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[[Berkas:Intestinal_organoid.PNG|jmpl|Organoid usus yang dibuat dari sel punca Lgr5+]]
[[Berkas:Intestinal_organoid.PNG|jmpl|Organoid usus yang dibuat dari sel punca Lgr5+]]
'''Organoid''' adalah versi miniatur dan sederhana dari sebuah [[organ]], yang diproduksi [[In vitro|di laboratorium]] dan memiliki bentuk tiga dimensi dan anatomi yang mirip organ sungguhan. Organoid dibuat dari satu atau sejumlah sel dari suatu [[jaringan]], dari [[sel punca embrionik]] atau dari [[sel induk pluripoten diinduksi]] (iPS), yang dapat dengan spontan membentuk susunan tiga dimensi karena kapasitas sel-sel tersebut membelah diri serta melakukan [[Diferensiasi sel|diferensiasi]]. Teknologi penumbuhan organoid telah mengalami banyak kemajuan sejak awal 2010an dan disebut majalah ''[[The Scientist (majalah)|The Scientist]]'' sebagai salah satu kemajuan ilmu pengetahuan terbesar pada tahun 2013.<ref>{{cite web|url=http://www.the-scientist.com/?articles.view/articleNo/38747/title/2013-s-Big-Advances-in-Science/|title=2013's Big Advances in Science|last=Grens|first=Kerry|date=December 24, 2013|website=[[The Scientist (magazine)|The Scientist]]|access-date=26 December 2013|name-list-format=vanc}}</ref> Organoid digunakan para ilmuwan untuk meneliti penyakit dan pengobatannya di laboratorium.
'''Organoid''' adalah versi miniatur sederhana dari sebuah [[Organ (anatomi)|organ]], yang diproduksi [[In vitro|di laboratorium]] dan memiliki bentuk tiga dimensi dan anatomi yang mirip organ sungguhan. Organoid dibuat dari satu atau sejumlah sel dari suatu [[jaringan]], dari [[sel punca embrionik]] atau dari [[sel induk pluripoten diinduksi]] (iPS), yang dapat dengan spontan membentuk susunan tiga dimensi karena kapasitas sel-sel tersebut membelah diri serta melakukan [[Diferensiasi sel|diferensiasi]]. Teknologi penumbuhan organoid telah mengalami banyak kemajuan sejak awal 2010an dan disebut majalah ''[[The Scientist (majalah)|The Scientist]]'' sebagai salah satu kemajuan ilmu pengetahuan terbesar pada tahun 2013.<ref>{{cite web|url=http://www.the-scientist.com/?articles.view/articleNo/38747/title/2013-s-Big-Advances-in-Science/|title=2013's Big Advances in Science|last=Grens|first=Kerry|date=December 24, 2013|website=[[The Scientist (magazine)|The Scientist]]|access-date=26 December 2013|name-list-format=vanc}}</ref> Organoid digunakan para ilmuwan untuk meneliti penyakit dan pengobatannya di laboratorium.


== Jenis-jenis organoid ==
== Jenis-jenis organoid ==
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=== Organoid otak ===
=== Organoid otak ===
[[Organoid otak]] adalah organoid yang menyerupai otak yang ditumbuhkan di laboratorium. Organoid otak dibuat dengan menggunakan sel punca pluripoten manusia dalam sebuah [[bioreaktor]] rotasional tiga dimensi dan tumbuh dalam beberapa bulan.
[[Organoid otak]] adalah organoid yang menyerupai [[otak]] yang ditumbuhkan di laboratorium. Organoid otak dibuat dengan menggunakan sel punca pluripoten manusia dalam sebuah [[bioreaktor]] rotasional tiga dimensi dan tumbuh dalam beberapa bulan.


