Homeoboksni protein NANOG

NANOG
Dostupne strukture
PDB	Pretraga ortologa: PDBe RCSB
Spisak PDB ID kodova
	2KT0
Identifikatori
Aliasi	NANOG
Vanjski ID-jevi	OMIM: 607937 MGI: 1919200 HomoloGene: 78027 GeneCards: NANOG
Lokacija gena (čovjek)
Hrom.	Hromosom 12 (čovjek)
Kraj	7,799,146 bp
Lokacija gena (miš)
Hrom.	Hromosom 6 (miš)
Kraj	122,691,592 bp
Obrazac RNK ekspresije
	Više referentnih podataka o ekspresiji
Ontologija gena
Molekularna funkcija	• GO:0001131, GO:0001151, GO:0001130, GO:0001204 DNA-binding transcription factor activity; • vezivanje sa DNK; • sequence-specific DNA binding; • GO:0001948, GO:0016582 vezivanje za proteine; • GO:0001106 transcription corepressor activity; • RNA polymerase II transcription regulatory region sequence-specific DNA binding; • GO:0001077, GO:0001212, GO:0001213, GO:0001211, GO:0001205 DNA-binding transcription activator activity, RNA polymerase II-specific; • GO:0001200, GO:0001133, GO:0001201 DNA-binding transcription factor activity, RNA polymerase II-specific; • GO:0000980 RNA polymerase II cis-regulatory region sequence-specific DNA binding;
Ćelijska komponenta	• Jedarce; • jedro; • nukleoplazma;
Biološki proces	• Regulacija ekspresije gena; • regulation of cell differentiation; • multicellular organism development; • Ćelijska diferencijacija; • Ćelijska proliferacija; • GO:0009373 regulation of transcription, DNA-templated; • somatic stem cell population maintenance; • transcription, DNA-templated; • GO:0003257, GO:0010735, GO:1901228, GO:1900622, GO:1904488 positive regulation of transcription by RNA polymerase II; • endodermal cell fate specification; • negative regulation of nucleic acid-templated transcription; • stem cell population maintenance; • transcription by RNA polymerase II; • cytokine-mediated signaling pathway; • GO:0044324, GO:0003256, GO:1901213, GO:0046019, GO:0046020, GO:1900094, GO:0061216, GO:0060994, GO:1902064, GO:0003258, GO:0072212 regulation of transcription by RNA polymerase II; • positive regulation of stem cell proliferation;
	Izvori:Amigo / QuickGO
Ortolozi
	79923
	71950
	ENSG00000111704
	ENSMUSG00000012396
	Q9H9S0
	Q80Z64
	NM_001297698; NM_024865
	NM_028016; NM_001289828; NM_001289830; NM_001289831
	NP_001284627; NP_079141
	NP_001276757; NP_001276759; NP_001276760; NP_082292
	Wikipodaci
Pogledaj/uredi – čovjek	Pogledaj/uredi – miš

Homeoboksni protein NANOG je transkripcijski faktor koji pomaže embrionskim matičnim ćelijama (ESC) održavanje pluripotencije putem supresije faktora ćelijke determinacije.^[5] Several types of cancer are associated with NANOG.

Etimologija

Ime NANOG potječe od Tír na nÓg (irski za "Zemlja mladih"), ime dato keltskom došljaku u irskoj i škotskoj mitologiji.^[6]^[7]

Aminokiselinska sekvenca

Dužina polipeptidnog lanca je 305 aminokiselina, a molekulska težina 34.620 Da.^[8]

10	20	30	40	50
MSVDPACPQS	LPCFEASDCK	ESSPMPVICG	PEENYPSLQM	SSAEMPHTET
VSPLPSSMDL	LIQDSPDSST	SPKGKQPTSA	EKSVAKKEDK	VPVKKQKTRT
VFSSTQLCVL	NDRFQRQKYL	SLQQMQELSN	ILNLSYKQVK	TWFQNQRMKS
KRWQKNNWPK	NSNGVTQKAS	APTYPSLYSS	YHQGCLVNPT	GNLPMWSNQT
WNNSTWSNQT	QNIQSWSNHS	WNTQTWCTQS	WNNQAWNSPF	YNCGEESLQS
CMQFQPNSPA	SDLEAALEAA	GEGLNVIQQT	TRYFSTPQTM	DLFLNYSMNM
QPEDV

