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CYR61

CYR61
Dostupne strukture
PDBPretraga ortologa: PDBe RCSB
Spisak PDB ID kodova

4D0Z, 4D11

Identifikatori
AliasiCCN1
Vanjski ID-jeviOMIM: 602369 MGI: 88613 HomoloGene: 1194 GeneCards: CCN1
Lokacija gena (čovjek)
Hromosom 1 (čovjek)
Hrom.Hromosom 1 (čovjek)[1]
Hromosom 1 (čovjek)
Genomska lokacija za CYR61
Genomska lokacija za CYR61
Bend1p22.3Početak85,580,761 bp[1]
Kraj85,584,589 bp[1]
Lokacija gena (miš)
Hromosom 3 (miš)
Hrom.Hromosom 3 (miš)[2]
Hromosom 3 (miš)
Genomska lokacija za CYR61
Genomska lokacija za CYR61
Bend3 H2|3 70.18 cMPočetak145,352,731 bp[2]
Kraj145,355,736 bp[2]
Obrazac RNK ekspresije


Više referentnih podataka o ekspresiji
Ontologija gena
Molekularna funkcija• insulin-like growth factor binding
• growth factor binding
• extracellular matrix binding
• integrin binding
• heparin binding
• extracellular matrix structural constituent
Ćelijska komponenta• GO:0005578 Vanćelijski matriks
• extracellular region
• Vanćelijsko
• endoplasmic reticulum lumen
• collagen-containing extracellular matrix
Biološki proces• positive regulation of osteoblast proliferation
• apoptotic process involved in heart morphogenesis
• positive regulation of ceramide biosynthetic process
• positive regulation of protein phosphorylation
• ventricular septum development
• positive regulation of cell migration
• anatomical structure morphogenesis
• extracellular matrix organization
• negative regulation of apoptotic process
• labyrinthine layer blood vessel development
• reactive oxygen species metabolic process
• chorio-allantoic fusion
• chondroblast differentiation
• Hemotaksija
• positive regulation of phospholipase activity
• atrial septum morphogenesis
• positive regulation of cysteine-type endopeptidase activity involved in apoptotic process
• intussusceptive angiogenesis
• positive regulation of osteoblast differentiation
• atrioventricular valve morphogenesis
• positive regulation of cartilage development
• regulation of cell growth
• osteoblast differentiation
• positive regulation of cell differentiation
• regulation of ERK1 and ERK2 cascade
• positive regulation of BMP signaling pathway
• positive regulation of apoptotic process
• Ćelijska proliferacija
• wound healing, spreading of cells
• GO:0003257, GO:0010735, GO:1901228, GO:1900622, GO:1904488 positive regulation of transcription by RNA polymerase II
• positive regulation of protein kinase activity
• positive regulation of cell-substrate adhesion
• Posttranslacione modifikacije
• Ćelijska adhezija
• cell-cell signaling
• negative regulation of cell death
• cell-cell adhesion
• positive regulation of bone mineralization
Izvori:Amigo / QuickGO
Ortolozi
VrsteČovjekMiš
Entrez

3491

16007

Ensembl

ENSG00000142871

ENSMUSG00000028195

UniProt

O00622

P18406

RefSeq (mRNK)

NM_001554

NM_010516

RefSeq (bjelančevina)

NP_001545

NP_034646

Lokacija (UCSC)Chr 1: 85.58 – 85.58 MbChr 3: 145.35 – 145.36 Mb
PubMed pretraga[3][4]
Wikipodaci
Pogledaj/uredi – čovjekPogledaj/uredi – miš

Cisteinom bogati angiogeni induktor 61 (CYR61) ili član 1 porodice proteina CCN (CCN1), jest matrićelijski protein koji je kod ljudi kodiran genom CYR61 sa hromosoma 1.[5]

CYR61 je izlučeni, vanćelijskomatriksni (ECM)-vezan signalni protein iz CCN porodice (unutarćelijski signalni protein CCN).[6][7] CYR61 je sposoban da reguliše širok spektar ćelijskih aktivnosti, uključujući ćelijsku adheziju, migraciju, proliferaciju, diferencijaciju, apoptozu i starenje putem interakcije sa integrinskim receptorima na površini ćelije i heparan-sulfatnim proteoglikanima. Tokom embrionskog razvoja, CYR61 je kritičan za morfogenezu srčanog septuma, formiranje krvnih sudova u placenti i vaskularni integritet. U odrasloj dobi, CYR61 ima važnu ulogu u upalama i obnavljanju tkiva, a u vezi je s bolestima povezanim s hroničnom upalom, uključujući reumatoidni artritis, aterosklerozu, dijabetes povezanu nefropatiju i retinopatiju, te mnoge različite oblike kancera.