=== Organoid saluran pencernaan ===
=== Organoid saluran pencernaan ===
Dalam saluran pencernaan, organoid yang telah dibuat di antaranya organoid [[usus]] yang dibuat langsung dari sel punca pluripoten,<ref name="Lancaster, M. A. 2014">{{cite journal | vauthors = Lancaster MA, Knoblich JA | title = Organogenesis in a dish: modeling development and disease using organoid technologies | journal = Science | volume = 345 | issue = 6194 | pages = 1247125 | date = July 2014 | pmid = 25035496 | doi = 10.1126/science.1247125 }}</ref><ref name="pmid19329995"/> serta organoid [[lambung]].<ref name="Mccracken, K. W. 2014">{{cite journal | vauthors = McCracken KW, Catá EM, Crawford CM, Sinagoga KL, Schumacher M, Rockich BE, Tsai YH, Mayhew CN, Spence JR, Zavros Y, Wells JM | display-authors = 6 | title = Modelling human development and disease in pluripotent stem-cell-derived gastric organoids | journal = Nature | volume = 516 | issue = 7531 | pages = 400–4 | date = December 2014 | pmid = 25363776 | pmc = 4270898 | doi = 10.1038/nature13863 | bibcode = 2014Natur.516..400M }}</ref>
Dalam saluran pencernaan, organoid yang telah dibuat di antaranya organoid [[usus]] yang dibuat langsung dari sel punca pluripoten,<ref name="Lancaster, M. A. 2014">{{cite journal | vauthors = Lancaster MA, Knoblich JA | title = Organogenesis in a dish: modeling development and disease using organoid technologies | journal = Science | volume = 345 | issue = 6194 | pages = 1247125 | date = July 2014 | pmid = 25035496 | doi = 10.1126/science.1247125 }}</ref><ref name="pmid19329995">{{cite journal | vauthors = Sato T, Vries RG, Snippert HJ, van de Wetering M, Barker N, Stange DE, van Es JH, Abo A, Kujala P, Peters PJ, Clevers H | display-authors = 6 | title = Single Lgr5 stem cells build crypt-villus structures in vitro without a mesenchymal niche | journal = Nature | volume = 459 | issue = 7244 | pages = 262–5 | date = May 2009 | pmid = 19329995 | doi = 10.1038/nature07935 | bibcode = 2009Natur.459..262S | authorlink10 = Peter J. Peters }}</ref> serta organoid [[lambung]].<ref name="Mccracken, K. W. 2014">{{cite journal | vauthors = McCracken KW, Catá EM, Crawford CM, Sinagoga KL, Schumacher M, Rockich BE, Tsai YH, Mayhew CN, Spence JR, Zavros Y, Wells JM | display-authors = 6 | title = Modelling human development and disease in pluripotent stem-cell-derived gastric organoids | journal = Nature | volume = 516 | issue = 7531 | pages = 400–4 | date = December 2014 | pmid = 25363776 | pmc = 4270898 | doi = 10.1038/nature13863 | bibcode = 2014Natur.516..400M }}</ref>