Simboli

C: Cistein
D: Asparaginska kiselina
E: Glutaminska kiselina
F: Fenilalanin
G: Glicin
H: Histidin
I: Izoleucin
K: Lizin
L: Leucin
M: Metionin
N: Asparagin
P: Prolin
Q: Glutamin
R: Arginin
S: Serin
T: Treonin
V: Valin
W: Triptofan
Y: Tirozin

Struktura

Ljudski protein NANOG kodiran genom NANOG1, sastoji se od 305 aminokiselina i ina tri funkcionalna domena: N-terminalni, C-terminalni i konzervirani homeodomenski motiv. Homeodomenska regija olakšava vezivanje DNK. Ljudski gen Nanog 1 nalazi se na hromosomu 12, a iRNK sadrži 915 baznih parova otvorenog okvira čitanja (ORF) sa četiri egzona i tri introna.^[7]

N-terminalna regija ljudskog NANOG-a bogata je ostacima serina, treonina i prolina, a C-kraj sadrži domen bogat triptofanom. 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.^[9]

Funkcija

Transkripcijski programi u embrionskim matičnim ćelijama

NANOG je transkripcijski faktor u embrionskim matičnim ćelijama (ESC) i smatra se da je ključni faktor u održavanju pluripotencije. Smatra se da NANOG funkcionira u skladu s drugim faktorima, kao što su 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 gastrulacije eksprimira u stražnjoj strani epiblasta.^[10] Tamo je NANOG umiješan u inhibiranje embrionske hematopoeze potiskivanjem ekspresije transkripcijskog faktora Tal1.^[11] U ovoj fazi embriogeneze, NANOG potiskuje Pou3f1 , transkripcijski faktor ključan za formiranje prednje-stražnje osi.^[10]

Analiza usporenih embriona pokazala je da eksprimiraju gene markera pluripotencije, kao što su 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.^[12]

POU5F1, TDGF1 (CRIPTO), SALL4, LECT1 i BUB1 su također povezani geni koji su svi odgovorni za samoobnavljanje i pluripotentnu diferencijaciju.^[13]

Utvrđeno je da je protein NANOG transkripcijski aktivator promotora Rex1, koji ima ključnu ulogu u održavanju ekspresije Rex1. Nokaut NANOG u embrionskim matičnim ćelijama rezultira smanjenjem ekspresije Rex1, dok nadekspresija NANOG -a stimulira ekspresiju Rex1.^[14]

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.^[15]^[16]^[17]^[18]^[19]

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.^[20]^[21]^[22]

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.^[9]

Dijagnostika

Gen NANOG p8 ocijenjen je kao prognostički i prediktivni biomarker raka.^[23]

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.^[9]

Evolucija

Ljudi i obične čimpanze dijele deset NANOG pseudogena (NanogP2-P11) tokom evaluacije, dva se nalaze na X hromosomu i karakterišu ih 5' promotorske sekvence i odsustvo introna, kao posljedica retrotranspozicije iRNK ^[7] sve na istim mjestima: jedan duplicirani pseudogen i devet retropseudogena. Od devet zajedničkih NANOG retropseudogena, dva nemaju 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 genoma, evolucijski biolozi ukazuju na to da NANOG i njegovi pseudogeni pružaju dokaz zajedničkog porijekla ljudi čimpanzi.^[24]

