Organoid - Knowledge

982:, and led to the first application of organoids for personalised treatment. Cystic fibrosis is an inherited disease that is caused by gene mutations of the cystic fibrosis transmembrane conductance regulator gene that encodes an epithelial ion channel necessary for healthy epithelial surface fluids. Studies by the laboratory of Jeffrey Beekman (Wilhelmina Children's Hospital, University Medical Center Utrecht, The Netherlands) described in 2013 that stimulation of colorectal organoids with cAMP-raising agonists such as forskolin or cholera toxin induced rapid swelling of organoids in a fully CFTR dependent manner. Whereas organoids from non-cystic fibrosis subjects swell in response to forskolin as a consequence of fluid transport into the organoids' lumens, this is severely reduced or absent in organoids derived from people with cystic fibrosis. Swelling could be restored by therapeutics that repair the CFTR protein (CFTR modulators), indicating that individual responses to CFTR modulating therapy could be quantitated in a preclinical laboratory setting. Schwank et al. also demonstrated that the intestinal cystic fibrosis organoid phenotype could be repaired by CRISPR-Cas9 gene editing in 2013. 986:

function. In addition, the authors demonstrated that CFTR modulator responses in intestinal organoids with specific CFTR mutations correlated with published clinical trial data of these treatments. This led to preclinical studies where organoids from patients with extremely rare CFTR mutations for who no treatment was registered were found to respond strongly to a clinically available CFTR modulator. The suggested clinical benefit of treatment for these subjects based on the preclinical organoid test was subsequently confirmed upon clinical introduction of treatment by members of the clinical CF center under supervision of Kors van der Ent (Department of Paediatric Pulmonology, Wilhelmina Children's Hospital, University Medical Center Utrecht, The Netherlands). These studies show for the first time that organoids can be used for the individual tailoring of therapy or

872:. These CRISPR-modified pluripotent stem cells were subsequently grown into human kidney organoids, which exhibited disease-specific phenotypes. Kidney organoids from stem cells with polycystic kidney disease mutations formed large, translucent cyst structures from kidney tubules. When cultured in the absence of adherent cues (in suspension), these cysts reached sizes of 1 cm in diameter over several months. Kidney organoids with mutations in a gene linked to focal segmental glomerulosclerosis developed junctional defects between podocytes, the filtering cells affected in that disease. Importantly, these disease phenotypes were absent in control organoids of identical genetic background, but lacking the CRISPR mutations. Comparison of these organoid phenotypes to diseased tissues from mice and humans suggested similarities to defects in early development. 801:

transporters, and drug efflux pumps, and can therefore be used to model drug transport across the BBB. Also, they can serve as a model for evaluating the interactions between the BBB and adjacent brain tissue and provide a platform for understanding the combined abilities of a new drug to overcome the BBB and its effect on brain tissue. In addition, such models are highly scalable and easier to manufacture and operate than microfluidic devices. However, they have limited ability to reconstruct the morphology and physiology of the BBB and are unable to simulate physiological flow and

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established colon organoids as tools for high-throughput drug screening, toxicity testing, and oral drug development. Or recently, Sakib, S., and Zou, S. developed graphene oxide nanoparticles for delivering siRNA regulating expression of tumor necrosis factor-α, that aimed to treat intestinal organoids exhibiting an inflammatory phenotype. Intestinal organoids also recapitulate the

398:. They propose that further evolution of the technique needs to be subject to a rigorous oversight procedure. In 2023, researchers have built a hybrid biocomputer that combines laboratory-grown human brain organoids with conventional circuits, and can complete tasks such as voice recognition. Cerebral Organoids are currently being used to research and develop 346:

microenvironments that are close to their counterparts, physiologically. Vasculature systems that can facilitate oxygen or nutrients to the inner mass of organoids can be achieved through microfluidic systems, vascular endothelial growth factor delivery systems, and endothelial cell-coated modules. With patient-derived

879:(iPSC) can also be reprogrammed from patient skin fibroblasts. These stem cells carry the exact genetic background of the patient including any genetic mutations which might contribute to the development of human disease. Differentiation of these cells into kidney organoids has been performed from patients with 1033:

using 2D cultures. These advancements in PSC fate direction, coupled with the advancements in 3D culturing techniques allowed for the creation of organoids that recapitulate the properties of various specific subregions of a multitude of organs. The use of these organoids has thus greatly contributed

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The self-renewal and regenerative properties of intestinal organoids make them promising candidates for transplantation therapies, particularly for disease involving epithelial barrier disruption. Notably, transplantation of organoids into the duodenum of mice has been shown to aid in the recovery of

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culture makes this system easy to manipulate and facilitates their monitoring. While organs are difficult to culture because their size limits the penetration of nutrients, the small size of organoids limits this problem. On the other hand, they do not exhibit all organ features and interactions with

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has in maintaining in intestinal stem cells, along with demonstrating the roles of other cell types like neurons and fibroblasts in maintenance of intestinal stem cells. In the field of infection biology, different intestinal organoid-based model systems have been explored. On one hand, organoids can

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and develop over the course of months. The procedure has potential applications in the study of brain development, physiology and function. Cerebral organoids may experience "simple sensations" in response to external stimulation and neuroscientists are among those expressing concern that such organs

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demonstrated that single Lgr5+ cells from damaged mouse liver can be clonally expanded as liver organoids in Rspo1-based culture medium over several months. In 2014, Artem Shkumatov et al. at the University of Illinois at Urbana-Champaign demonstrated that cardiovascular organoids can be formed from

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types. The advent of the field of organoids, started with a shift from culturing and differentiating stem cells in two dimensional (2D) media, to three dimensional (3D) media to allow for the development of the complex 3-dimensional structures of organs. Utilization of 3D media culture media methods

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development, for example, organoids have contributed to our understanding of the physical forces that underlie retinal cup formation. More recent work has extended cortical organoid growth periods extensively and at nearly a year under specific differentiation conditions, the organoids persist and

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control, which can be performed simultaneously with iPSC reprogramming. Comparison of a patient iPSC derived organoid against an isogenic control is the current gold standard in the field as it permits isolation of the mutation of interest as the only variable within the experimental model. In one

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structure to such a high degree of fidelity that they have been successfully transplanted to mouse intestines, and are hence highly regarded as a valuable model for research. One of the fields of research that intestinal organoids have been utilized is that of stem cell niche. Intestinal organoids

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demonstrated the production of renal organoids from murine fetus-derived renogenic stem cells. In 2014, Qun Wang and co-workers engineered collagen-I and laminin based gels and synthetic foam biomaterials for the culture and delivery of intestinal organoids and encapsulated DNA-functionalized gold

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signaling pathways as they have been demonstrated to promote posterior gut fate. Intestinal organoids have also been generated from intestinal stem cells, extracted from adult tissue and cultured in 3D media. These adult stem cell-derived organoids are often referred to as enteroids or colonoids,

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in a 3D medium. Stem cells have the ability to self-renew and differentiate into various cell subtypes, and they enable understanding the processes of development and disease progression. Therefore organoids derived from stem cells enable studying biology and physiology at the organ level. The 3D

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Follow-up studies by Dekkers et al. in 2016 revealed that quantitative differences in forskolin-induced swelling between intestinal organoids derived from people with cystic fibrosis associate with known diagnostic and prognostic markers such as CFTR gene mutations or in vivo biomarkers of CFTR

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cells surrounding a central lumen. As such, recapitulate the crypt-villus structure of the intestine, by recapitulating its function, physiology and organization, and maintaining all the cell types found normally in the structure including intestinal stem cells. Thus, intestinal organoids are a

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A multitude of organ structures have been recapitulated using organoids. This section aims to outline the state of the field as of now through providing an abridged list of the organoids that have been successfully created, along with a brief outline based on the most recent literature for each