=== Organoid lidah ===
=== Organoid lidah ===
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* Organoid [[epitelium]]<ref name="pmid19329995" /><ref name="pmid17934449">{{cite journal|display-authors=6|date=October 2007|title=Identification of stem cells in small intestine and colon by marker gene Lgr5|journal=Nature|volume=449|issue=7165|pages=1003–7|bibcode=2007Natur.449.1003B|doi=10.1038/nature06196|pmid=17934449|vauthors=Barker N, van Es JH, Kuipers J, Kujala P, van den Born M, Cozijnsen M, Haegebarth A, Korving J, Begthel H, Peters PJ, Clevers H|authorlink10=Peter J. Peters}}</ref>
* Organoid [[epitelium]]<ref name="pmid19329995" /><ref name="pmid17934449">{{cite journal|display-authors=6|date=October 2007|title=Identification of stem cells in small intestine and colon by marker gene Lgr5|journal=Nature|volume=449|issue=7165|pages=1003–7|bibcode=2007Natur.449.1003B|doi=10.1038/nature06196|pmid=17934449|vauthors=Barker N, van Es JH, Kuipers J, Kujala P, van den Born M, Cozijnsen M, Haegebarth A, Korving J, Begthel H, Peters PJ, Clevers H|authorlink10=Peter J. Peters}}</ref>
* Organoid [[paru-paru]]<ref>{{cite journal|display-authors=6|date=January 2014|title=Lung stem cell differentiation in mice directed by endothelial cells via a BMP4-NFATc1-thrombospondin-1 axis|journal=Cell|volume=156|issue=3|pages=440–55|doi=10.1016/j.cell.2013.12.039|pmc=3951122|pmid=24485453|vauthors=Lee JH, Bhang DH, Beede A, Huang TL, Stripp BR, Bloch KD, Wagers AJ, Tseng YH, Ryeom S, Kim CF}}</ref>
* Organoid [[paru-paru]]<ref>{{cite journal|display-authors=6|date=January 2014|title=Lung stem cell differentiation in mice directed by endothelial cells via a BMP4-NFATc1-thrombospondin-1 axis|journal=Cell|volume=156|issue=3|pages=440–55|doi=10.1016/j.cell.2013.12.039|pmc=3951122|pmid=24485453|vauthors=Lee JH, Bhang DH, Beede A, Huang TL, Stripp BR, Bloch KD, Wagers AJ, Tseng YH, Ryeom S, Kim CF}}</ref>
* Organoid [[ginjal]]<ref name="sciencedirect.com" /><ref>{{cite journal|display-authors=6|date=October 2015|title=Kidney organoids from human iPS cells contain multiple lineages and model human nephrogenesis|journal=Nature|volume=526|issue=7574|pages=564–8|bibcode=2015Natur.526..564T|doi=10.1038/nature15695|pmid=26444236|vauthors=Takasato M, Er PX, Chiu HS, Maier B, Baillie GJ, Ferguson C, Parton RG, Wolvetang EJ, Roost MS, Chuva de Sousa Lopes SM, Little MH}}</ref><ref name="ReferenceA" /><ref>{{cite journal|date=November 2015|title=Nephron organoids derived from human pluripotent stem cells model kidney development and injury|journal=Nature Biotechnology|volume=33|issue=11|pages=1193–200|doi=10.1038/nbt.3392|pmc=4747858|pmid=26458176|vauthors=Morizane R, Lam AQ, Freedman BS, Kishi S, Valerius MT, Bonventre JV}}</ref>
* Organoid [[ginjal]]<ref name="sciencedirect.com">{{cite journal | vauthors = Unbekandt M, Davies JA | title = Dissociation of embryonic kidneys followed by reaggregation allows the formation of renal tissues | url = https://archive.org/details/sim_kidney-international_2010-03_77_5/page/407 | journal = Kidney International | volume = 77 | issue = 5 | pages = 407–16 | date = March 2010 | pmid = 20016472 | doi = 10.1038/ki.2009.482 }}</ref><ref>{{cite journal|display-authors=6|date=October 2015|title=Kidney organoids from human iPS cells contain multiple lineages and model human