Izvorni kôd stranice Homeoboksni protein NANOG

{{Infokutija gen}}
'''[[Homeoboks]]ni protein NANOG''' je [[transkripcijski faktor]] koji pomaže [[Embrionska matična ćelija|embrionskim matičnim ćelijama]] (ESC) održavanje [[Ćelijska potencija|pluripotencije]] putem supresije faktora ćelijke determinacije.<ref>Heurtier, V., Owens, N., Gonzalez, I. et al. The molecular logic of Nanog-induced self-renewal in mouse embryonic stem cells. Nat Commun 10, 1109 (2019). https://doi.org/10.1038/s41467-019-09041-z</ref> Several types of cancer are associated with NANOG.

== Etimologija ==
Ime NANOG potječe od [[Tir na n-Og|Tír na nÓg]] ([[irski]] za "Zemlja mladih"), ime dato keltskom došljaku u irskoj i škotskoj mitologiji.<ref>{{cite web|url=https://www.sciencedaily.com/releases/2003/06/030602024530.htm|title=ScienceDaily: Cells Of The Ever Young: Getting Closer To The Truth|access-date=26. 7. 2007}}</ref><ref name="The role of NANOG transcriptional f">{{cite journal | vauthors = Gawlik-Rzemieniewska N, Bednarek I | title = The role of NANOG transcriptional factor in the development of malignant phenotype of cancer cells | journal = Cancer Biology & Therapy | volume = 17 | issue = 1 | pages = 1–10 | date = 30. 11. 2015 | pmid = 26618281 | pmc = 4848008 | doi = 10.1080/15384047.2015.1121348 }}</ref>

==Aminokiselinska sekvenca==
Dužina [[polipeptid]]nog lanca je 305 [[aminokiselina]], a  [[molekularna masa|molekulska težina]]   34.620 [[dalton (jedinica)|Da]].<ref name="UniProt">{{cite web|url=http://www.uniprot.org/uniprot/Q9H9S0#sequences |title=UniProt, Q9H9S0 |access-date=12. 8. 2021}}</ref>

<table  style="font-family:monospace;">
<tr style="text-align:right;"><th>10</th><th></th><th>20</th><th></th><th>30</th><th></th><th>40</th><th></th><th>50</th></tr>
<tr style="background-color: #eee;"><td>MSVDPACPQS</td><td style="background-color: #f9f9f9;"></td><td>LPCFEASDCK</td><td style="background-color: #f9f9f9;"></td><td>ESSPMPVICG</td><td style="background-color: #f9f9f9;"></td><td>PEENYPSLQM</td><td style="background-color: #f9f9f9;"></td><td>SSAEMPHTET</td></tr>
<tr style="background-color: #eee;"><td>VSPLPSSMDL</td><td style="background-color: #f9f9f9;"></td><td>LIQDSPDSST</td><td style="background-color: #f9f9f9;"></td><td>SPKGKQPTSA</td><td style="background-color: #f9f9f9;"></td><td>EKSVAKKEDK</td><td style="background-color: #f9f9f9;"></td><td>VPVKKQKTRT</td></tr>
<tr style="background-color: #eee;"><td>VFSSTQLCVL</td><td style="background-color: #f9f9f9;"></td><td>NDRFQRQKYL</td><td style="background-color: #f9f9f9;"></td><td>SLQQMQELSN</td><td style="background-color: #f9f9f9;"></td><td>ILNLSYKQVK</td><td style="background-color: #f9f9f9;"></td><td>TWFQNQRMKS</td></tr>
<tr style="background-color: #eee;"><td>KRWQKNNWPK</td><td style="background-color: #f9f9f9;"></td><td>NSNGVTQKAS</td><td style="background-color: #f9f9f9;"></td><td>APTYPSLYSS</td><td style="background-color: #f9f9f9;"></td><td>YHQGCLVNPT</td><td style="background-color: #f9f9f9;"></td><td>GNLPMWSNQT</td></tr>
<tr style="background-color: #eee;"><td>WNNSTWSNQT</td><td style="background-color: #f9f9f9;"></td><td>QNIQSWSNHS</td><td style="background-color: #f9f9f9;"></td><td>WNTQTWCTQS</td><td style="background-color: #f9f9f9;"></td><td>WNNQAWNSPF</td><td style="background-color: #f9f9f9;"></td><td>YNCGEESLQS</td></tr>
<tr style="background-color: #eee;"><td>CMQFQPNSPA</td><td style="background-color: #f9f9f9;"></td><td>SDLEAALEAA</td><td style="background-color: #f9f9f9;"></td><td>GEGLNVIQQT</td><td style="background-color: #f9f9f9;"></td><td>TRYFSTPQTM</td><td style="background-color: #f9f9f9;"></td><td>DLFLNYSMNM</td></tr>
<tr style="background-color: #eee;"><td>QPEDV</td></tr>
</table>
:Simboli
{{refbegin|4}}
'''C''':  [[Cistein]] <br>
'''D''': [[Asparaginska kiselina]] <br>
'''E''': [[Glutaminska kiselina]] <br>
'''F''':  [[Fenilalanin]] <br>
'''G''':  [[Glicin]] <br>
'''H''':  [[Histidin]] <br>
'''I''':  [[Izoleucin]] <br>
'''K''': [[Lizin]] <br>
'''L''': [[Leucin]] <br>
'''M''':  [[Metionin]] <br>
'''N''':  [[Asparagin]] <br>
'''P''':  [[Prolin]]<br>
'''Q''':  [[Glutamin]] <br>
'''R''':  [[Arginin]] <br>
'''S''':  [[Serin]] <br>
'''T''':  [[Treonin]] <br>
'''V''':  [[Valin]] <br>
'''W''':  [[Triptofan]] <br>
'''Y''':  [[Tirozin]]
{{refend}}