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and colleagues showed that a laminin rich gel can be used as a basement membrane for differentiation and morphogenesis in cell cultures of mammary epithelial cells. Since 1987, researchers have devised different methods for 3D culturing, and were able to utilize different types of stem cells to

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and nanomedicine, incretin hormone secretion, and infection by various enteropathogens. For example, Qun Wang's team rationally designed artificial virus nanoparticles as oral drug delivery vehicles (ODDVs) with gut organoid-derived mucosal models and demonstrated a new concept of using newly

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Self-assembled cell aggregates consisting of BMECs, astrocytes, and pericytes are emerging as a potential alternative to transwell and microfluidic models for certain applications. These organoides can generate many features of the BBB, such as the expression of tight junctions, molecular

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Biochemical cues have been incorporated in 3D organoid cultures and with exposure of morphogenes, morphogen inhibitors, or growth factors, organoid models can be developed using embryonic stem cells (ESCs) or adult stem cells (ASCs). Vascularization techniques can be utilized to embody

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Recent advances in cell repellent microtiter plates has allowed rapid, cost-effective screening of large small molecule drug like libraries against 3D models of pancreas cancer. These models are consistent in phenotype and expression profiles with those found in the lab of Dr.

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gene mutated. To further assess the role of TGFBR2 in the metastasis, the investigators created organoids where TGFBR2 expression is knocked down, through which they were able to demonstrate that reduced TGFBR2 activity leads to invasion and metastasis of cancerous tumors both

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Organoids provide an opportunity to create cellular models of human disease, which can be studied in the laboratory to better understand the causes of disease and identify possible treatments. The power of organoids in this regard was first shown for a genetic form of

864:, which were smaller and showed abnormalities in early generation of neurons. In another example, the genome editing system called CRISPR was applied to human pluripotent stem cells to introduce targeted mutations in genes relevant to two different kidney diseases, 942:

morphology consistent with existing animal models which was rescued to wild type morphology in the gene corrected organoids. Comparative transcriptional profiling of epithelial cells purified from patient and control organoids highlighted pathways involved in

342:. Those embryoid bodies are then pharmacologically treated with patterning factors to drive the formation of the desired organoid identity. Organoids have also been created using adult stem cells extracted from the target organ, and cultured in 3D media. 777:

3D organoid models of brain cancer derived from either patient derived explants (PDX) or direct from cancer tissue is now easily achievable and affords high-throughput screening of these tumors against the current panel of approved drugs form around the

676:, which is a lymphoid organ where T cells mature. Thymic organoids have been generated through the seeding of thymic stromal cells in 3-dimensional culture. Thymic organoids seem to successfully recapitulate the thymus' function, as co-culturing human 456:

Intestinal organoids have thus far been among the gut organoids generated directly from intestinal tissues or pluripotent stem cells. One way human pluripotent stem cells can be driven to form intestinal organoids is through first the application of

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generate organoids resembling a multitude of organs. In the 1990s, in addition to their role in physical support, the role of ECM components in gene expression by their interaction with integrin-based focal adhesion pathways was reported. In 2006,

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Lancaster and Knoblich define an organoid as a collection of organ-specific cell types that develops from stem cells or organ progenitors, self-organizes through cell sorting and spatially restricted lineage commitment in a manner similar to

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Dekkers JF, Berkers G, Kruisselbrink E, Vonk A, de Jonge HR, Janssens HM, et al. (June 2016). "Characterizing responses to CFTR-modulating drugs using rectal organoids derived from subjects with cystic fibrosis".

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mutations. This report compared kidney organoids differentiated from patient iPSC to unrelated control iPSC and demonstrated an inability of patient kidney cells to mobilise transcription factor SIX2 from the

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demonstrated that mechanically dissociated sponge cells can reaggregate and self-organize to generate a whole organism. In the subsequent decades, multiple labs were able to generate different types of organs

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Davoudi Z, Atherly T, Borcherding DC, Jergens AE, Wannemuehler M, Barrett TA, et al. (December 2023). "Study Transportation of Drugs within Newly Established Murine Colon Organoid Systems".

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and cultured as 2D monolayers in order to make both the apical and basolateral sides of the epithelium more easily accessible. Intestinal organoids have also demonstrated therapeutic potential.

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Dekkers JF, Wiegerinck CL, de Jonge HR, Bronsveld I, Janssens HM, de Winter-de Groot KM, et al. (July 2013). "A functional CFTR assay using primary cystic fibrosis intestinal organoids".

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nanoparticles into intestinal organoids to form an intestinal Trojan horse for drug delivery and gene therapy. Subsequent reports showed significant physiological function of these organoids

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Beccari L, Moris N, Girgin M, Turner DA, Baillie-Johnson P, Cossy AC, et al. (October 2018). "Multi-axial self-organization properties of mouse embryonic stem cells into gastruloids".

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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".

418:, which during development forms a tube that can be divided in three distinct regions, which give rise to, along with other organs, the following sections of the gastrointestinal tract: 4901:

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".

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and 3T3 fibroblasts. The phenomena of mechanically dissociated cells aggregating and reorganizing to reform the tissue they were obtained from subsequently led to the development of the

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Yui S, Nakamura T, Sato T, Nemoto Y, Mizutani T, Zheng X, et al. (March 2012). "Functional engraftment of colon epithelium expanded in vitro from a single adult Lgr5⁺ stem cell".

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assembly, some of which had been implicated for other genotypes within the phenotypic family of renal ciliopathies. Another report utilising an isogenic control demonstrated abnormal

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Takahashi K, Tanabe K, Ohnuki M, Narita M, Ichisaka T, Tomoda K, et al. (November 2007). "Induction of pluripotent stem cells from adult human fibroblasts by defined factors".

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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".

354:-based genome editing technologies, genome-edited or mutated pluripotent stem cells (PSCs) with altered signaling cues can be generated to control intrinsic cues within organoids. 642:-expressing taste stem/progenitor cells. and indirectly, through the isolation, digestion, and subsequent culturing of CV tissue containing Lgr5+ or CD44+ stem/progenitor cells. 827:

and regulation of stemness was the first field of application of intestinal organoids, they are now also used to study e.g. uptake of nutrients, drug transport and secretion of

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depending on their segment of origin, and have been established from both the human and murine intestine. Intestinal organoids consist of a single layer of polarized intestinal

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Watanabe S, Kobayashi S, Ogasawara N, Okamoto R, Nakamura T, Watanabe M, et al. (March 2022). "Transplantation of intestinal organoids into a mouse model of colitis".

334:-rich extracellular matrix that is secreted by the Engelbreth-Holm-Swarm tumor line. Organoid bodies can then be made through embedding stem cells in the 3D medium. When 5265: 5163:"Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids" 265:

through modulation of the substrate stiffness, to which they adhere. Physiological stiffness promoted three-dimensionality of EBs and cardiomyogenic differentiation.

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Nahmias Y, Schwartz RE, Hu WS, Verfaillie CM, Odde DJ (June 2006). "Endothelium-mediated hepatocyte recruitment in the establishment of liver-like tissue in vitro".

630:, as these cells do not arise from Bmi1 expressing epithelial stem cells. Lingual taste bud organoids containing taste cells, however, have been created using the 2682:

Jung P, Sato T, Merlos-Suárez A, Barriga FM, Iglesias M, Rossell D, et al. (September 2011). "Isolation and in vitro expansion of human colonic stem cells".

86:, for drug discovery and development in industry, personalized diagnostics and medicine, gene and cell therapies, tissue engineering, and regenerative medicine. 814:

Organoids enable to study how cells interact together in an organ, their interaction with their environment, how diseases affect them and the effect of drugs.