CCN porodica proteina

CYR61 je prvi put identificiran kao protein kodiran genom koji se inducira u serumu u fibroblastima miša.[6][8] Kasnije su identifikovani i drugi visoko konzervirani homolozi, koji obuhvataju porodicu proteina CCN (CCN unutarćelijski signalni protein).[9][10][11] CCN akronim je izveden od prva tri identifikovana člana porodice, CYR61 (CCN1), CTGF (faktor rasta vezivnog tkiva ili CCN2) i NOV (preeksprimirani nefroblastom ili CCN3). Ovi proteini, zajedno sa WISP1 (CCN4), WISP2 (CCN5) i WISP3 (CCN6) čine šest članova kičmenjačke porodice i preimenovani su, međunarodnim konsenzusom, u CCN1-6 po redu njihovog otkrića.[12] CCN proteini funkcionišu kao matrićelijskii proteini, proteini vanćelijskog matriksa koji imaju regulatorne uloge, posebno u kontekstu zacjeljivanja rana.[13]

Amiokiselininska sekvenca

Dužina polipeptidnog lanca je 381 aminokiselina, a molekulska težina 42.027 Da.[14]

1020304050
MSSRIARALALVVTLLHLTRLALSTCPAACHCPLEAPKCAPGVGLVRDGC
GCCKVCAKQLNEDCSKTQPCDHTKGLECNFGASSTALKGICRAQSEGRPC
EYNSRIYQNGESFQPNCKHQCTCIDGAVGCIPLCPQELSLPNLGCPNPRL
VKVTGQCCEEWVCDEDSIKDPMEDQDGLLGKELGFDASEVELTRNNELIA
VGKGSSLKRLPVFGMEPRILYNPLQGQKCIVQTTSWSQCSKTCGTGISTR
VTNDNPECRLVKETRICEVRPCGQPVYSSLKKGKKCSKTKKSPEPVRFTY
AGCLSVKKYRPKYCGSCVDGRCCTPQLTRTVKMRFRCEDGETFSKNVMMI
QSCKCNYNCPHANEAAFPFYRLFNDIHKFRD

Struktura i regulacija gena

CYR61 se nalazi na ljudskom hromosomu 1, regija p22.3, dok se gen Cyr61 miša nalazi na hromosomu 3, 72.9cM.[15] Kodirajuća regija mišjeg CYR61 prostire se na ~3,2 Kb i sadrži pet međuprostornih egzona sa četiri introna.[16] Prvi egzon kodira sekvencu 5' UTR i prvih nekoliko aminokiselina u sekretornom signalnom peptidu. Preostala četiri egzona kodiraju različit CCN1 domen. Peti egzon također sadrži sekvencu 3' UTR, koja ima pet kopija AU-bogatih elermenata koji daje kratkopoluživotnu iRNK i mir-155 ciljnu lokaciju.[17] Promotor CYR61 je TATA-kutija koja sadrži promotor, sa veznim mjestima za mnoge transkripcijske faktore uključujući AP1, ATF, E2F, HNF3b, NF1, NFκB, SP1 i SRF i dva poli(CA) dijela koji mogu formirati Z-DNK strukturu. Transkripcijska aktivacija CYR61 je izuzetno osjetljiva na širok spektar perturbacija okoline, uključujući stimulaciju trombocitni faktor rasta i osnovni faktor rasta fibroblasta, transformirajući faktor rasta β1 (TGF-β1) , hormon rasta, forbol-ester 12-O-tetradekanoilforbol-13-acetat (TPA), cAMP, vitamin D3, estrogen i tamoksifen, angiotenzin II, hipoksiju, UV svjetlo i mehaničko rastezanje.[7][11]