== Struktura ==
Ljudski protein NANOG kodiran genom NANOG1, sastoji se od 305 aminokiselina i ina tri funkcionalna domena: [[N-terminal]]ni, [[C-terminal]]ni  i konzervirani [[homeodomen]]ski motiv. [[Homeodomen]]ska regija olakšava vezivanje [[DNK]]. Ljudski gen Nanog 1 nalazi se na [[hromosom 12|hromosomu 12]], a [[iRNK]] sadrži 915 baznih parova [[otvoreni okvir čitanja|otvorenog okvira čitanja]] (ORF) sa četiri [[egzon]]a i tri [[intron]]a.<ref name="The role of NANOG transcriptional f"/>

[[N-terminal]]na regija ljudskog NANOG-a bogata je ostacima [[serin]]a, [[treonin]]a i [[prolin]]a, a [[C-kraj]] sadrži domen bogat [[triptofan]]om. Homeodomen u hNANOG-u kreće se od 95 do 155 ostataka. Postoje i dodatni NANOG geni (NANOG2, NANOG p8) koji potencijalno utiču na diferencijaciju ESC -a. Pokazano je da je NANOG1 temelj za samoobnavljanje i pluripotenciju, a NANOG p8 je visoko eksprimirann u ćelijama raka.<ref name="Nanog gene: A propeller for stemnes">{{cite journal | vauthors = Zhang W, Sui Y, Ni J, Yang T | title = Nanog gene: A propeller for stemness in primitive stem cells | journal = International Journal of Biological Sciences | volume = 12 | issue = 11 | pages = 1372–1381 | date = 2016 | pmid = 27877089 | pmc = 5118783 | doi = 10.7150/ijbs.16349 }}</ref>

== Funkcija ==
[[slika:Transcription programs in stem cells..jpg|280px|thumb|Transkripcijski programi u embrionskim matičnim ćelijama]]
NANOG je [[transkripcijski faktor]] u embrionskim matičnim ćelijama (ESC) i smatra se da je ključni faktor u održavanju [[pluripotentnost|pluripotencije]]. Smatra se da NANOG funkcionira u skladu s drugim faktorima, kao što su [[okt-4|POU5F1 (okt-4)]] i [[SOX2]], radi uspostavljanja ESC identiteta. Ove ćelije nude važno područje proučavanja, zbog svoje sposobnosti održavanja pluripotencije. Drugim riječima, ove ćelije imaju sposobnost da postanu gotovo bilo koja ćelija bilo kojeg od tri zametna sloja ([[endoderm]], [[ektoderm]], [[mezoderm]]). Iz tog razloga je razumijevanje mehanizama koji održavaju pluripotenciju ćelije ključno da se razumije kako funkcioniraju [[matične ćelije]] i može dovesti do budućeg napretka u liječenju degenerativnih bolesti.