4208:"From Pluripotent Stem Cells to Organoids and Bioprinting: Recent Advances in Dental Epithelium and Ameloblast Models to Study Tooth Biology and Regeneration" 2355:"Three-Dimensional Gastrointestinal Organoid Culture in Combination with Nerves or Fibroblasts: A Method to Characterize the Gastrointestinal Stem Cell Niche" 5301:

Rivron NC, Frias-Aldeguer J, Vrij EJ, Boisset JC, Korving J, Vivié J, et al. (May 2018). "Blastocyst-like structures generated solely from stem cells".

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Lingual organoids are organoids that recapitulate, at least partly, aspects of the tongue physiology. Epithelial lingual organoids have been generated using

638:+ stem/progenitor cells of circumvallate (CV) papilla tissue. These taste bud organoids have been successfully created both directly from isolated Lgr5- or 4032:"Establishment of a novel lingual organoid culture system: generation of organoids having mature keratinized epithelium from adult epithelial stem cells" 3477:

Dutta D, Heo I, O'Connor R (September 2019). "Studying Cryptosporidium Infection in 3D Tissue-derived Human Organoid Culture Systems by Microinjection".

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Rahmani S, Breyner NM, Su HM, Verdu EF, Didar TF (February 2019). "Intestinal organoids: A new paradigm for engineering intestinal epithelium in vitro".

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through the dissociation and reaggregation of organ tissues obtained from amphibians and embryonic chicks. The formation of first tissue-like colonies

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Takahashi K, Yamanaka S (August 2006). "Induction of pluripotent stem cells from mouse embryonic and adult fibroblast cultures by defined factors".

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Peng H, Poovaiah N, Forrester M, Cochran E, Wang Q (December 2014). "Ex Vivo Culture of Primary Intestinal Stem Cells in Collagen Gels and Foams".

2540: 6146:"Patient-iPSC-Derived Kidney Organoids Show Functional Validation of a Ciliopathic Renal Phenotype and Reveal Underlying Pathogenetic Mechanisms" 3299:"Attenuation of Chronic Inflammation in Intestinal Organoids with Graphene Oxide-Mediated Tumor Necrosis Factor-α_Small Interfering RNA Delivery" 5401:

Lee EJ, Kim DE, Azeloglu EU, Costa KD (February 2008). "Engineered cardiac organoid chambers: toward a functional biological model ventricle".

6447:"Organoids transplantation attenuates intestinal ischemia/reperfusion injury in mice through L-Malic acid-mediated M2 macrophage polarization" 5710:"Druggable growth dependencies and tumor evolution analysis in patient-derived organoids of neuroendocrine neoplasms from multiple body sites" 2291:"Influence of a reconstituted basement membrane and its components on casein gene expression and secretion in mouse mammary epithelial cells" 1499:"Influence of a reconstituted basement membrane and its components on casein gene expression and secretion in mouse mammary epithelial cells" 82:

named it one of the biggest scientific advancements of 2013. Scientists and engineers use organoids to study development and disease in the

6031:"Kidney-differentiated cells derived from Lowe Syndrome patient's iPSCs show ciliogenesis defects and Six2 retention at the Golgi complex" 4253:

Martin A, Barbesino G, Davies TF (1999). "T-cell receptors and autoimmune thyroid disease--signposts for T-cell-antigen driven diseases".

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Rheinwald JG, Green H (November 1975). "Formation of a keratinizing epithelium in culture by a cloned cell line derived from a teratoma".

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in three dimensions that mimics the key functional, structural, and biological complexity of that organ. It is derived from one or a few

6198:"Organoids from Nephrotic Disease-Derived iPSCs Identify Impaired NEPHRIN Localization and Slit Diaphragm Formation in Kidney Podocytes" 3009:"Organoids to Study Intestinal Nutrient Transport, Drug Uptake and Metabolism - Update to the Human Model and Expansion of Applications" 585:

along with human disease and development. For example, one study investigated the underlying genetic alterations behind a patient's

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Intestinal organoids grown from rectal biopsies using culture protocols established by the Clevers group have been used to model

1896:"Transport of organic anions and cations in murine embryonic kidney development and in serially-reaggregated engineered kidneys" 1226:

Lancaster MA, Knoblich JA (July 2014). "Organogenesis in a dish: modeling development and disease using organoid technologies".

4532:"Recapitulating hepatitis E virus-host interactions and facilitating antiviral drug discovery in human liver-derived organoids" 3428:

Geiser P, Di Martino ML, Samperio Ventayol P, Eriksson J, Sima E, Al-Saffar AK, et al. (January 2021). Sperandio V (ed.).

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Hepatic organoid. A recent study showed the usefulness of the technology for identifying novel medication for the treatment of

2831:"Intestinal organoids containing poly(lactic-co-glycolic acid) nanoparticles for the treatment of inflammatory bowel diseases" 6714: 5437: 4359: 502:

is required. In addition, the polarity of intestinal organoids can be inverted, and they can even be dissociated into single

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variant giving rise to a non-viable mRNA transcript was corrected by CRISPR. Patient kidney organoids demonstrated abnormal

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of interest. However, to model infection via a more natural route starting from the intestinal lumen, microinjection of the

3611:"A primary human macrophage-enteroid co-culture model to investigate mucosal gut physiology and host-pathogen interactions" 237: 3774:"Bioengineered Human Organ-on-Chip Reveals Intestinal Microenvironment and Mechanical Forces Impacting Shigella Infection" 3430:"Salmonella enterica Serovar Typhimurium Exploits Cycling through Epithelial Cells To Colonize Human and Murine Enteroids" 751:

Cardiac organoid – In 2018 hollow cardiac organoids were made to beat, and to respond to stimuli to beat faster or slower.

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Zilova L, Weinhardt V, Tavhelidse T, Schlagheck C, Thumberger T, Wittbrodt J (July 2021). Arias AM, Stainier DY (eds.).

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van den Brink SC, Baillie-Johnson P, Balayo T, Hadjantonakis AK, Nowotschin S, Turner DA, et al. (November 2014).

1337:"Reconstitution of Complete Organs From Single-Cell Suspensions of Chick Embryos in Advanced Stages of Differentiation" 869: 1558:"Functional differentiation and alveolar morphogenesis of primary mammary cultures on reconstituted basement membrane" 553:. Gastric organoids have been generated directly from pluripotent stem cells through the temporal manipulation of the 5463:"cGMP production of patient-specific iPSCs and photoreceptor precursor cells to treat retinal degenerative blindness" 915:

Other studies have used CRISPR gene editing to correct the patient's mutation in the patient iPSC cells to create an

6494: 908:) could be related to alteration in nephron patterning arising from nephron progenitor cells lacking this important 5161:

Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, et al. (November 2017).

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stem cells are used for the creation of the organoid, the cells are usually, but not all the time, allowed to form

121: 2643:"Long-term expansion of epithelial organoids from human colon, adenoma, adenocarcinoma, and Barrett's epithelium" 6287: 5065:"Symmetry breaking, germ layer specification and axial organisation in aggregates of mouse embryonic stem cells" 4959:"Modelling kidney disease with CRISPR-mutant kidney organoids derived from human pluripotent epiblast spheroids" 4656:"Advanced Development of Primary Pancreatic Organoid Tumor Models for High-Throughput Phenotypic Drug Screening" 3978:"Metastatic tumor evolution and organoid modeling implicate TGFBR2 as a cancer driver in diffuse gastric cancer" 2207:"Differentiation of embryonic stem cells to clinically relevant populations: lessons from embryonic development" 5520:"Fish primary embryonic pluripotent cells assemble into retinal tissue mirroring in vivo early eye development" 4854:"Lung stem cell differentiation in mice directed by endothelial cells via a BMP4-NFATc1-thrombospondin-1 axis" 4754:"A microenvironment-inspired synthetic three-dimensional model for pancreatic ductal adenocarcinoma organoids" 573:

signalling pathways in three-dimensional culture conditions. Gastric organoids have also been generated using

3725:"Human Colon-on-a-Chip Enables Continuous In Vitro Analysis of Colon Mucus Layer Accumulation and Physiology" 2880:"Gut Organoid as a New Platform to Study Alginate and Chitosan Mediated PLGA Nanoparticles for Drug Delivery" 2829:

Davoudi Z, Peroutka-Bigus N, Bellaire B, Wannemuehler M, Barrett TA, Narasimhan B, et al. (April 2018).

