Proliferirajući ćelijski jedarni antigen
Proliferirajući ćelijski jedarni antigen (PCNA) je DNK-stezaljka koja djeluje kao procesivni faktor za DNK-polimerazu δ u eukariotske ćelije i neophodan je za replikaciju. PCNA je homotrimer i svoju procesnost postiže okružujući DNK, gdje djeluje kao skela za regrutovanje proteina uključenih u replikaciju DNK, popravak DNK, remodeliranje hromatina i epigenetičke uticaje. Kod ljudi, genski lokus PCNA nalazi se na kratkom (p) kraku hromosoma 20.[5]
Mnogi proteini stupaju u interakciju sa PCNA preko dva poznata PCNA-interagirajuća motiva PCNA-interaktivne peptidne (PIP) kutije [6] i AlkB homolog 2 PCNA motiva u interakciji (APIM).[7] Proteini koji se vezuju za PCNA preko PIP-kutije su uglavnom uključeni u replikaciju DNK, dok su proteini koji se vezuju za PCNA preko APIM-a uglavnom važni u kontekstu genotoksičnog stresa.[8]
Funkcija
[uredi | uredi izvor]Protein koji je kodiran ovim genom nalazi se u jedru i kofaktor je delta DNK-polimeraza. Kodirani protein djeluje kao homotrimer i pomaže u povećanju procesivnosti sinteze vodećeg lanca tokom replikacije DNK. Kao odgovor na oštećenje DNK, ovaj protein je ubikvitinski uključen, kao i u RAD6-ovisni put popravk DNK. Za ovaj gen su pronađene dvije varijante transkripta koje kodiraju isti protein. Pseudogeni ovog gena su opisani na hromosomu 4 i na hromosomu X.[9]
PCNA se također nalazi u arhejama, kao faktor procesivnosti polD, jedinstvene multifunkcionalne DNK-polimeraze u ovoj domeni života.[10]
Ekspresija u jedru tokom sinteze DNK
[uredi | uredi izvor]PCNA je prvobitno identifikovana kao antigen koji se eksprimira u ćelijskom jedru tokom faze sinteze DNK ćelijskog ciklusa. Dio proteina je sekvenciran i ta sekvenca je korištena da se omogući izolacija klona cDNK.[11] PCNA pomaže u držanju DNK-polimeraze delta (Pol δ) na DNK. PCNA je stegnut [12][13] koji je heteropentamerni član AAA+ klase ATPaza. Ekspresija PCNA je pod kontrolom kompleksa koji sadrže E2F transkripcijski faktor.[14][15]
Uloga u popravku DNK
[uredi | uredi izvor]Budući da je DNK-polimeraza epsilon uključena u resintezu izrezanih oštećenih lanaca DNK tokom popravka DNK, PCNA je važna i za sintezu DNK i za popravak DNK.[16][17]
PCNA je također uključen u put tolerancije oštećenja DNK poznat kao replikacijski popravak (PRR).[18] U PRR postoje dva potputa: (1) put translezije, koji se provodi specijaliziranim DNK-polimerazama koje su sposobne da inkorporiraju oštećene baze DNK u svoja aktivna mjesta (za razliku od normalne replikativne polimeraze, koja se zaustavlja), i time zaobilaze oštećenje, i (2) predloženi put "prebacivanja šablona" za koji se smatra da uključuje premoštavanje oštećenja angažovanjem mehanizma za homolognu rekombinaciju. PCNA je ključan za aktivaciju ovih puteva i izbor koji put će ćelija koristiti. PCNA postaje posttranslacijski modificiran pomoću ubikvitina.[19] Monoubikvitin lizina broj 164 na PCNA aktivira put sinteze translezije. Smatra se da proširenje ovog monoubikvitina pomoću nekanonskog lanca poliubikvitina vezanog uz lizin-63 na PCNA[19] aktivira put prebacivanja šablona. Nadalje, sumoilacija (pomoću malog modifikatora sličnog ubikvitinu, SUMO) PCNA lizina-164 (i u manjoj mjeri, lizina-127) inhibira put prebacivanja šablona.[19] Ovaj antagonistički efekt nastaje jer sumoilirana PCNA regrutuje DNK-helikazu zvanu Srs2,[20] koja ima ulogu u razbijanju Rad51 nukleoproteinskih filamenata koji su fundamentalni za iniciranje homologne rekombinacije.