Opisano je da se NANOG na početku [[gastrulacija|gastrulacije]] eksprimira u stražnjoj strani epiblasta.<ref name = "Barral_2009">{{cite journal | vauthors = Barral A, Rollan I, Sanchez-Iranzo H, Jawaid W, Badia-Careaga C, Menchero S, Gomez MJ, Torroja C, Sanchez-Cabo F, Göttgens B, Manzanares M, Sainz de Aja J | display-authors = 6 | title = Pou3f1 expression at the exit from pluripotency during gastrulation | journal = Biology Open | volume = 8 | issue = 11 | pages = bio046367 | date = decembar 2019 | pmid = 31791948 | doi = 10.1242/bio.046367 | pmc = 6899006 | doi-access = free }}</ref> Tamo je NANOG  umiješan u inhibiranje embrionske [[hematopoeza|hematopoeze]] potiskivanjem ekspresije transkripcijskog faktora ''Tal1''.<ref>{{cite journal | vauthors = Sainz de Aja J, Menchero S, Rollan I, Barral A, Tiana M, Jawaid W, Cossio I, Alvarez A, Carreño-Tarragona G, Badia-Careaga C, Nichols J, Göttgens B, Isern J, Manzanares M | display-authors = 6 | title = Tal1 | journal = The EMBO Journal | volume = 38 | issue = 7 | date = april 2019 | pmid = 30814124 | pmc = 6443201 | doi = 10.15252/embj.201899122 }}</ref> U ovoj  fazi embriogeneze, NANOG potiskuje '' Pou3f1 '', [[transkripcijski faktor]] ključan za formiranje prednje-stražnje osi.<ref name = "Barral_2009"/>

Analiza usporenih embriona pokazala je da eksprimiraju gene markera pluripotencije, kao što su [[Oct-4|POU5F1]], NANOG i [[Rex1]]. Izvedene ljudske ESC linije također eksprimiraju specifične markere pluripotencije:
* TRA-1-60
* TRA-1-81
* SSEA4
alkalna fosfataza
* TERT
* [[Rex1]]

Ovi markeri su omogućili diferencijaciju stanja ''[[in vitro]]'' i ''[[in vivo]]'' u derivate sva tri zametna sloja.<ref name="pmid16990582">{{cite journal | vauthors = Zhang X, Stojkovic P, Przyborski S, Cooke M, Armstrong L, Lako M, Stojkovic M | title = Derivation of human embryonic stem cells from developing and arrested embryos | journal = Stem Cells | volume = 24 | issue = 12 | pages = 2669–76 | date = decembar 2006 | pmid = 16990582 | doi = 10.1634/stemcells.2006-0377 | s2cid = 32587408 | doi-access = free }}</ref>

[[Oct-4|POU5F1]], TDGF1 (CRIPTO), [[SALL4]], LECT1 i BUB1 su također povezani geni koji su svi odgovorni za samoobnavljanje i pluripotentnu diferencijaciju.<ref name="pmid16978057">{{cite journal | vauthors = Li SS, Liu YH, Tseng CN, Chung TL, Lee TY, Singh S | title = Characterization and gene expression profiling of five new human embryonic stem cell lines derived in Taiwan | journal = Stem Cells and Development | volume = 15 | issue = 4 | pages = 532–55 | date = august 2006 | pmid = 16978057 | doi = 10.1089/scd.2006.15.532 }}</ref>

Utvrđeno je da je protein NANOG transkripcijski aktivator promotora [[Rex1]], koji ima ključnu ulogu u održavanju ekspresije [[Rex1]]. Nokaut NANOG  u [[embrionske matične ćelije|embrionskim matičnim ćelijama]]  rezultira smanjenjem ekspresije [[Rex1]], dok nadekspresija NANOG -a stimulira ekspresiju [[Rex1]].<ref>{{cite journal | vauthors = Shi W, Wang H, Pan G, Geng Y, Guo Y, Pei D | title = Regulation of the pluripotency marker Rex-1 by Nanog and Sox2 | journal = The Journal of Biological Chemistry | volume = 281 | issue = 33 | pages = 23319–25 | date = august 2006 | pmid = 16714766 | doi = 10.1074/jbc.M601811200 | doi-access = free }}</ref>