964: 876: 347: 61: 486: 2581:

Cai H, Ao Z, Tian C, et al. (2023). "Brain organoid reservoir computing for artificial intelligence".

241: 6881: 1023: 6406:"Functional repair of CFTR by CRISPR/Cas9 in intestinal stem cell organoids of cystic fibrosis patients" 3826:

McCracken KW, Catá EM, Crawford CM, Sinagoga KL, Schumacher M, Rockich BE, et al. (December 2014).

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Qi Y, Lohman J, Bratlie KM, Peroutka-Bigus N, Bellaire B, Wannemuehler M, et al. (September 2019).

5661:"A Cytotoxic Three-Dimensional-Spheroid, High-Throughput Assay Using Patient-Derived Glioma Stem Cells" 880: 562: 5862:"Organoid cystogenesis reveals a critical role of microenvironment in human polycystic kidney disease" 4607:"Unlimited in vitro expansion of adult bi-potent pancreas progenitors through the Lgr5/R-spondin axis" 4092:"Characterization of stem/progenitor cell cycle using murine circumvallate papilla taste bud organoid" 3381:"Salmonella-infected crypt-derived intestinal organoid culture system for host-bacterial interactions" 865: 614:

expressing epithelial stem cells in three-dimensional culture conditions through the manipulation of

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A to drive the cells into a mesoendodermal identity, followed by the pharmacological upregulation of

6855: 3885:"Lgr5(+ve) stem cells drive self-renewal in the stomach and build long-lived gastric units in vitro" 1605:

Streuli CH, Schmidhauser C, Bailey N, Yurchenco P, Skubitz AP, Roskelley C, et al. (May 1995).

6728:"Vascularized and Complex Organ Buds from Diverse Tissues via Mesenchymal Cell-Driven Condensation" 3883:

Barker N, Huch M, Kujala P, van de Wetering M, Snippert HJ, van Es JH, et al. (January 2010).

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Sontheimer-Phelps A, Chou DB, Tovaglieri A, Ferrante TC, Duckworth T, Fadel C, et al. (2020).

3567:"Transport of artificial virus-like nanocarriers through intestinal monolayers via microfold cells" 2040:

Lancaster MA, Renner M, Martin CA, Wenzel D, Bicknell LS, Hurles ME, et al. (September 2013).

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Peng H, Wang C, Xu X, Yu C, Wang Q (January 2015). "An intestinal Trojan horse for gene delivery".

1785:"Dissociation of embryonic kidneys followed by reaggregation allows the formation of renal tissues" 1698: 1136: 1027: 554: 145: 78: 73: 5708:

Dayton TL, Alcala N, Moonen L, den Hartigh L, Geurts V, Mangiante L, et al. (December 2023).

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Wiley LA, Burnight ER, DeLuca AP, Anfinson KR, Cranston CM, Kaalberg EE, et al. (July 2016).

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Ren W, Lewandowski BC, Watson J, Aihara E, Iwatsuki K, Bachmanov AA, et al. (November 2014).

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Grassart A, Malardé V, Gobaa S, Sartori-Rupp A, Kerns J, Karalis K, et al. (September 2019).

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Sato T, Stange DE, Ferrante M, Vries RG, Van Es JH, Van den Brink S, et al. (November 2011).

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van de Wetering M, Francies HE, Francis JM, Bounova G, Iorio F, Pronk A, et al. (May 2015).

5016:"Nephron organoids derived from human pluripotent stem cells model kidney development and injury" 921: 677: 615: 589:, and identified that unlike the patient's primary tumor, the metastasis had both alleles of the 570: 104: 6196:

Tanigawa S, Islam M, Sharmin S, Naganuma H, Yoshimura Y, Haque F, et al. (September 2018).

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capacities. The technique for growing organoids has rapidly improved since the early 2010s, and

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Cruz NM, Song X, Czerniecki SM, Gulieva RE, Churchill AJ, Kim YK, et al. (November 2017).

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Huch M, Bonfanti P, Boj SF, Sato T, Loomans CJ, van de Wetering M, et al. (October 2013).

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Huch M, Gehart H, van Boxtel R, Hamer K, Blokzijl F, Verstegen MM, et al. (January 2015).

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Sakib S, Uchida A, Valenzuela-Leon P, Yu Y, Valli-Pulaski H, Orwig K, et al. (June 2019).

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Qi Y, Shi E, Peroutka-Bigus N, Bellaire B, Wannemuehler M, Jergens A, et al. (May 2018). "

1039: 900:, the authors concluded that renal disease frequently seen in Lowe Syndrome (global failure of 411: 5266:"Blastoid: The backstory of the formation of blastocyst-like structure solely from stem cells" 4288:

Bredenkamp N, Ulyanchenko S, O'Neill KE, Manley NR, Vaidya HJ, Blackburn CC (September 2014).

4149:"Single Lgr5- or Lgr6-expressing taste stem/progenitor cells generate taste bud cells ex vivo" 1951:

Xinaris C, Benedetti V, Rizzo P, Abbate M, Corna D, Azzollini N, et al. (November 2012).

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Nadauld LD, Garcia S, Natsoulis G, Bell JM, Miotke L, Hopmans ES, et al. (August 2014).

3162:"TNFα regulates intestinal organoids from mice with both defined and conventional microbiota" 2097:

Huch M, Dorrell C, Boj SF, van Es JH, Li VS, van de Wetering M, et al. (February 2013).

1062: 1019: 1003: 987: 925: 619: 558: 399: 6715:"Orchestrating Organoids. A guide to crafting tissues in a dish that reprise in vivo organs" 3828:"Modelling human development and disease in pluripotent stem-cell-derived gastric organoids" 3609:

Noel G, Baetz NW, Staab JF, Donowitz M, Kovbasnjuk O, Pasetti MF, et al. (March 2017).

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Schwank G, Koo BK, Sasselli V, Dekkers JF, Heo I, Demircan T, et al. (December 2013).

6042: 5873: 5811: 5474: 5310: 5222: 4970: 4910: 4810: 4543: 4160: 4103: 4043: 3839: 3679: 3668:"FOXO1 inhibition yields functional insulin-producing cells in human gut organoid cultures" 3622: 3071: 2608:

Smirnova L, Caffo BS, Gracias DH, Huang Q, Morales Pantoja IE, Tang B, et al. (2023).

2302: 2159: 2053: 1907: 1738: 1510: 1453: 1348: 1282: 783: 154: 6662:"Long-term maturation of human cortical organoids matches key early postnatal transitions" 5922:

Kim YK, Refaeli I, Brooks CR, Jing P, Gulieva RE, Hughes MR, et al. (December 2017).