nephrogenesis|journal=Nature|volume=526|issue=7574|pages=564–8|bibcode=2015Natur.526..564T|doi=10.1038/nature15695|pmid=26444236|vauthors=Takasato M, Er PX, Chiu HS, Maier B, Baillie GJ, Ferguson C, Parton RG, Wolvetang EJ, Roost MS, Chuva de Sousa Lopes SM, Little MH}}</ref><ref name="ReferenceA">{{cite journal | vauthors = Freedman BS, Brooks CR, Lam AQ, Fu H, Morizane R, Agrawal V, Saad AF, Li MK, Hughes MR, Werff RV, Peters DT, Lu J, Baccei A, Siedlecki AM, Valerius MT, Musunuru K, McNagny KM, Steinman TI, Zhou J, Lerou PH, Bonventre JV | display-authors = 6 | title = Modelling kidney disease with CRISPR-mutant kidney organoids derived from human pluripotent epiblast spheroids | journal = Nature Communications | volume = 6 | pages = 8715 | date = October 2015 | pmid = 26493500 | pmc = 4620584 | doi = 10.1038/ncomms9715 | bibcode = 2015NatCo...6E8715F }}</ref><ref>{{cite journal|date=November 2015|title=Nephron organoids derived from human pluripotent stem cells model kidney development and injury|journal=Nature Biotechnology|volume=33|issue=11|pages=1193–200|doi=10.1038/nbt.3392|pmc=4747858|pmid=26458176|vauthors=Morizane R, Lam AQ, Freedman BS, Kishi S, Valerius MT, Bonventre JV}}</ref>
* [[Gastruloid]] (Organoid embrionik)<ref>{{cite journal|display-authors=6|date=November 2014|title=Symmetry breaking, germ layer specification and axial organisation in aggregates of mouse embryonic stem cells|journal=Development|volume=141|issue=22|pages=4231–42|doi=10.1242/dev.113001|pmc=4302915|pmid=25371360|vauthors=van den Brink SC, Baillie-Johnson P, Balayo T, Hadjantonakis AK, Nowotschin S, Turner DA, Martinez Arias A}}</ref><ref>{{cite journal|date=February 2016|title=Organoids and the genetically encoded self-assembly of embryonic stem cells|journal=BioEssays|volume=38|issue=2|pages=181–91|doi=10.1002/bies.201500111|pmc=4737349|pmid=26666846|vauthors=Turner DA, Baillie-Johnson P, Martinez Arias A}}</ref><ref>{{cite journal|display-authors=6|date=November 2017|title=Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids|journal=Development|volume=144|issue=21|pages=3894–3906|doi=10.1242/dev.150391|pmc=5702072|pmid=28951435|vauthors=Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, Hayward PC, Collignon J, Gustavsen C, Serup P, Steventon B, P Lutolf M, Arias AM}}</ref><ref name=":0">{{cite journal|display-authors=6|date=October 2018|title=Multi-axial self-organization properties of mouse embryonic stem cells into gastruloids|journal=Nature|language=En|volume=562|issue=7726|pages=272–276|bibcode=2018Natur.562..272B|doi=10.1038/s41586-018-0578-0|pmid=30283134|vauthors=Beccari L, Moris N, Girgin M, Turner DA, Baillie-Johnson P, Cossy AC, Lutolf MP, Duboule D, Arias AM}}</ref>
* [[Gastruloid]] (Organoid embrionik)<ref>{{cite journal|display-authors=6|date=November 2014|title=Symmetry breaking, germ layer specification and axial organisation in aggregates of mouse embryonic stem cells|journal=Development|volume=141|issue=22|pages=4231–42|doi=10.1242/dev.113001|pmc=4302915|pmid=25371360|vauthors=van den Brink SC, Baillie-Johnson P, Balayo T, Hadjantonakis AK, Nowotschin S, Turner DA, Martinez Arias A}}</ref><ref>{{cite journal|date=February 2016|title=Organoids and the genetically encoded self-assembly of embryonic stem cells|journal=BioEssays|volume=38|issue=2|pages=181–91|doi=10.1002/bies.201500111|pmc=4737349|pmid=26666846|vauthors=Turner