Osim učinaka NANOG-a u embrionskim fazama života, ektopijska ekspresija NANOG-a u odraslim matičnim ćelijama može vratiti potencijal proliferacije i diferencijacije koji se gubi uslijed starenja organizma ili ćelijskog starenja.<ref>{{cite journal | vauthors = Shahini A, Choudhury D, Asmani M, Zhao R, Lei P, Andreadis ST | title = NANOG restores the impaired myogenic differentiation potential of skeletal myoblasts after multiple population doublings | journal = Stem Cell Research | volume = 26 | pages = 55–66 | date = januar 2018 | pmid = 29245050 | doi = 10.1016/j.scr.2017.11.018 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Shahini A, Mistriotis P, Asmani M, Zhao R, Andreadis ST | title = NANOG Restores Contractility of Mesenchymal Stem Cell-Based Senescent Microtissues | journal = Tissue Engineering. Part A | volume = 23 | issue = 11–12 | pages = 535–545 | date = juni 2017 | pmid = 28125933 | pmc =5467120  | doi = 10.1089/ten.TEA.2016.0494 }}</ref><ref>{{cite journal | vauthors = Mistriotis P, Bajpai VK, Wang X, Rong N, Shahini A, Asmani M, Liang MS, Wang J, Lei P, Liu S, Zhao R, Andreadis ST | title = NANOG Reverses the Myogenic Differentiation Potential of Senescent Stem Cells by Restoring ACTIN Filamentous Organization and SRF-Dependent Gene Expression | journal = Stem Cells | volume = 35 | issue = 1 | pages = 207–221 | date = januar 2017 | pmid = 27350449 | doi = 10.1002/stem.2452 | s2cid = 4482665 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Han J, Mistriotis P, Lei P, Wang D, Liu S, Andreadis ST | title = Nanog reverses the effects of organismal aging on mesenchymal stem cell proliferation and myogenic differentiation potential | journal = Stem Cells | volume = 30 | issue = 12 | pages = 2746–59 | date = decembar 2012 | pmid = 22949105 | pmc = 3508087 | doi = 10.1002/stem.1223 }}</ref><ref>{{cite journal | vauthors = Münst B, Thier MC, Winnemöller D, Helfen M, Thummer RP, Edenhofer F | title = Nanog induces suppression of senescence through downregulation of p27KIP1 expression | journal = Journal of Cell Science | volume = 129 | issue = 5 | pages = 912–20 | date = mart 2016 | pmid = 26795560 | pmc = 4813312 | doi = 10.1242/jcs.167932 }}</ref>

== Klinički značaj ==

=== Kancer===

NANOG je visoko eksprimiran u matičnim ćelijama raka i stoga može funkcionirati kao [[onkogen]] za poticanje karcinogeneze. Visoka ekspresija NANOG -a korelira sa lošim preživljavanjem kod pacijenata obolelih od raka.<ref name="pmid26013997">{{cite journal | vauthors = Gong S, Li Q, Jeter CR, Fan Q, Tang DG, Liu B | title = Regulation of NANOG in cancer cells | journal = Molecular Carcinogenesis | volume = 54 | issue = 9 | pages = 679–87 | date = septembar 2015 | pmid = 26013997 | pmc = 4536084 | doi = 10.1002/mc.22340 }}</ref><ref name="pmid25821200">{{cite journal | vauthors = Jeter CR, Yang T, Wang J, Chao HP, Tang DG | title = Concise Review: NANOG in Cancer Stem Cells and Tumor Development: An Update and Outstanding Questions | journal = Stem Cells | volume = 33 | issue = 8 | pages = 2381–90 | date = august 2015 | pmid = 25821200 | pmc = 4509798 | doi = 10.1002/stem.2007 }}</ref><ref name="pmid26618281">{{cite journal | vauthors = Gawlik-Rzemieniewska N, Bednarek I | title = The role of NANOG transcriptional factor in the development of malignant phenotype of cancer cells | journal = Cancer Biology & Therapy | volume = 17 | issue = 1 | pages = 1–10 | year = 2016 | pmid = 26618281 | pmc = 4848008 | doi = 10.1080/15384047.2015.1121348 }}</ref>