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Organoids have been created for the following structures of the gastrointestinal tract:

6787: 6762: 6686: 6661: 6637: 6610: 6547: 6386: 6339: 6222: 6197: 6170: 6145: 6114: 6089: 6065: 6030: 6006: 5948: 5923: 5894: 5861: 5832: 5800:"Intestinal organoids for assessing nutrient transport, sensing and incretin secretion" 5799: 5775: 5750: 5685: 5660: 5636: 5611: 5546: 5519: 5495: 5462: 5378: 5353: 5334: 5246: 5187: 5162: 5138: 5113: 5089: 5064: 5040: 5015: 4991: 4958: 4934: 4878: 4853: 4834: 4778: 4753: 4729: 4704: 4680: 4655: 4631: 4606: 4587: 4574: 4531: 4507: 4482: 4458: 4433: 4409: 4384: 4314: 4289: 4183: 4148: 4124: 4091: 4064: 4031: 4004: 3977: 3950: 3925: 3860: 3827: 3803: 3749: 3724: 3700: 3667: 3643: 3610: 3542: 3517: 3498: 3454: 3429: 3405: 3380: 3186: 3161: 3142: 3130: 3094: 3060:"Intestinal organoids for assessing nutrient transport, sensing and incretin secretion" 3059: 3035: 3008: 2906: 2879: 2878:

Davoudi Z, Peroutka-Bigus N, Bellaire B, Jergens A, Wannemuehler M, Wang Q (May 2021).

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3648: 3588: 3547: 3502: 3490: 3459: 3410: 3328: 3279: 3240: 3211:"Rational design of oral drugs targeting mucosa delivery with gut organoid platforms" 3191: 3134: 3099: 3040: 2989: 2950: 2911: 2860: 2811: 2758: 2711: 2699: 2664: 2521: 2480: 2431: 2386: 2330: 2275: 2263: 2228: 2187: 2128: 2079: 2017: 1982: 1933: 1876: 1841: 1806: 1754: 1671: 1636: 1587: 1538: 1479: 1411: 1407: 1376: 1298: 1243: 1175: 1112: 844: 763: 475:

valuable model to study intestinal nutrient transport, drug absorption and delivery,

372: 245: 125: 65: 6390: 6247:"Best Practices for Translational Disease Modeling Using Human iPSC-Derived Neurons" 5338: 5250: 3226: 3146: 1423: 1255: 6825: 6782: 6774: 6739: 6681: 6673: 6632: 6622: 6583: 6531: 6466: 6458: 6417: 6370: 6343: 6323: 6258: 6217: 6209: 6165: 6157: 6109: 6101: 6060: 6050: 6010: 5988: 5980: 5943: 5935: 5889: 5881: 5827: 5819: 5770: 5762: 5721: 5680: 5672: 5631: 5623: 5612:"Prospective derivation of a living organoid biobank of colorectal cancer patients" 5582: 5541: 5531: 5490: 5482: 5410: 5373: 5365: 5318: 5230: 5182: 5174: 5133: 5125: 5084: 5076: 5035: 5027: 4986: 4978: 4938: 4918: 4873: 4865: 4838: 4818: 4773: 4765: 4724: 4716: 4675: 4667: 4626: 4618: 4569: 4559: 4551: 4502: 4494: 4453: 4445: 4404: 4396: 4347: 4309: 4301: 4262: 4225: 4215: 4178: 4168: 4119: 4111: 4059: 4051: 3999: 3989: 3945: 3937: 3896: 3855: 3847: 3785: 3744: 3736: 3695: 3687: 3638: 3630: 3578: 3537: 3529: 3482: 3449: 3441: 3400: 3392: 3359: 3318: 3310: 3275: 3267: 3230: 3222: 3181: 3177: 3173: 3126: 3089: 3079: 3030: 3020: 2981: 2942: 2901: 2891: 2850: 2842: 2801: 2793: 2748: 2738: 2691: 2654: 2621: 2590: 2511: 2470: 2462: 2421: 2413: 2376: 2366: 2320: 2310: 2255: 2218: 2177: 2167: 2118: 2110: 2069: 2061: 2009: 1972: 1964: 1923: 1915: 1868: 1833: 1796: 1766: 1746: 1663: 1626: 1618: 1577: 1569: 1528: 1518: 1469: 1461: 1403: 1366: 1356: 1290: 1235: 1167: 1102: 1094: 1052: 905: 481: 318: 149: 53: 4290:"An organized and functional thymus generated from FOXN1-reprogrammed fibroblasts" 3924:

Li X, Nadauld L, Ootani A, Corney DC, Pai RK, Gevaert O, et al. (July 2014).

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Gastrointestinal organoids refer to organoids that recapitulate structures of the

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Zhang FL, Hu Z, Wang YF, Zhang WJ, Zhou BW, Sun QS, et al. (October 2023).

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Proceedings of the National Academy of Sciences of the United States of America

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An intestinal organoid (Minigut) grows up in 7 days. The scale bars are 200 μm.

503: 495: 490: 249: 163: 49: 6587: 6495:"World's first mini organ transportation to a patient with ulcerative colitis" 5322: 5234: 4720: 4266: 3994: 3025: 2727:"Effects of six common dietary nutrients on murine intestinal organoid growth" 2626: 2609: 6870: 6763:"Organoids and the genetically encoded self-assembly of embryonic stem cells" 6627: 5766: 5676: 5114:"Organoids and the genetically encoded self-assembly of embryonic stem cells" 4671: 2545: 2289:

Li ML, Aggeler J, Farson DA, Hatier C, Hassell J, Bissell MJ (January 1987).

1667: 1497:

Li ML, Aggeler J, Farson DA, Hatier C, Hassell J, Bissell MJ (January 1987).

1057: 1035: 948: 944: 916: 832: 757: 713: 566: 476: 339: 175: 141: 5571:"A Living Biobank of Breast Cancer Organoids Captures Disease Heterogeneity" 5283: 4400: 4173: 1239: 1081:

Zhao Z, Chen X, Dowbaj AM, Sljukic A, Bratlie K, Lin L, et al. (2022).

304:

its cells are grouped together and spatially organized, similar to an organ.

190:

cells that self-organize into distinctive layers. In 2009 the Laboratory of

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started with one of the first dissociation-reaggregation experiments where

5414: 4530:

Li P, Li Y, Wang Y, Liu J, Lavrijsen M, Li Y, et al. (January 2022).

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it is capable of recapitulating some specific function of the organ (e.g.

153:

for the structural organization was made possible with the development of

148:

can organize into different structures resembling those found in multiple

4564: 3427: 2828: 2449:

Ran FA, Hsu PD, Wright J, Agarwala V, Scott DA, Zhang F (November 2013).

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niche, making them the first organoids. In 2010, Mathieu Unbekandt &

211: 199: 5536: 5369: 4922: 4822: 3851: 3396: 2985: 2846: 2797: 2114: 2065: 1801: 1784: 1750: 244:

established a protocol starting from pluripotent stem cells to generate

5178: 5080: 4982: 3722: 3691: 3583: 3566: 2896: 2426: 1872: 1607:"Laminin mediates tissue-specific gene expression in mammary epithelia" 897: 732: 586: 471: 390: 314: 183: 83: 22: 5993: 5939: 5823: 5486: 4287: 4115: 4055: 3634: 3084: 1919: 1837: 1270: 896:

is a well characterised marker of nephron progenitor cells in the cap

509: 28: 5885: 5609: 5031: 4385:"Formation of organotypic testicular organoids in microwell culture†" 824: 789: 549:

Gastric organoids recapitulate at least partly the physiology of the

395: 386: 298: 294: 136:

was realized early on with the observation that when stem cells form

129: 69: 6830: 6805: 6327: 5160: 4434:"Organoid culture systems for prostate epithelial and cancer tissue" 4305: 4205: 3941: 2877: 2695: 2562:"Scientists 'may have crossed ethical line' in growing human brains" 2013: 6760: 6608: 5517: 5111: 2725:

Cai T, Qi Y, Jergens A, Wannemuehler M, Barrett TA, Wang Q (2018).