Proteinska struktura i funkcija

Strukturni domeni

CYR61 protein pune dužine sadrži 381 aminokiselinu sa N-terminalnim sekretornim signalnim peptidom, nakon čega slijede četiri strukturno različita domena.[18] Četiri domena CYR61 su, od N– do C-kraja, domen proteina koji vezuje faktor rasta nalik insulinu (IGFBP), von Willebrandov tip C-ponavljanja vWC domena, trombospondin tip 1 ponavljanja domena (TSR) i C-terminalni (CT) domen koji sadrži motiv cisteinskog čvora. CCN1 ima neobično visok sadržaj cisteinskih ostataka (10% ili 38 ukupno). Broj i razmak cisteinskih ostataka potpuno su konzervirano među CYR61 (CCN1), CTGF (CCN2), NOV (CCN3) i WISP-1 (CCN4), a u velikoj mjeri konzervirani su kod WISP-2 (CCN5), kojem nedostaje upravo CT domen i WISP3 (CCN6), kojem nedostaju četiri cisteina u vWC domenu. CYR61 je glikoziliran, iako su regulacija i funkcija gilkozilacije nepoznati.

Struktura proteina CCN1

Vezanje integrina

CYR61 se direktno vezuje za različite integrinske receptore na način ovisan o tipu ćelije, uključujući integrin αvβ3 u endotelnim ćelijama,[19] α6β1 i heparan-sulfatne proteoglikane (HSPGs) u fibroblastima i ćelijama glatkih mišića .[20][21] αIIbβ3 u aktiviranim trombocitima,[22] αMβ2 u monocitima i makrofagima,[23][24] i αDβ2 u pjenastim ćelijama makrofaga.[25] Kada je ispitan, sindekan-4 je identificiran kao HSPG kritičan za funkcije CCN1.[26][27] Mjesta vezanja CYR61 za neke od ovih integrina su mapirana (Slika 1). Zbog specifičnosti ćelijskog tipa ekspresije integrina, CYR61 djeluje kroz različite integrine, kako bi posredovao u specifičnim funkcijama u različitim tipovima ćelija. Na primjer, CYR61 inducira angiogene funkcije u endotelnim ćelijama putem αvβ3,[28] a u fibroblastima podstiče ćelijsko starenje i omogućava TNFα da indukuje apoptozu, vezivanjem za α6β1-HSPGs.[27][29] Međutim, CYR61 podržava ćelijsku adheziju putem svih gore identificiranih integrina.

Ćelijska signalizacija i funkcija

Kao ćelijski adhezivni supstrat, CYR61 inducira aktivaciju fokusne adhezijske kinaze, paksilina, RAC i trajnu aktivaciju MAPK/ERK1-2.[30] U makrofagima, CYR61 također aktivira transkripcijski faktor NFκB i stimulira polarizaciju M1.[24] CYR61 aktivira Akt signalizaciju u epitelnim ćelijama timusa, promovirajući njihovu proliferaciju i time rast veličine timusa.[31] CYR61 ima snažnu angiogenu aktivnost na endotelne ćelije i inducira neovaskularizaciju, što je prvi put pokazano u testu mikrodžepova implantata rožnjače [32] i naknadno potvrđeno na modelu kunićne ishemije stražnjeg mozga.[33] CYR61 također ubrzava i podstiče hondrogenu diferencijaciju mezenhimskih ćelijaa pupoljaka mišjih ekstremiteta,[34] i stimulira diferencijaciju osteoblasta, ali inhibira osteoklastogenezu.[35][36][37] Cyr61 je snažan induktor akumulacije reaktivnih vrsta kisika u fibroblastnim ćelijama, a ova aktivnost je u osnovi mnogih CYR61 induciranih apoptoza i starenja.[27][29] CYR61 je u stanju podržati ćelijsku adheziju, stimulira migraciju ćelija, promovira proliferaciju i diferencijaciju ćelija izazvanu faktorom rasta u nekim tipovima ćelija, promovira apoptozu u sinergiji sa citokinima porodice TNF i inducira ćelijsko starenje u fibroblastima.