PCNA-vezujući proteini
[uredi | uredi izvor]PCNA stupa u interakciju s mnogim proteinima.[21]
- Apoptotski faktori
- ATPaza
- Enzimi popravka ekscizijom baza
- Regulatori ćelijskog ciklusa
- Faktor hromatinskog remodeliranja
- Utovarivač stezaljke
- Kohezin
- DNK-ligaza
- DNK-metiltransferaza
- DNK-polimeraza
- E2 SUMO-konjugirajući enzim
- E3 ubikvitin-ligaza
- Flap endonukleaza
- Helikaze
- Histon-acetiltransferaza
- Histonski šaperon
- Histon-deacetilaza
- Enzimi Popravak neusklađenosti
- Receptor NKp44
- Enzim popravka ekscizijom nukleotida
- Poly ADP riboza polimeraza
- Prokaspaze[22]
- Protein-kinaza
- Faktor licenciranja
- TCP proteinski domen
- Topoizomeraza
Interakcije
[uredi | uredi izvor]Pokazalo se da PCNA interraguje sa:
- Annexin A2,[23]
- CAF-1,[24][25][26]
- CDC25C,[27]
- CHTF18,[23]
- Ciklin D1,[28][29]
- Ciklin O,[23][30]
- Ciklin-ovisna kinaza 4,[29][31]
- Ciklin-ovisni inhibitor kinaze 1C,[32]
- DNMT1,[33][34][35]
- EP300,[36]
- Uspostava kohezije sestrinskih hromatida 2,[37]
- Zakrilna strukturno-specifična endonukleaza 1,[38][39][40][41][42][43][44]
- GADD45A,[45][46][47][48][49]
- GADD45G,[50][51]
- HDAC1,[52]
- HUS1,[53]
- ING1,[54]
- KCTD13,[55]
- KIAA0101,[44]
- Ku70,[23][56]
- Ku80,[23][56][57]
- MCL1,[58]
- MSH3,[23][59][60]
- MSH6,[23][59][60]
- MUTYH,[61]
- P21,[32][40][44][62][63][64][65][66]
- POLD2,[67]
- POLD3,[23][68]
- POLDIP2,[69]
- POLH,[70]
- POLL,[71][72][73]
- RFC1,[23][62][74][75][76]
- RFC2,[23][77][78]
- RFC3,[23][79]
- RFC4,[23][77]
- RFC5,[23][75][77]
- Ubikvitin C[80][81][82]
- ATP-ovisna helikaza Wernerovog sindroma,[83][84]
- XRCC1,[85] i
- Y-kutijski vezani protein 1.[86]
Proteini koji komuniciraju sa PCNA preko APIM-a uključuju ljudski AlkB homolog 2, TFIIS-L, TFII-I, Rad51B,[7] XPA,[87] ZRANB3,[88] and FBH1.[89]
Upotreba
[uredi | uredi izvor]Antitijela protiv proliferirajućeg ćelijskog jedarnog antigena (PCNA) ili monoklonsko antitijelo zvano Ki-67 može se koristiti za procjenu različitih neoplazmi, npr. astrocitoma. Mogu imati dijagnostičku i prognostičku vrijednost. Snimanje jedarne distribucije PCNA (putem označavanja antitijela) može se koristiti za razlikovanje između ranog, srednje i kasne S-faze ćelijskog ciklusa.[90] Međutim, važno ograničenje antitijela je da ćelije moraju biti fiksirane što dovodi do potencijalnih artefakata.
S druge strane, proučavanje dinamike replikacije i popravke u živim ćelijama može se obaviti uvođenjem translacijskih fuzija PCNA. Da bi se eliminisala potreba za transfekcijama i zaobišao problem ćelija koje je teško transficirati i/ili kratko žive, mogu se koristiti markeri za replikaciju i/ili reparaciju permeabilnih ćelija. Ovi peptidi nude izrazitu prednost koja se može koristiti "in situ" u živom tkivu i čak razlikovati ćelije koje su podvrgnute replikaciji od ćelija koje su podvrgnute popravljanju..[91]
PCNA je potencijalna terapijska meta u terapiji kancera.[92]
Također pogledajte
[uredi | uredi izvor]- Ki-67 – ćelijski marker za proliferaciju
- Transkripcija
Reference
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Vanjski linkovi
[uredi | uredi izvor]- PCNA na US National Library of Medicine Medical Subject Headings (MeSH)
- "ANA: Cell cycle related (Mitotic): PCNA type 1 and type 2 Antibody Patterns". Antibody Patterns.com. Arhivirano s originala, 23. 9. 2015. Pristupljeno 15. 4. 2008.
- Dan Krotz. "Structure of a clamp–loader complex". Advanced Light Source News. Lawrence Berkeley National Laboratory. Pristupljeno 15. 4. 2008.
- "Movie showing a model of clamp loading of PCNA onto DNA". Pubmed Central.[trajno mrtav link]
- P12004