Nedavna istraživanja pokazala su da lokalizacija NANOG -a i drugih transkripcijskih faktora ima potencijalne posljedice na ćelijsku funkciju. Eksperimentalni dokazi su pokazali da je razina ekspresije NANOG p8 posebno povišena u ćelijama raka, što znači da je gen NANOG p8 kritični član u (CSC) matičnim ćelijama raka, pa bi njegovo poništavanje moglo smanjiti malignitet raka.<ref name="Nanog gene: A propeller for stemnes"/>

=== Dijagnostika ===

Gen NANOG p8 ocijenjen je kao prognostički i prediktivni biomarker raka.<ref name="pmid24375318">{{cite journal | vauthors = Iv Santaliz-Ruiz LE, Xie X, Old M, Teknos TN, Pan Q | title = Emerging role of nanog in tumorigenesis and cancer stem cells | journal = International Journal of Cancer | volume = 135 | issue = 12 | pages = 2741–8 | date = decembar 2014 | pmid = 24375318 | pmc = 4065638 | doi = 10.1002/ijc.28690 }}</ref>

== Matične ćelije raka ==

Nanog je [[transkripcijski faktor]] koji kontrolira i samoobnavljanje i [[pluripotentnost]] embrionskih matičnih čelija. Slično, ekspresija proteina porodice Nanog povećana je kod mnogih tipova karcinoma i korelira s lošijom prognozom.<ref name="Nanog gene: A propeller for stemnes"/>

== Evolucija ==
[[Ljudi]] i [[čimpanze|obične čimpanze]] dijele deset NANOG [[pseudogen]]a (NanogP2-P11) tokom evaluacije, dva se nalaze na X hromosomu i karakterišu ih 5' promotorske sekvence i odsustvo introna, kao posljedica retrotranspozicije [[iRNK]] <ref name="The role of NANOG transcriptional f"/> sve na istim mjestima: jedan duplicirani pseudogen i devet retropseudogena. Od devet zajedničkih NANOG retropseudogena, dva nemaju [[poliadenilacija|poli- (A) repove]] karakteristične za većinu retropseudogena, što ukazuje na to da je došlo do grešaka pri kopiranju tokom njihovog stvaranja. Zbog velike vjerovatnoće da bi isti pseudogeni (uključujući greške pri kopiranju) postojali na istim mjestima u dva nepovezana [[genom]]a, [[Evolucijska biologija|evolucijski biolozi]] ukazuju na to da NANOG i njegovi pseudogeni pružaju [[dokaz zajedničkog porijekla]]  ljudi  čimpanzi.<ref name="isbn1-59102-564-83">{{cite book|title=Relics of Eden: The Powerful Evidence of Evolution in Human DNA| first = Daniel J. | last = Fairbanks | name-list-style = vanc |publisher=Prometheus Books|year=2007|isbn=978-1-59102-564-1|location=Buffalo, N.Y|pages=94–96, 177–182}}</ref>

== Također pogledajte ==
* [[Histon]]
* [[Okt-4]]
* [[promotor (genetika)|Promotor]]
* [[RNK-polimeraza]]
* [[Brahiurija]]
* [[Transkripcijski faktori]]
*[[Bioinformatika]]