DA, Baillie-Johnson P, Martinez Arias A}}</ref><ref>{{cite journal|display-authors=6|date=November 2017|title=Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids|journal=Development|volume=144|issue=21|pages=3894–3906|doi=10.1242/dev.150391|pmc=5702072|pmid=28951435|vauthors=Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, Hayward PC, Collignon J, Gustavsen C, Serup P, Steventon B, P Lutolf M, Arias AM}}</ref><ref name=":0">{{cite journal|display-authors=6|date=October 2018|title=Multi-axial self-organization properties of mouse embryonic stem cells into gastruloids|journal=Nature|language=En|volume=562|issue=7726|pages=272–276|bibcode=2018Natur.562..272B|doi=10.1038/s41586-018-0578-0|pmid=30283134|vauthors=Beccari L, Moris N, Girgin M, Turner DA, Baillie-Johnson P, Cossy AC, Lutolf MP, Duboule D, Arias AM}}</ref>
* Organoid [[jantung]]<ref>{{cite journal|date=February 2008|title=Engineered cardiac organoid chambers: toward a functional biological model ventricle|journal=Tissue Engineering. Part A|volume=14|issue=2|pages=215–25|doi=10.1089/tea.2007.0351|pmid=18333774|vauthors=Lee EJ, Kim DE, Azeloglu EU, Costa KD}}</ref> - Pada 2018 sebuah organoid jantung dapat berdetak dan merespons stimulus dengan berdetak lebih cepat atau lambat<ref>{{Cite news|url=https://www.wired.com/story/these-beating-mini-hearts-could-save-big-bucksand-maybe-lives/|title=These Beating Mini-Hearts Could Save Big Bucks—And Maybe Lives|last=Molteni|first=Megan|name-list-format=vanc|date=2018-06-27|work=WIRED|access-date=2018-06-30}}</ref>
* Organoid [[jantung]]<ref>{{cite journal|date=February 2008|title=Engineered cardiac organoid chambers: toward a functional biological model ventricle|journal=Tissue Engineering. Part A|volume=14|issue=2|pages=215–25|doi=10.1089/tea.2007.0351|pmid=18333774|vauthors=Lee EJ, Kim DE, Azeloglu EU, Costa KD}}</ref> - Pada 2018 sebuah organoid jantung dapat berdetak dan merespons stimulus dengan berdetak lebih cepat atau lambat<ref>{{Cite news|url=https://www.wired.com/story/these-beating-mini-hearts-could-save-big-bucksand-maybe-lives/|title=These Beating Mini-Hearts Could Save Big Bucks—And Maybe Lives|last=Molteni|first=Megan|name-list-format=vanc|date=2018-06-27|work=WIRED|access-date=2018-06-30}}</ref>
* Organoid [[retina]] <ref>{{cite journal|display-authors=6|date=July 2016|title=cGMP production of patient-specific iPSCs and photoreceptor precursor cells to treat retinal degenerative blindness|journal=Scientific Reports|volume=6|pages=30742|bibcode=2016NatSR...630742W|doi=10.1038/srep30742|pmc=4965859|pmid=27471043|vauthors=Wiley LA, Burnight ER, DeLuca AP, Anfinson KR, Cranston CM, Kaalberg EE, Penticoff JA, Affatigato LM, Mullins RF, Stone EM, Tucker BA}}</ref>
* Organoid [[retina]] <ref>{{cite journal|display-authors=6|date=July 2016|title=cGMP production of patient-specific iPSCs and photoreceptor precursor cells to treat retinal degenerative blindness|journal=Scientific Reports|volume=6|pages=30742|bibcode=2016NatSR...630742W|doi=10.1038/srep30742|pmc=4965859|pmid=27471043|vauthors=Wiley LA, Burnight ER, DeLuca AP, Anfinson KR, Cranston CM, Kaalberg EE, Penticoff JA, Affatigato LM, Mullins RF, Stone EM, Tucker BA}}</ref>
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== Referensi ==
== Referensi ==
{{reflist}}
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[[Kategori:Biologi]]