Također pogledajte

Reference

^ ^a ^b ^c GRCh38: Ensembl release 89: ENSG00000111704 - Ensembl, maj 2017
^ ^a ^b ^c GRCm38: Ensembl release 89: ENSMUSG00000012396 - Ensembl, maj 2017
^ "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^ "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^ 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
^ "ScienceDaily: Cells Of The Ever Young: Getting Closer To The Truth". Pristupljeno 26. 7. 2007.
^ ^a ^b ^c Gawlik-Rzemieniewska N, Bednarek I (30. 11. 2015). "The role of NANOG transcriptional factor in the development of malignant phenotype of cancer cells". Cancer Biology & Therapy. 17 (1): 1–10. doi:10.1080/15384047.2015.1121348. PMC 4848008. PMID 26618281.
^ "UniProt, Q9H9S0". Pristupljeno 12. 8. 2021.
^ ^a ^b ^c Zhang W, Sui Y, Ni J, Yang T (2016). "Nanog gene: A propeller for stemness in primitive stem cells". International Journal of Biological Sciences. 12 (11): 1372–1381. doi:10.7150/ijbs.16349. PMC 5118783. PMID 27877089.
^ ^a ^b Barral A, Rollan I, Sanchez-Iranzo H, Jawaid W, Badia-Careaga C, Menchero S, et al. (decembar 2019). "Pou3f1 expression at the exit from pluripotency during gastrulation". Biology Open. 8 (11): bio046367. doi:10.1242/bio.046367. PMC 6899006. PMID 31791948.
^ Sainz de Aja J, Menchero S, Rollan I, Barral A, Tiana M, Jawaid W, et al. (april 2019). "Tal1". The EMBO Journal. 38 (7). doi:10.15252/embj.201899122. PMC 6443201. PMID 30814124.
^ Zhang X, Stojkovic P, Przyborski S, Cooke M, Armstrong L, Lako M, Stojkovic M (decembar 2006). "Derivation of human embryonic stem cells from developing and arrested embryos". Stem Cells. 24 (12): 2669–76. doi:10.1634/stemcells.2006-0377. PMID 16990582. S2CID 32587408.
^ Li SS, Liu YH, Tseng CN, Chung TL, Lee TY, Singh S (august 2006). "Characterization and gene expression profiling of five new human embryonic stem cell lines derived in Taiwan". Stem Cells and Development. 15 (4): 532–55. doi:10.1089/scd.2006.15.532. PMID 16978057.
^ Shi W, Wang H, Pan G, Geng Y, Guo Y, Pei D (august 2006). "Regulation of the pluripotency marker Rex-1 by Nanog and Sox2". The Journal of Biological Chemistry. 281 (33): 23319–25. doi:10.1074/jbc.M601811200. PMID 16714766.
^ Shahini A, Choudhury D, Asmani M, Zhao R, Lei P, Andreadis ST (januar 2018). "NANOG restores the impaired myogenic differentiation potential of skeletal myoblasts after multiple population doublings". Stem Cell Research. 26: 55–66. doi:10.1016/j.scr.2017.11.018. PMID 29245050.