== Reference ==
{{reflist|30em}}

== Dopunska literatura ==
{{refbegin | 2}}
* {{cite journal | vauthors = Cavaleri F, Schöler HR | title = Nanog: a new recruit to the embryonic stem cell orchestra | journal = Cell | volume = 113 | issue = 5 | pages = 551–2 | date = maj 2003 | pmid = 12787492 | doi = 10.1016/S0092-8674(03)00394-5 | s2cid = 16254995 | doi-access = free }}
* {{cite journal | vauthors = Constantinescu S | title = Stemness, fusion and renewal of hematopoietic and embryonic stem cells | journal = Journal of Cellular and Molecular Medicine | volume = 7 | issue = 2 | pages = 103–12 | year = 2004 | pmid = 12927049 | doi = 10.1111/j.1582-4934.2003.tb00209.x | pmc = 6740230 }}
* {{cite journal | vauthors = Pan G, Thomson JA | title = Nanog and transcriptional networks in embryonic stem cell pluripotency | journal = Cell Research | volume = 17 | issue = 1 | pages = 42–9 | date = januar 2007 | pmid = 17211451 | doi = 10.1038/sj.cr.7310125 | doi-access = free }}
* {{cite journal | vauthors = Mitsui K, Tokuzawa Y, Itoh H, Segawa K, Murakami M, Takahashi K, Maruyama M, Maeda M, Yamanaka S | title = The homeoprotein Nanog is required for maintenance of pluripotency in mouse epiblast and ES cells | journal = Cell | volume = 113 | issue = 5 | pages = 631–42 | date = maj 2003 | pmid = 12787504 | doi = 10.1016/S0092-8674(03)00393-3 | s2cid = 18836242 | doi-access = free }}
* {{cite journal | vauthors = Chambers I, Colby D, Robertson M, Nichols J, Lee S, Tweedie S, Smith A | title = Functional expression cloning of Nanog, a pluripotency sustaining factor in embryonic stem cells | journal = Cell | volume = 113 | issue = 5 | pages = 643–55 | date = maj 2003 | pmid = 12787505 | doi = 10.1016/S0092-8674(03)00392-1 | hdl = 1842/843 | s2cid = 2236779 | hdl-access = free }}
* {{cite journal | vauthors = Clark AT, Rodriguez RT, Bodnar MS, Abeyta MJ, Cedars MI, Turek PJ, Firpo MT, Reijo Pera RA | title = Human STELLAR, NANOG, and GDF3 genes are expressed in pluripotent cells and map to chromosome 12p13, a hotspot for teratocarcinoma | journal = Stem Cells | volume = 22 | issue = 2 | pages = 169–79 | year = 2004 | pmid = 14990856 | doi = 10.1634/stemcells.22-2-169 | s2cid = 38136098 | doi-access = free }}
* {{cite journal | vauthors = Hart AH, Hartley L, Ibrahim M, Robb L | title = Identification, cloning and expression analysis of the pluripotency promoting Nanog genes in mouse and human | journal = Developmental Dynamics | volume = 230 | issue = 1 | pages = 187–98 | date = maj 2004 | pmid = 15108323 | doi = 10.1002/dvdy.20034 | s2cid = 21502533 | doi-access = free }}
* {{cite journal | vauthors = Booth HA, Holland PW | title = Eleven daughters of NANOG | journal = Genomics | volume = 84 | issue = 2 | pages = 229–38 | date = august 2004 | pmid = 15233988 | doi = 10.1016/j.ygeno.2004.02.014 }}
* {{cite journal | vauthors = Hatano SY, Tada M, Kimura H, Yamaguchi S, Kono T, Nakano T, Suemori H, Nakatsuji N, Tada T | title = Pluripotential competence of cells associated with Nanog activity | journal = Mechanisms of Development | volume = 122 | issue = 1 | pages = 67–79 | date = januar 2005 | pmid = 15582778 | doi = 10.1016/j.mod.2004.08.008 | hdl = 2433/144766 | s2cid = 17490059 | hdl-access = free }}
* {{cite journal | vauthors = Deb-Rinker P, Ly D, Jezierski A, Sikorska M, Walker PR | title = Sequential DNA methylation of the Nanog and Oct-4 upstream regions in human NT2 cells during neuronal differentiation | journal = The Journal of Biological Chemistry | volume = 280 | issue = 8 | pages = 6257–60 | date = februar 2005 | pmid = 15615706 | doi = 10.1074/jbc.C400479200 | doi-access = free }}
* {{cite journal | vauthors = Zaehres H, Lensch MW, Daheron L, Stewart SA, Itskovitz-Eldor J, Daley GQ | title = High-efficiency RNA interference in human embryonic stem cells | journal = Stem Cells | volume = 23 | issue = 3 | pages = 299–305 | date = mart 2005 | pmid = 15749924 | doi = 10.1634/stemcells.2004-0252 | s2cid = 1395518 | doi-access = free }}
* {{cite journal | vauthors = Hoei-Hansen CE, Almstrup K, Nielsen JE, Brask Sonne S, Graem N, Skakkebaek NE, Leffers H, Rajpert-De Meyts E | title = Stem cell pluripotency factor NANOG is expressed in human fetal gonocytes, testicular carcinoma in situ and germ cell tumours | journal = Histopathology | volume = 47 | issue = 1 | pages = 48–56 | date = juli 2005 | pmid = 15982323 | doi = 10.1111/j.1365-2559.2005.02182.x | s2cid = 10164525 }}
* {{cite journal | vauthors = Hyslop L, Stojkovic M, Armstrong L, Walter T, Stojkovic P, Przyborski S, Herbert M, Murdoch A, Strachan T, Lako M | title = Downregulation of NANOG induces differentiation of human embryonic stem cells to extraembryonic lineages | journal = Stem Cells | volume = 23 | issue = 8 | pages = 1035–43 | date = septembar 2005 | pmid = 15983365 | doi = 10.1634/stemcells.2005-0080 | s2cid = 29881293 }}
* {{cite journal | vauthors = Oh JH, Do HJ, Yang HM, Moon SY, Cha KY, Chung HM, Kim JH | title = Identification of a putative transactivation domain in human Nanog | journal = Experimental & Molecular Medicine | volume = 37 | issue = 3 | pages = 250–4 | date = juni 2005 | pmid = 16000880 | doi = 10.1038/emm.2005.33 | doi-access = free }}
* {{cite journal | vauthors = Boyer LA, Lee TI, Cole MF, Johnstone SE, Levine SS, Zucker JP, Guenther MG, Kumar RM, Murray HL, Jenner RG, Gifford DK, Melton DA, Jaenisch R, Young RA | title = Core transcriptional regulatory circuitry in human embryonic stem cells | journal = Cell | volume = 122 | issue = 6 | pages = 947–56 | date = septembar 2005 | pmid = 16153702 | pmc = 3006442 | doi = 10.1016/j.cell.2005.08.020 }}
* {{cite journal | vauthors = Kim JS, Kim J, Kim BS, Chung HY, Lee YY, Park CS, Lee YS, Lee YH, Chung IY | title = Identification and functional characterization of an alternative splice variant within the fourth exon of human nanog | journal = Experimental & Molecular Medicine | volume = 37 | issue = 6 | pages = 601–7 | date = decembar 2005 | pmid = 16391521 | doi = 10.1038/emm.2005.73 | doi-access = free }}
* {{cite journal | vauthors = Darr H, Mayshar Y, Benvenisty N | title = Overexpression of NANOG in human ES cells enables feeder-free growth while inducing primitive ectoderm features | journal = Development | volume = 133 | issue = 6 | pages = 1193–201 | date = mart 2006 | pmid = 16501172 | doi = 10.1242/dev.02286 | doi-access = free }}
* {{cite journal | vauthors = Saunders A, Li D, Faiola F, Huang X, Fidalgo M, Guallar D, Ding J, Yang F, Xu Y, Zhou H, Wang J | title = Context-Dependent Functions of NANOG Phosphorylation in Pluripotency and Reprogramming | journal = Stem Cell Reports | volume = 8 | issue = 5 | pages = 1115–1123 | date = maj 2017 | pmid = 28457890 | doi = 10.1016/j.stemcr.2017.03.023 | pmc = 5425684 }}
{{refend}}