Revisi terkini sejak 13 September 2021 19.44

Organoid usus yang dibuat dari sel punca Lgr5+

Organoid adalah versi miniatur sederhana dari sebuah organ, yang diproduksi di laboratorium dan memiliki bentuk tiga dimensi dan anatomi yang mirip organ sungguhan. Organoid dibuat dari satu atau sejumlah sel dari suatu jaringan, dari sel punca embrionik atau dari sel induk pluripoten diinduksi (iPS), yang dapat dengan spontan membentuk susunan tiga dimensi karena kapasitas sel-sel tersebut membelah diri serta melakukan diferensiasi. Teknologi penumbuhan organoid telah mengalami banyak kemajuan sejak awal 2010an dan disebut majalah The Scientist sebagai salah satu kemajuan ilmu pengetahuan terbesar pada tahun 2013.[1] Organoid digunakan para ilmuwan untuk meneliti penyakit dan pengobatannya di laboratorium.

Jenis-jenis organoid

[sunting | sunting sumber]

Banyak struktur organ telah berhasil diminaturkan menjadi organoid.[2]

Organoid otak

[sunting | sunting sumber]

Organoid otak adalah organoid yang menyerupai otak yang ditumbuhkan di laboratorium. Organoid otak dibuat dengan menggunakan sel punca pluripoten manusia dalam sebuah bioreaktor rotasional tiga dimensi dan tumbuh dalam beberapa bulan.

Organoid saluran pencernaan

[sunting | sunting sumber]

Dalam saluran pencernaan, organoid yang telah dibuat di antaranya organoid usus yang dibuat langsung dari sel punca pluripoten,[2][3] serta organoid lambung.[4]

Organoid lidah

[sunting | sunting sumber]

Organoid lidah telah dibuat menggunakan protein sel punca epitelium yang mengekspresikan BMI1 dalam kultur tiga dimensi melalui manipulasi EGF, WNT, and TGF-β.[5] Kultur rganoid ini tidak memiliki sel reseptor rasa, karena sel tersebut tidak berasal dari sel punca epitelium yang mengekspresikan BMI1.[5] Namun, organoid lidah dengan indra pengecap telah dibuat dengan menggunakan sel punca LGR5+ atau CD44+.[6]

