue= 41 |pages= 39921–30 |year= 2003 |pmid= 12869548 |doi= 10.1074/jbc.M306553200  | author-separator=,  | author2=Gau CL  | author3=Machado IM  | display-authors=3  | last4=Uhlmann  | first4=EJ  | last5=Gutmann  | first5=DH  | last6=Guo  | first6=L  | last7=Tamanoi  | first7=F }}
*{{cite journal  | author=Inoki K, Li Y, Xu T, Guan KL |title=Rheb GTPase is a direct target of TSC2 GAP activity and regulates mTOR signaling |journal=Genes Dev. |volume=17 |issue= 15 |pages= 1829–34 |year= 2003 |pmid= 12869586 |doi= 10.1101/gad.1110003  | pmc=196227 }}
*{{cite journal  | author=Tee AR |title=Tuberous sclerosis complex gene products, Tuberin and Hamartin, control mTOR signaling by acting as a GTPase-activating protein complex toward Rheb |journal=Curr. Biol. |volume=13 |issue= 15 |pages= 1259–68 |year= 2004 |pmid= 12906785 |doi=10.1016/S0960-9822(03)00506-2  | author-separator=,  | author2=Manning BD  | author3=Roux PP  | display-authors=3  | last4=Cantley  | first4=Lewis C  | last5=Blenis  | first5=John  }}
*{{cite journal  | author=Ota T |title=Complete sequencing and characterization of 21,243 full-length human cDNAs |journal=Nat. Genet. |volume=36 |issue= 1 |pages= 40–5 |year= 2004 |pmid= 14702039 |doi= 10.1038/ng1285  | author-separator=,  | author2=Suzuki Y  | author3=Nishikawa T  | display-authors=3  | last4=Otsuki  | first4=Tetsuji  | last5=Sugiyama  | first5=Tomoyasu  | last6=Irie  | first6=Ryotaro  | last7=Wakamatsu  | first7=Ai  | last8=Hayashi  | first8=Koji  | last9=Sato  | first9=Hiroyuki }}
{{refend}}

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