2541:"Miniature brains grown in the lab have human-like neural activity" 2042:"Cerebral organoids model human brain development and microcephaly" 828: 701:

as it allows to allows to recapitulate the entire viral life cycle.

668:

Thymic organoids recapitulate at least partly the architecture and

499: 415: 377: 327: 323: 137: 99: 45: 6313: 6090:"Simultaneous reprogramming and gene editing of human fibroblasts" 3486: 3345: 2352: 1604: 1556:

Barcellos-Hoff MH, Aggeler J, Ram TG, Bissell MJ (February 1989).

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such report, kidney organoids derived from iPSC of a patient with

623: 529:

models combine intestinal organoids with other cell types such as

6521: 2970:"Manipulate intestinal organoids with niobium carbide nanosheets" 2500:"Cerebral organoids: ethical issues and consciousness assessment" 1689: 1440:

Yi SA, Zhang Y, Rathnam C, Pongkulapa T, Lee KB (November 2021).

1010:

whose cells were used for the organoid, was carried out in 2022.

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Turner DA, Baillie-Johnson P, Martinez Arias A (February 2016).

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Turner DA, Baillie-Johnson P, Martinez Arias A (February 2016).

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organoid, and examples of how it has been utilized in research.

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that mimic the developing human brain's cellular organization.

187: 6806:"The boom in mini stomachs, brains, breasts, kidneys and more" 1442:"Bioengineering Approaches for the Advanced Organoid Research" 6028: 5707: 5354:"Organoids to model the endometrium: implantation and beyond" 4702: 3825: 2775: 581:. Gastric organoids have been used as model for the study of 489:, and research done with them demonstrated the positive role 462: 382: 179: 167: 6725: 6195: 6029:

Hsieh WC, Ramadesikan S, Fekete D, Aguilar RC (2018-02-14).

5751:"Astrocytes and human artificial blood-brain barrier models" 5351: 4382: 3006: 2681: 1823: 5970: 5460: 4705:"Dynamic changes during the treatment of pancreatic cancer" 4653: 4089: 2928: 2039: 1950: 1653: 999:

mucosal damages on mice with ischemia-reperfusion injury.

970:

Things such as epithelial metabolism can also be modelled.

639: 635: 631: 611: 574: 466: 203: 5352:

Rawlings TM, Makwana K, Tryfonos M, Lucas ES (July 2021).

4146: 3975: 3564: 1271:"A New Method by Which Sponges May Be Artificially Reared" 1158:

Mullard A (March 2023). "Mini-organs attract big pharma".

1018:

Organoids offer researchers an exceptional model to study

934:

were compared to an isogenic control organoid in which an

680:

with mouse thymic organoids resulted in the production of

435:

gives rise to the small intestines and the ascending colon

6403: 6143: 5859: 5658: 4800: 4480: 2607: 5568: 5212: 5013: 4751: 4431: 4029: 3821: 3819: 3817: 2724: 1439: 1221: 1219: 1217: 1215: 1213: 1211: 1209: 739:

gene expression patterns along the anteroposterior axis.

6609:

Martinez-Morales JR, Cavodeassi F, Bovolenta P (2017).

2640: 1207: 1205: 1203: 1201: 1199: 1197: 1195: 1193: 1191: 1189: 1013: 831:

hormones. This is of great relevance in the context of

517:

In order to more accurately recapitulate the intestine

232:

Other significant early advancements included in 2013,

6659: 5798:

Zietek T, Rath E, Haller D, Daniel H (November 2015).

5797: 5748: 3971: 3969: 3729:

Cellular and Molecular Gastroenterology and Hepatology

3608: 3515: 3159: 3058:

Zietek T, Rath E, Haller D, Daniel H (November 2015).

3057: 1043:

have some features of human fetal development stages.

875:

As first developed by Takahashi and Yamanaka in 2007,

4252: 3923: 3814: 3208: 3116: 2288: 1999: 1496: 1080: 5921: 3665: 3476: 2448: 2096: 1778: 1776: 1186: 385:. Cerebral organoids are created by culturing human 313:

Organoid formation generally requires culturing the

166:

and David Odde showed the self-assembly of vascular

132:

biology, the potential of stem cells to form organs

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as new biomaterials to alter intestinal stem cells"