^ Shahini A, Mistriotis P, Asmani M, Zhao R, Andreadis ST (juni 2017). "NANOG Restores Contractility of Mesenchymal Stem Cell-Based Senescent Microtissues". Tissue Engineering. Part A. 23 (11–12): 535–545. doi:10.1089/ten.TEA.2016.0494. PMC 5467120. PMID 28125933.
^ Mistriotis P, Bajpai VK, Wang X, Rong N, Shahini A, Asmani M, Liang MS, Wang J, Lei P, Liu S, Zhao R, Andreadis ST (januar 2017). "NANOG Reverses the Myogenic Differentiation Potential of Senescent Stem Cells by Restoring ACTIN Filamentous Organization and SRF-Dependent Gene Expression". Stem Cells. 35 (1): 207–221. doi:10.1002/stem.2452. PMID 27350449. S2CID 4482665.
^ Han J, Mistriotis P, Lei P, Wang D, Liu S, Andreadis ST (decembar 2012). "Nanog reverses the effects of organismal aging on mesenchymal stem cell proliferation and myogenic differentiation potential". Stem Cells. 30 (12): 2746–59. doi:10.1002/stem.1223. PMC 3508087. PMID 22949105.
^ Münst B, Thier MC, Winnemöller D, Helfen M, Thummer RP, Edenhofer F (mart 2016). "Nanog induces suppression of senescence through downregulation of p27KIP1 expression". Journal of Cell Science. 129 (5): 912–20. doi:10.1242/jcs.167932. PMC 4813312. PMID 26795560.
^ Gong S, Li Q, Jeter CR, Fan Q, Tang DG, Liu B (septembar 2015). "Regulation of NANOG in cancer cells". Molecular Carcinogenesis. 54 (9): 679–87. doi:10.1002/mc.22340. PMC 4536084. PMID 26013997.
^ Jeter CR, Yang T, Wang J, Chao HP, Tang DG (august 2015). "Concise Review: NANOG in Cancer Stem Cells and Tumor Development: An Update and Outstanding Questions". Stem Cells. 33 (8): 2381–90. doi:10.1002/stem.2007. PMC 4509798. PMID 25821200.
^ Gawlik-Rzemieniewska N, Bednarek I (2016). "The role of NANOG transcriptional factor in the development of malignant phenotype of cancer cells". Cancer Biology & Therapy. 17 (1): 1–10. doi:10.1080/15384047.2015.1121348. PMC 4848008. PMID 26618281.
^ Iv Santaliz-Ruiz LE, Xie X, Old M, Teknos TN, Pan Q (decembar 2014). "Emerging role of nanog in tumorigenesis and cancer stem cells". International Journal of Cancer. 135 (12): 2741–8. doi:10.1002/ijc.28690. PMC 4065638. PMID 24375318.
^ Fairbanks DJ (2007). Relics of Eden: The Powerful Evidence of Evolution in Human DNA. Buffalo, N.Y: Prometheus Books. str. 94–96, 177–182. ISBN 978-1-59102-564-1.