Embrionalni razvoj

Tokom razvoja embriona kod miševa, "Cyr61" je visoko eksprimiran u kardiovaskularnom, skeletnom i neuronskom sistemu.[38][39] Cyr61 nokaut-miševa su embrionalno smrtonosni zbog defekata u morfogenezi srčanog septuma, manjkavog formiranja krvnih sudova u placenti i ugroženog vaskularnog integriteta.[40][41] Kod Xenopus laevis, Cyr61 je potreban za normalnu gastrulaciju i modulaciju Wnt-signalizaciju.[42]

Klinički značaj

CYR61 je visoko eksprimiran na mjestima upale i popravke rana, a povezan je s bolestima koje uključuju hroničnu upalu i ozljedu tkiva.[7]

Zacjeljivanje rana i fibroza

U kožnon zacjeljivanju rana, CYR61 je visoko eksprimiran u granulastom tkivu pomoću miofibroblasta, koji proliferiraju i brzo sintetiziraju ECM, kako bi održali integritet tkiva i promovirali regeneraciju parenhimskih ćelija.[43][44] Međutim, prekomjerno taloženje matriksa može dovesti do fibroze, ožiljaka i gubitka funkcije tkiva. U ranama na koži, CYR61 se akumulira u granulastom tkivu, kako miofibroblasti proliferiraju i na kraju dostiže dovoljno visok nivo da dovede do starenja same miofibroblaste, nakon čega ove ćelije prestaju da se razmnožavaju i eksprimiraju enzime koji razgrađuju matriks.[29] Dakle, CYR61 ograničava sintezu i taloženje ECM putem miofibroblasta, smanjujući rizik od fibroze tokom zarastanja rana.[45] Osim zacjeljivanja rana na koži, ekspresija CYR61 je povišena u remodeliranju kardiomiocita nakon infarkta miokarda,[46] kod vaskularnih ozljeda,[21] i kod dugih kostiju tokom sanacije prijeloma.[47][48] Blokada CYR61 antitelima inhibira zarastanje prijeloma kostiju kod miševa.[49] U bubrezima, CYR61 se eksprimira u podocitima u normalnim odraslim i embrionalnim glomerulima, ali je ekspresija smanjena u IgA-nefropatijama, dijabeteskoj i membranoznoj nefropatiji, posebno u bolesnim bubrezima s teškim mezangijnim ekspanzijama.[50] CYR61 indukcija ćelijskog starenja u bubrezima potencijalna je terapija za ograničavanje fibroze.[51]

Upala

CYR61 promovira apoptotske funkcije upalnih citokina kao što su TNFα, FasL i TRAIL.[27][52][53] Također reprogramira makrofage prema M1 polarizaciji preko αMβ2 posredovane aktivacije NF-κB.[24] CYR61 je pojačan kod pacijenata sa Crohnovom bolešću i ulceroznim kolitisom.[54] CYR61 podržava ponašanje patroliranja mišjih rezidentnih Ly6Cniskih monocita duž endotela u stabilnom stanju i potreban je za njihovu akumulaciju pod vaskularnom upalom koja oponaša virus.[55]

Artritis

CYR61 je visoko eksprimiran u artritisu izazvanom kolagenom kod glodara, a inhibicija ekspresije CCN1 korelira sa supresijom upalnog artritisa.[56] CYR61 se također nalazi u zglobnoj hrskavici pacijenata s osteoartritisom i čini se da potiskuje aktivnost ADAMTS4 (agrekanaze), što može dovesti do kloniranja ćelija hrskavica (hondrocita)..[57]

Vaskularne bolesti

CYR61 je prekomjerno eksprimiran u ćelijama vaskularnih glatkih mišića, aterosklerotskih lezija i u restenozama noointime nakon balonske angioplastike, kako kod modela glodra, tako i kod ljudi.[21][23][58][59] Supresija ekspresije CYR61 rezultira smanjenom neointimnom hiperplazijom nakon balonske angioplastike, efekt koji se poništava isporukom CYR61 putem prijenosa gena[60][61] Na mišjem modelu retinopatije izazvane kisikom, ekspresija CYR61 u staklastom tijelu dala je značajne korisne efekte u popravljanju oštećene vaskulature.[62]

Kancer

Za opskrbu kisikom i hranjivim tvarima za ishranu rastućeg tumora, neophodna je angiogeneza.[63] CYR61 je snažan angiogeni induktor in vivo,[32][33] i također može promovirati proliferaciju ćelija raka, invaziju, preživljavanje, epitelno-mezenhimalni prijelaz i metastaze.[64][65][66][67] Shodno tome, prisilna prekomjerna ekspresija CYR61 pojačala je rast tumora u ksenotransplantatnim ćelijama raka dojke,[68] prostate,[65] ćelijama raka jajnika[69] ćelijama raka pločastih ćelija.[70] Klinički, ekspresija CYR61 korelira sa stadijem i veličinom tumora, pozitivnošću limfnih čvorova i lošom prognozom kod nekoliko karcinoma, uključujući rak dojke,[68][71][72][73][74] rak prostate,[75] gliom,[76] želučani adenokarcinom[77] i karcinom skvamoznih ćelija.[70]