2348: 2346: 2344: 2245: 1893: 1034:to expanding our understanding of the processes of 494:be infected in bulk by simply mixing them with the 4851: 4085: 4083: 3166:International Journal of Biological Macromolecules 963:of kidney organoids generated from a patient with 442:gives rise to the rectum and the rest of the colon 389:in a three-dimensional structure using rotational 6244: 6191: 6189: 4341: 4025: 4023: 2451:"Genome engineering using the CRISPR-Cas9 system" 2403: 2145: 1773: 1225: 322:medium can be made using an extracellular matrix 6868: 6444: 4337: 4335: 4333: 2974:Journal of Biomedical Materials Research. Part A 2967: 2835:Journal of Biomedical Materials Research. Part A 2786:Journal of Biomedical Materials Research. Part A 2497: 2341: 116:was observed for the first time by co-culturing 21:For Organoids appearing in an anime series, see 6245:Engle SJ, Blaha L, Kleiman RJ (November 2018). 6139: 6137: 6135: 6133: 5208: 5206: 4080: 3379:Zhang YG, Wu S, Xia Y, Sun J (September 2014). 1894:Lawrence ML, Chang CH, Davies JA (March 2015). 1782: 1724: 1722: 1720: 1718: 1716: 835:diseases as well as metabolic diseases such as 40:is a miniaturised and simplified version of an 32:Intestinal organoid grown from Lgr5+ stem cells 16:Miniaturized and simplified version of an organ 6355: 6353: 6309: 6307: 6186: 6024: 6022: 6020: 5917: 5915: 5913: 5855: 5853: 5851: 4020: 1393: 6602: 6569: 6567: 6565: 6563: 6561: 6088:Howden SE, Thomson JA, Little MH (May 2018). 4952: 4950: 4948: 4330: 3378: 3013:Frontiers in Bioengineering and Biotechnology 1957:Journal of the American Society of Nephrology 1647: 1038:, and the field of developmental biology. In 428:gives rise to the oral cavity and the stomach 414:. The gastrointestinal tract arises from the 6130: 5652: 5203: 4794: 4696: 4647: 4598: 4474: 4246: 3604: 3602: 2139: 2035: 2033: 2031: 1858: 1713: 1006:of an organoid into a human, a patient with 405: 68:in three-dimensional culture owing to their 6350: 6304: 6285: 6017: 5910: 5848: 4529: 3659: 3352:Biology of Blood and Marrow Transplantation 2204: 1334: 6573: 6558: 4945: 2933:Study of Telluride Nanowires in Minigut". 2146:Shkumatov A, Baek K, Kong H (April 2014). 1826:ACS Biomaterials Science & Engineering 1315: 521:, co-cultures of intestinal organoids and 282:it has multiple organ-specific cell types; 278:, and exhibits the following properties: 6829: 6786: 6743: 6685: 6636: 6626: 6470: 6421: 6262: 6221: 6169: 6113: 6064: 6054: 5992: 5947: 5893: 5831: 5774: 5755:Bosnian Journal of Basic Medical Sciences 5725: 5684: 5635: 5586: 5545: 5535: 5494: 5377: 5186: 5137: 5088: 5039: 4990: 4877: 4777: 4728: 4679: 4630: 4573: 4563: 4506: 4457: 4408: 4313: 4229: 4219: 4182: 4172: 4123: 4063: 4003: 3993: 3949: 3900: 3859: 3789: 3748: 3699: 3642: 3599: 3582: 3541: 3453: 3404: 3363: 3322: 3234: 3185: 3093: 3083: 3034: 3024: 2905: 2895: 2854: 2805: 2752: 2742: 2658: 2625: 2580: 2515: 2498:Lavazza A, Massimini M (September 2018). 2474: 2425: 2380: 2370: 2324: 2314: 2222: 2181: 2171: 2122: 2073: 2028: 1976: 1927: 1800: 1630: 1581: 1532: 1522: 1473: 1435: 1433: 1370: 1360: 1106: 626:. This organoid culture, however, lacks 6803: 6292:Special Issue of the Metabolites Journal 5345: 3296: 1026:, there have been great advancements in 973: 860:, where patient cells were used to make 508: 27: 5435: 4199: 1157: 993: 678:hematopoietic or bone marrow stem cells 6869: 6856:"Organoid-based regenerative medicine" 5263: 2559: 2538: 1430: 1268: 451: 1783:Unbekandt M, Davies JA (March 2010). 1335:Weiss P, Taylor AC (September 1960). 1129: 1028:directing pluripotent stem cells fate 485:were used to study the nature of the 170:organoid maintained for over 50 days 2935:Journal of Biomedical Nanotechnology 2205:Murry CE, Keller G (February 2008). 1688: 1014:As a model for developmental biology 850: 366: 238:Institute of Molecular Biotechnology 6712: 6500:Tokyo Medical and Dental University 4255:International Reviews of Immunology 819:other organs are not recapitulated 605: 544: 13: 6705: 6150:American Journal of Human Genetics 3131:10.1016/j.biomaterials.2018.12.006 870:focal segmental glomerulosclerosis 795:Blood-brain barrier (BBB) organoid 525:have been developed. 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While research on 784:Neuroendocrine tumor 748:Endometrial organoid 146:differentiated cells 58:embryonic stem cells 6822:2015Natur.523..520W 6666:Nature Neuroscience 6047:2018PLoSO..1392635H 5878:2017NatMa..16.1112C 5816:2015NatSR...516831Z 5537:10.7554/eLife.66998 5479:2016NatSR...630742W 5370:10.1530/RAF-21-0023 5315:2018Natur.557..106R 5227:2018Natur.562..272B 4975:2015NatCo...6.8715F 4923:10.1038/nature15695 4915:2015Natur.526..564T 4823:10.1038/nature06196 4815:2007Natur.449.1003B 4809:(7165): 1003–1007. 4715:(19): 14764–14790. 4548:2022SciA....8.5908L 4294:Nature Cell Biology 4165:2014PNAS..11116401R 4159:(46): 16401–16406. 4108:2015NatSR...517185A 4048:2013NatSR...3E3224H 3852:10.1038/nature13863 3844:2014Natur.516..400M 3684:2014NatCo...5.4242B 3627:2017NatSR...745270N 3577:(30): 16339–16347. 3528:(9): 2509–2520.e4. 3397:10.14814/phy2.12147 3215:Bioactive Materials 3076:2015NatSR...516831Z 2986:10.1002/jbm.a.37032 2847:10.1002/jbm.a.36305 2798:10.1002/jbm.a.36715 2307:1987PNAS...84..136L 2164:2014PLoSO...994764S 2115:10.1038/nature11826 2066:10.1038/nature12517 2058:2013Natur.501..373L 1912:2015NatSR...5E9092L 1802:10.1038/ki.2009.482 1751:10.1038/nature07935 1743:2009Natur.459..262S 1692:(August 28, 2013). 1515:1987PNAS...84..136L 1458:2021AdM....3307949Y 1353:1960PNAS...46.1177W 1287:1907Sci....25..912W 949:cell-cell junctions 723:Epithelial organoid 704:Pancreatic organoid 691:Testicular organoid 577:expressing stomach 452:Intestinal organoid 94:Attempts to create 6882:Tissue engineering 5804:Scientific Reports 5467:Scientific Reports 5288:Nicolas Rivron Lab 5179:10.1242/dev.150391 5081:10.1242/dev.113001 4983:10.1038/ncomms9715 4096:Scientific Reports 4036:Scientific Reports 3692:10.1038/ncomms5242 3615:Scientific Reports 3584:10.1039/d0nr03680c 3064:Scientific Reports 2897:10.3390/md19050282 1900:Scientific Reports 1873:10.1039/c4nr06377e 1656:Tissue Engineering 1446:Advanced Materials 1008:ulcerative colitis 862:cerebral organoids 841:insulin resistance 653:tooth regeneration 515: 254:Hubrecht Institute 246:cerebral organoids 234:Madeline Lancaster 196:Hubrecht Institute 34: 6816:(7562): 520–522. 6202:Stem Cell Reports 5940:10.1002/stem.2707 5934:(12): 2366–2378. 5872:(11): 1112–1119. 5824:10.1038/srep16831 5487:10.1038/srep30742 5309:(7703): 106–111. 5221:(7726): 272–276. 5173:(21): 3894–3906. 5075:(22): 4231–4242. 5026:(11): 1193–1200. 4909:(7574): 564–568. 4617:(20): 2708–2721. 4361:978-1-59745-395-0 4116:10.1038/srep17185 4056:10.1038/srep03224 3838:(7531): 400–404. 3784:(3): 435–444.e4. 