Dopunska literatura

Cavaleri F, Schöler HR (maj 2003). "Nanog: a new recruit to the embryonic stem cell orchestra". Cell. 113 (5): 551–2. doi:10.1016/S0092-8674(03)00394-5. PMID 12787492. S2CID 16254995.
Constantinescu S (2004). "Stemness, fusion and renewal of hematopoietic and embryonic stem cells". Journal of Cellular and Molecular Medicine. 7 (2): 103–12. doi:10.1111/j.1582-4934.2003.tb00209.x. PMC 6740230. PMID 12927049.
Pan G, Thomson JA (januar 2007). "Nanog and transcriptional networks in embryonic stem cell pluripotency". Cell Research. 17 (1): 42–9. doi:10.1038/sj.cr.7310125. PMID 17211451.
Mitsui K, Tokuzawa Y, Itoh H, Segawa K, Murakami M, Takahashi K, Maruyama M, Maeda M, Yamanaka S (maj 2003). "The homeoprotein Nanog is required for maintenance of pluripotency in mouse epiblast and ES cells". Cell. 113 (5): 631–42. doi:10.1016/S0092-8674(03)00393-3. PMID 12787504. S2CID 18836242.
Chambers I, Colby D, Robertson M, Nichols J, Lee S, Tweedie S, Smith A (maj 2003). "Functional expression cloning of Nanog, a pluripotency sustaining factor in embryonic stem cells". Cell. 113 (5): 643–55. doi:10.1016/S0092-8674(03)00392-1. hdl:1842/843. PMID 12787505. S2CID 2236779.
Clark AT, Rodriguez RT, Bodnar MS, Abeyta MJ, Cedars MI, Turek PJ, Firpo MT, Reijo Pera RA (2004). "Human STELLAR, NANOG, and GDF3 genes are expressed in pluripotent cells and map to chromosome 12p13, a hotspot for teratocarcinoma". Stem Cells. 22 (2): 169–79. doi:10.1634/stemcells.22-2-169. PMID 14990856. S2CID 38136098.
Hart AH, Hartley L, Ibrahim M, Robb L (maj 2004). "Identification, cloning and expression analysis of the pluripotency promoting Nanog genes in mouse and human". Developmental Dynamics. 230 (1): 187–98. doi:10.1002/dvdy.20034. PMID 15108323. S2CID 21502533.
Booth HA, Holland PW (august 2004). "Eleven daughters of NANOG". Genomics. 84 (2): 229–38. doi:10.1016/j.ygeno.2004.02.014. PMID 15233988.
Hatano SY, Tada M, Kimura H, Yamaguchi S, Kono T, Nakano T, Suemori H, Nakatsuji N, Tada T (januar 2005). "Pluripotential competence of cells associated with Nanog activity". Mechanisms of Development. 122 (1): 67–79. doi:10.1016/j.mod.2004.08.008. hdl:2433/144766. PMID 15582778. S2CID 17490059.
Deb-Rinker P, Ly D, Jezierski A, Sikorska M, Walker PR (februar 2005). "Sequential DNA methylation of the Nanog and Oct-4 upstream regions in human NT2 cells during neuronal differentiation". The Journal of Biological Chemistry. 280 (8): 6257–60. doi:10.1074/jbc.C400479200. PMID 15615706.
Zaehres H, Lensch MW, Daheron L, Stewart SA, Itskovitz-Eldor J, Daley GQ (mart 2005). "High-efficiency RNA interference in human embryonic stem cells". Stem Cells. 23 (3): 299–305. doi:10.1634/stemcells.2004-0252. PMID 15749924. S2CID 1395518.
Hoei-Hansen CE, Almstrup K, Nielsen JE, Brask Sonne S, Graem N, Skakkebaek NE, Leffers H, Rajpert-De Meyts E (juli 2005). "Stem cell pluripotency factor NANOG is expressed in human fetal gonocytes, testicular carcinoma in situ and germ cell tumours". Histopathology. 47 (1): 48–56. doi:10.1111/j.1365-2559.2005.02182.x. PMID 15982323. S2CID 10164525.
Hyslop L, Stojkovic M, Armstrong L, Walter T, Stojkovic P, Przyborski S, Herbert M, Murdoch A, Strachan T, Lako M (septembar 2005). "Downregulation of NANOG induces differentiation of human embryonic stem cells to extraembryonic lineages". Stem Cells. 23 (8): 1035–43. doi:10.1634/stemcells.2005-0080. PMID 15983365. S2CID 29881293.
Oh JH, Do HJ, Yang HM, Moon SY, Cha KY, Chung HM, Kim JH (juni 2005). "Identification of a putative transactivation domain in human Nanog". Experimental & Molecular Medicine. 37 (3): 250–4. doi:10.1038/emm.2005.33. PMID 16000880.
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 (septembar 2005). "Core transcriptional regulatory circuitry in human embryonic stem cells". Cell. 122 (6): 947–56. doi:10.1016/j.cell.2005.08.020. PMC 3006442. PMID 16153702.
Kim JS, Kim J, Kim BS, Chung HY, Lee YY, Park CS, Lee YS, Lee YH, Chung IY (decembar 2005). "Identification and functional characterization of an alternative splice variant within the fourth exon of human nanog". Experimental & Molecular Medicine. 37 (6): 601–7. doi:10.1038/emm.2005.73. PMID 16391521.
Darr H, Mayshar Y, Benvenisty N (mart 2006). "Overexpression of NANOG in human ES cells enables feeder-free growth while inducing primitive ectoderm features". Development. 133 (6): 1193–201. doi:10.1242/dev.02286. PMID 16501172.
Saunders A, Li D, Faiola F, Huang X, Fidalgo M, Guallar D, Ding J, Yang F, Xu Y, Zhou H, Wang J (maj 2017). "Context-Dependent Functions of NANOG Phosphorylation in Pluripotency and Reprogramming". Stem Cell Reports. 8 (5): 1115–1123. doi:10.1016/j.stemcr.2017.03.023. PMC 5425684. PMID 28457890.