Međutim, CYR61 također može izazvati apoptoze[] i ćelijsko starenje,[26][29][78] dva dobro uspostavljena mehanizma supresije tumora.[79] Dakle, dok CYR61 može promovirati proliferaciju ćelija raka prostate, on također može pogoršati njihovu apoptozu u prisustvu molekula imunoskog nadzora TRAIL.[53][65][80] CYR61 ima inhibitorni efekat na neke tipove raka i potiskuje rast tumorskih ćelija nemaloćelijskog karcinoma pluća (NSCLC),[81] ćelija endometrijskog adenokarcinoma,[82] i u ćelijama melanoma.[83]

Reference

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Relationship between the Chinese People's Volunteer Army and the Korean People's Army This is the order of battle for Chinese People's Volunteer Army during major periods of hostilities in the Korean War. After the People's Republic of China entered the Korean War in October 1950 by designating the People's Liberation Army (PLA) North East Frontier Force as the People's Volunteer Army (PVA),[1] the PVA spent the next two years and nine months in combat operations and five years and th...

 

Yanomamita

Yanomamita Verde la yanomamita y blanca la escorodita.GeneralCategoría Minerales fosfatos - ArseniatosClase 8.CD.10 (Strunz)Fórmula química InAsO4·2H2OPropiedades físicasColor verde pálido, verde amarillo, azul-verdeRaya blancaLustre vítreoTransparencia semitransparenteSistema cristalino Ortorrómbico dipiramidalHábito cristalino cristales dipiramidales, típicamente en costrasExfoliación no observadaDureza 3.5–4 (Mohs)Densidad 3.87 g/cm³[editar datos en Wikidata] La yan...

Yoyes (film)

2000 filmYoyesSpanish film posterDirected byHelena TabernaWritten byHelena TabernaAndrés MartorellProduced byEnrique CerezoStarringAna TorrentErnesto AlterioFlorence PernelCinematographyFederico RibesEdited byRosario Sáinz de RozasMusic byÁngel IllarramendiRelease date 31 March 2000 (2000-03-31) (Spain) Running time104 minutesCountriesSpainFranceItalyLanguageSpanish Yoyes is a 2000 Spanish political drama film directed by Helena Taberna [es] consisting of a ...

 

Stomper the Maverick

StomperStomper the Maverick at Blakeslee StadiumUniversityMinnesota State University, MankatoConference * Northern Sun Intercollegiate Conference * Western College Hockey Association DescriptionMaverickFirst seen1977 Stomper the Maverick is the mascot for the Minnesota State Mavericks athletics teams of the Minnesota State University, Mankato and the associated club teams and charities. During the year, Stomper makes over 250 appearances and is at virtually all home games for University teams...

 

E. Brown House

Historic house in Massachusetts, United States United States historic placeE. Brown HouseU.S. National Register of Historic Places Show map of MassachusettsShow map of the United StatesLocationUxbridge, MassachusettsCoordinates42°5′24″N 71°39′0″W / 42.09000°N 71.65000°W / 42.09000; -71.65000Built1805Architectural styleFederalMPSUxbridge MRANRHP reference No.83004109 [1]Added to NRHPOctober 7, 1983 The E. Brown House is an historic hou...

Distrito 13 de Budapest

Distrito 13 de Budapest Distrito Escudo Coordenadas 47°32′11″N 19°04′19″E / 47.5364, 19.0719Capital BudapestIdioma oficial HúngaroEntidad Distrito • País  HungríaAlcalde József Tóth (MSZP)Superficie   • Total 13.43 km²Población (1 de enero de 2023)   • Total 120 256[1]​ hab. • Densidad 8,954 hab/km² Sitio web oficial [editar datos en Wikidata] El Distrito 13 de Budapest (en húngaro: Bu...

 

Columbia River Treaty

1961 agreement between Canada and the United States The Columbia River Treaty is a 1961 agreement between Canada and the United States on the development and operation of dams in the upper Columbia River basin for power and flood control benefits in both countries. Four dams were constructed under this treaty: three in the Canadian province of British Columbia (Duncan Dam, Mica Dam, Keenleyside Dam) and one in the U.S. state of Montana (Libby Dam). The treaty provided for the sharing with Can...

 
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