Referensi

[sunting | sunting sumber]
  1. ^ Grens K (December 24, 2013). "2013's Big Advances in Science". The Scientist. Diakses tanggal 26 December 2013. 
  2. ^ a b Lancaster MA, Knoblich JA (July 2014). "Organogenesis in a dish: modeling development and disease using organoid technologies". Science. 345 (6194): 1247125. doi:10.1126/science.1247125. PMID 25035496. 
  3. ^ a b Sato T, Vries RG, Snippert HJ, van de Wetering M, Barker N, Stange DE, et al. (May 2009). "Single Lgr5 stem cells build crypt-villus structures in vitro without a mesenchymal niche". Nature. 459 (7244): 262–5. Bibcode:2009Natur.459..262S. doi:10.1038/nature07935. PMID 19329995. 
  4. ^ McCracken KW, Catá EM, Crawford CM, Sinagoga KL, Schumacher M, Rockich BE, et al. (December 2014). "Modelling human development and disease in pluripotent stem-cell-derived gastric organoids". Nature. 516 (7531): 400–4. Bibcode:2014Natur.516..400M. doi:10.1038/nature13863. PMC 4270898alt=Dapat diakses gratis. PMID 25363776. 
  5. ^ a b Hisha, H., Tanaka, T., Kanno, S., Tokuyama, Y., Komai, Y., Ohe, S., . . . Ueno, H. (2013). Establishment of a Novel Lingual Organoid Culture System: Generation of Organoids Having Mature Keratinized Epithelium from Adult Epithelial Stem Cells. Scientific Reports, 3. doi:10.1038/srep03224
  6. ^ Aihara E, Mahe MM, Schumacher MA, Matthis AL, Feng R, Ren W, et al. (November 2015). "Characterization of stem/progenitor cell cycle using murine circumvallate papilla taste bud organoid". Scientific Reports. 5: 17185. Bibcode:2015NatSR...517185A. doi:10.1038/srep17185. PMC 4665766alt=Dapat diakses gratis. PMID 26597788. 
  7. ^ Martin A, Barbesino G, Davies TF (1999). "T-cell receptors and autoimmune thyroid disease--signposts for T-cell-antigen driven diseases". International Reviews of Immunology. 18 (1-2): 111–40. doi:10.3109/08830189909043021. PMID 10614741. 
  8. ^ Bredenkamp N, Ulyanchenko S, O'Neill KE, Manley NR, Vaidya HJ, Blackburn CC (September 2014). "An organized and functional thymus generated from FOXN1-reprogrammed fibroblasts". Nature Cell Biology. 16 (9): 902–8. doi:10.1038/ncb3023. PMC 4153409alt=Dapat diakses gratis. PMID 25150981. 
  9. ^ Huch M, Gehart H, van Boxtel R, Hamer K, Blokzijl F, Verstegen MM, et al. (January 2015). "Long-term culture of genome-stable bipotent stem cells from adult human liver". Cell. 160 (1-2): 299–312. doi:10.1016/j.cell.2014.11.050. PMC 4313365alt=Dapat diakses gratis. PMID 25533785. 
  10. ^ Huch M, Bonfanti P, Boj SF, Sato T, Loomans CJ, van de Wetering M, et al. (October 2013). "Unlimited in vitro expansion of adult bi-potent pancreas progenitors through the Lgr5/R-spondin axis". The EMBO Journal. 32 (20): 2708–21. doi:10.1038/emboj.2013.204. PMC 3801438alt=Dapat diakses gratis. PMID 24045232. 
  11. ^ Barker N, van Es JH, Kuipers J, Kujala P, van den Born M, Cozijnsen M, et al. (October 2007). "Identification of stem cells in small intestine and colon by marker gene Lgr5". Nature. 449 (7165): 1003–7. Bibcode:2007Natur.449.1003B. doi:10.1038/nature06196. PMID 17934449. 
  12. ^ Lee JH, Bhang DH, Beede A, Huang TL, Stripp BR, Bloch KD, et al. (January 2014). "Lung stem cell differentiation in mice directed by endothelial cells via a BMP4-NFATc1-thrombospondin-1 axis". Cell. 156 (3): 440–55. doi:10.1016/j.cell.2013.12.039. PMC 3951122alt=Dapat diakses gratis. PMID 24485453. 
  13. ^ Unbekandt M, Davies JA (March 2010). "Dissociation of embryonic kidneys followed by reaggregation allows the formation of renal tissues". Kidney International. 77 (5): 407–16. doi:10.1038/ki.2009.482. PMID 20016472. 
  14. ^ Takasato M, Er PX, Chiu HS, Maier B, Baillie GJ, Ferguson C, et al. (October 2015). "Kidney organoids from human iPS cells contain multiple lineages and model human nephrogenesis". Nature. 526 (7574): 564–8. Bibcode:2015Natur.526..564T. doi:10.1038/nature15695. PMID 26444236. 
  15. ^ Freedman BS, Brooks CR, Lam AQ, Fu H, Morizane R, Agrawal V, et al. (October 2015). "Modelling kidney disease with CRISPR-mutant kidney organoids derived from human pluripotent epiblast spheroids". Nature Communications. 6: 8715. Bibcode:2015NatCo...6E8715F. doi:10.1038/ncomms9715. PMC 4620584alt=Dapat diakses gratis. PMID 26493500. 
  16. ^ Morizane R, Lam AQ, Freedman BS, Kishi S, Valerius MT, Bonventre JV (November 2015). "Nephron organoids derived from human pluripotent stem cells model kidney development and injury". Nature Biotechnology. 33 (11): 1193–200. doi:10.1038/nbt.3392. PMC 4747858alt=Dapat diakses gratis. PMID 26458176. 
  17. ^ van den Brink SC, Baillie-Johnson P, Balayo T, Hadjantonakis AK, Nowotschin S, Turner DA, et al. (November 2014). "Symmetry breaking, germ layer specification and axial organisation in aggregates of mouse embryonic stem cells". Development. 141 (22): 4231–42. doi:10.1242/dev.113001. PMC 4302915alt=Dapat diakses gratis. PMID 25371360. 
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