== Vanjski linkovi ==
* {{MeshName|NANOG+protein,+human}}
* {{MeshName|Nanog+protein,+mouse}}
* https://www.factorbook.org/tf/human/NANOG/function
* {{cite web| url =http://jura.wi.mit.edu/young_public/hESregulation/index.html| title =Core Transcriptional Regulatory Circuitry in Human Embryonic Stem Cells| work =Young Lab| publisher =Whitehead Institute for Biomedical Research| archive-url =https://web.archive.org/web/20090628185523/http://jura.wi.mit.edu/young_public/hESregulation/index.html| archive-date =28. 6. 2009| access-date =28. 2. 2009| url-status =dead}}
* {{cite web | url = http://www.wi.mit.edu/research/summaries/jaenisch.html | title = Jaenisch Lab Research Summary | publisher = Whitehead Institute | access-date = 28. 2. 2009 | archive-date = 26. 6. 2012 | archive-url = https://web.archive.org/web/20120626025323/http://www.wi.mit.edu/research/summaries/jaenisch.html | url-status = dead }}
* [https://web.archive.org/web/20130525111306/http://www.wellcome.ac.uk/News/2009/News/WTX056296.htm Discovery reveals more about stem cells' immortality]

{{Transkripcijski faktori|g3}}

[[Kategorija:Ekspresija gena]]
[[Kategorija:Onkologija]]
[[Kategorija:Faktori transkripcije]]
[[Kategorija:Geni na hromosomu 12]]
[[Kategorija:Ljudski proteini]]
[[Kategorija:Proteomika]]