Vanjski linkovi

NANOG protein, human na US National Library of Medicine Medical Subject Headings (MeSH)
Nanog protein, mouse na US National Library of Medicine Medical Subject Headings (MeSH)
https://www.factorbook.org/tf/human/NANOG/function
"Core Transcriptional Regulatory Circuitry in Human Embryonic Stem Cells". Young Lab. Whitehead Institute for Biomedical Research. Arhivirano s originala, 28. 6. 2009. Pristupljeno 28. 2. 2009.
"Jaenisch Lab Research Summary". Whitehead Institute. Arhivirano s originala, 26. 6. 2012. Pristupljeno 28. 2. 2009.
Discovery reveals more about stem cells' immortality

[refGRCh38Ensembl-1] GRCh38: Ensembl release 89: ENSG00000111704 - Ensembl, maj 2017

[refGRCm38Ensembl-2] GRCm38: Ensembl release 89: ENSMUSG00000012396 - Ensembl, maj 2017

[3] "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.

[4] "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.

[5] 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

[6] "ScienceDaily: Cells Of The Ever Young: Getting Closer To The Truth". Pristupljeno 26. 7. 2007.

[The_role_of_NANOG_transcriptional_f-7] Gawlik-Rzemieniewska N, Bednarek I (30. 11. 2015). "The role of NANOG transcriptional factor in the development of malignant phenotype of cancer cells". Cancer Biology & Therapy. 17 (1): 1–10. doi:10.1080/15384047.2015.1121348. PMC 4848008. PMID 26618281.

[UniProt-8] "UniProt, Q9H9S0". Pristupljeno 12. 8. 2021.

[Nanog_gene:_A_propeller_for_stemnes-9] Zhang W, Sui Y, Ni J, Yang T (2016). "Nanog gene: A propeller for stemness in primitive stem cells". International Journal of Biological Sciences. 12 (11): 1372–1381. doi:10.7150/ijbs.16349. PMC 5118783. PMID 27877089.

[Barral_2009-10] Barral A, Rollan I, Sanchez-Iranzo H, Jawaid W, Badia-Careaga C, Menchero S, et al. (decembar 2019). "Pou3f1 expression at the exit from pluripotency during gastrulation". Biology Open. 8 (11): bio046367. doi:10.1242/bio.046367. PMC 6899006. PMID 31791948.

[11] Sainz de Aja J, Menchero S, Rollan I, Barral A, Tiana M, Jawaid W, et al. (april 2019). "Tal1". The EMBO Journal. 38 (7). doi:10.15252/embj.201899122. PMC 6443201. PMID 30814124.

[pmid16990582-12] Zhang X, Stojkovic P, Przyborski S, Cooke M, Armstrong L, Lako M, Stojkovic M (decembar 2006). "Derivation of human embryonic stem cells from developing and arrested embryos". Stem Cells. 24 (12): 2669–76. doi:10.1634/stemcells.2006-0377. PMID 16990582. S2CID 32587408.

[pmid16978057-13] Li SS, Liu YH, Tseng CN, Chung TL, Lee TY, Singh S (august 2006). "Characterization and gene expression profiling of five new human embryonic stem cell lines derived in Taiwan". Stem Cells and Development. 15 (4): 532–55. doi:10.1089/scd.2006.15.532. PMID 16978057.

[14] Shi W, Wang H, Pan G, Geng Y, Guo Y, Pei D (august 2006). "Regulation of the pluripotency marker Rex-1 by Nanog and Sox2". The Journal of Biological Chemistry. 281 (33): 23319–25. doi:10.1074/jbc.M601811200. PMID 16714766.

[15] Shahini A, Choudhury D, Asmani M, Zhao R, Lei P, Andreadis ST (januar 2018). "NANOG restores the impaired myogenic differentiation potential of skeletal myoblasts after multiple population doublings". Stem Cell Research. 26: 55–66. doi:10.1016/j.scr.2017.11.018. PMID 29245050.

[16] Shahini A, Mistriotis P, Asmani M, Zhao R, Andreadis ST (juni 2017). "NANOG Restores Contractility of Mesenchymal Stem Cell-Based Senescent Microtissues". Tissue Engineering. Part A. 23 (11–12): 535–545. doi:10.1089/ten.TEA.2016.0494. PMC 5467120. PMID 28125933.

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