3635:10.1038/srep45270 3085:10.1038/srep16831 2690:(10): 1225–1227. 2589:(12): 1032–1039. 2461:(11): 2281–2308. 2109:(7436): 247–250. 2052:(7467): 373–379. 1963:(11): 1857–1868. 1920:10.1038/srep09092 1867:(10): 4354–4360. 1838:10.1021/ab500041d 1737:(7244): 262–265. 1234:(6194): 1247125. 912:gene expression. 851:Models of disease 792:(Myelin organoid) 764:Colorectal cancer 694:Prostate organoid 373:cerebral organoid 367:Cerebral organoid 126:Malcolm Steinberg 6889: 6863: 6862:. 30 March 2024. 6843: 6833: 6800: 6790: 6757: 6747: 6722: 6700: 6699: 6689: 6657: 6651: 6650: 6640: 6630: 6606: 6600: 6599: 6571: 6556: 6555: 6524:Nature Protocols 6519: 6513: 6512: 6510: 6508: 6491: 6485: 6484: 6474: 6442: 6436: 6435: 6425: 6401: 6395: 6394: 6369:(344): 344ra84. 6357: 6348: 6347: 6311: 6302: 6301: 6299: 6298: 6283: 6277: 6276: 6266: 6242: 6236: 6235: 6225: 6193: 6184: 6183: 6173: 6141: 6128: 6127: 6117: 6094:Nature Protocols 6085: 6079: 6078: 6068: 6058: 6026: 6015: 6014: 5996: 5968: 5962: 5961: 5951: 5919: 5908: 5907: 5897: 5886:10.1038/nmat4994 5866:Nature Materials 5857: 5846: 5845: 5835: 5795: 5789: 5788: 5778: 5746: 5740: 5739: 5729: 5705: 5699: 5698: 5688: 5656: 5650: 5649: 5639: 5607: 5601: 5600: 5590: 5566: 5560: 5559: 5549: 5539: 5515: 5509: 5508: 5498: 5458: 5452: 5451: 5449: 5448: 5433: 5427: 5426: 5398: 5392: 5391: 5381: 5349: 5343: 5342: 5298: 5292: 5291: 5280: 5274: 5273: 5261: 5255: 5254: 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3457: 3425: 3419: 3418: 3408: 3376: 3370: 3369: 3367: 3343: 3337: 3336: 3326: 3309:(7): 3402–3413. 3294: 3288: 3287: 3278: 10840714. 3266:(12): e2300103. 3260:Advanced Biology 3255: 3249: 3248: 3238: 3206: 3200: 3199: 3189: 3157: 3151: 3150: 3114: 3108: 3107: 3097: 3087: 3055: 3049: 3048: 3038: 3028: 3004: 2998: 2997: 2965: 2959: 2958: 2926: 2920: 2919: 2909: 2899: 2875: 2869: 2868: 2858: 2826: 2820: 2819: 2809: 2792:(9): 1886–1897. 2778:"Vitamin C and B 2773: 2767: 2766: 2756: 2746: 2722: 2716: 2715: 2679: 2673: 2672: 2662: 2653:(5): 1762–1772. 2647:Gastroenterology 2638: 2632: 2631: 2629: 2605: 2599: 2598: 2578: 2572: 2571: 2557: 2551: 2550: 2549:. 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Netherlands. 5275: 5256: 5202: 5153: 5124:(2): 181–191. 5104: 5055: 5006: 4944: 4893: 4864:(3): 440–455. 4844: 4793: 4764:(1): 110–119. 4744: 4695: 4666:(6): 574–584. 4660:SLAS Discovery 4646: 4597: 4522: 4473: 4444:(2): 347–358. 4424: 4375: 4360: 4329: 4300:(9): 902–908. 4280: 4245: 4198: 4139: 4079: 4019: 3982:Genome Biology 3965: 3936:(7): 769–777. 3916: 3889:Cell Stem Cell 3875: 3813: 3764: 3735:(3): 507–526. 3715: 3658: 3598: 3557: 3508: 3481:(151): 59610. 3469: 3420: 3371: 3358:(2): S53–S54. 3338: 3289: 3250: 3201: 3152: 3109: 3050: 2999: 2980:(4): 479–487. 2960: 2941:(5): 978–986. 2921: 2870: 2841:(4): 876–886. 2821: 2779: 2768: 2717: 2674: 2633: 2600: 2573: 2552: 2531: 2510:(9): 606–610. 2490: 2441: 2412:(4): 663–676. 2396: 2340: 2301:(1): 136–140. 2281: 2254:(3): 245–249. 2238: 2217:(4): 661–680. 2197: 2138: 2089: 2027: 2008:(4): 618–623. 1992: 1943: 1886: 1851: 1816: 1795:(5): 407–416. 1772: 1712: 1681: 1646: 1617:(3): 591–603. 1597: 1568:(2): 223–235. 1548: 1509:(1): 136–140. 1489: 1429: 1402:(3): 317–330. 1386: 1327: 1318:Rev. 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Biol 1308: 1261: 1185: 1166:(3): 175–176. 1150: 1122: 1072: 1070: 1067: 1066: 1065: 1060: 1055: 1048: 1045: 1015: 1012: 995: 992: 975: 972: 852: 849: 811: 810:Basic research 808: 797: 796: 793: 787: 774: 773: 767: 761: 755: 752: 749: 746: 740: 730: 727: 724: 720: 719: 718: 717: 706: 705: 702: 695: 692: 688: 687: 686: 685: 663: 662: 659: 656: 647: 644: 607: 604: 546: 543: 496:enteropathogen 453: 450: 446: 445: 444: 443: 436: 429: 407: 404: 402:technologies. 394:could develop 368: 365: 359: 356: 310: 307: 306: 305: 302: 283: 270: 267: 250:Meritxell Huch 164:Yaakov Nahmias 91: 88: 15: 9: 6: 4: 3: 2: 6894: 6883: 6880: 6878: 6875: 6874: 6872: 6861: 6857: 6853: 6852: 6841: 6837: 6832: 6827: 6823: 6819: 6815: 6811: 6807: 6802: 6798: 6794: 6789: 6784: 6780: 6776: 6772: 6768: 6764: 6759: 6755: 6751: 6746: 6741: 6737: 6733: 6729: 6724: 6720: 6719:The Scientist 6716: 6711: 6710: 6697: 6693: 6688: 6683: 6679: 6675: 6671: 6667: 6663: 6656: 6648: 6644: 6639: 6634: 6629: 6624: 6620: 6616: 6612: 6605: 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2615: 2611: 2604: 2596: 2592: 2588: 2584: 2577: 2570:. p. 15. 2569: 2568: 2563: 2556: 2548: 2547: 2546:New Scientist 2542: 2535: 2527: 2523: 2518: 2513: 2509: 2505: 2501: 2494: 2486: 2482: 2477: 2472: 2468: 2464: 2460: 2456: 2452: 2445: 2437: 2433: 2428: 2423: 2419: 2415: 2411: 2407: 2400: 2392: 2388: 2383: 2378: 2373: 2368: 2364: 2360: 2356: 2349: 2347: 2345: 2336: 2332: 2327: 2322: 2317: 2312: 2308: 2304: 2300: 2296: 2292: 2285: 2277: 2273: 2269: 2265: 2261: 2257: 2253: 2249: 2242: 2234: 2230: 2225: 2220: 2216: 2212: 2208: 2201: 2193: 2189: 2184: 2179: 2174: 2169: 2165: 2161: 2158:(4): e94764. 2157: 2153: 2149: 2142: 2134: 2130: 2125: 2120: 2116: 2112: 2108: 2104: 2100: 2093: 2085: 2081: 2076: 2071: 2067: 2063: 2059: 2055: 2051: 2047: 2043: 2036: 2034: 2032: 2023: 2019: 2015: 2011: 2007: 2003: 1996: 1988: 1984: 1979: 1974: 1970: 1966: 1962: 1958: 1954: 1947: 1939: 1935: 1930: 1925: 1921: 1917: 1913: 1909: 1905: 1901: 1897: 1890: 1882: 1878: 1874: 1870: 1866: 1862: 1855: 1847: 1843: 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1100: 1096: 1092: 1088: 1084: 1077: 1073: 1064: 1061: 1059: 1058:Organ culture 1056: 1054: 1051: 1050: 1044: 1041: 1037: 1036:organogenesis 1032: 1029: 1025: 1021: 1011: 1009: 1005: 1000: 991: 989: 983: 981: 971: 968: 966: 962: 958: 954: 950: 946: 945:cell polarity 941: 937: 933: 932: 928:mutations in 927: 923: 918: 913: 911: 907: 903: 899: 895: 891: 890:golgi complex 886: 882: 881:Lowe Syndrome 878: 873: 871: 867: 863: 859: 848: 846: 842: 838: 834: 833:malabsorption 830: 826: 822: 817: 806: 804: 794: 791: 788: 785: 782: 781: 779: 771: 768: 765: 762: 759: 758:Breast cancer 756: 753: 750: 747: 744: 741: 738: 734: 731: 728: 726:Lung organoid 725: 722: 721: 715: 714:David Tuveson 710: 709: 708: 707: 703: 700: 696: 693: 690: 689: 683: 679: 675: 671: 667: 666: 665: 664: 660: 657: 654: 650: 649: 643: 641: 637: 633: 629: 625: 621: 617: 613: 603: 601: 597: 592: 588: 584: 580: 576: 572: 568: 567:retinoic acid 564: 560: 556: 552: 542: 540: 536: 532: 528: 524: 520: 511: 507: 505: 501: 497: 492: 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In 2008, 6877:Stem cells 6871:Categories 6297:2022-10-16 5994:2433/49782 5928:Stem Cells 5530:: e66998. 5447:2018-06-30 5284:"Blastoid" 4709:Oncotarget 3988:(8): 428. 3019:: 577656. 2890:(5): 282. 1324:: 220–250. 1069:References 898:mesenchyme 892:. Because 825:stem cells 733:Gastruloid 472:epithelial 391:bioreactor 352:CRISPR/Cas 315:stem cells 293:activity, 269:Properties 184:stem cells 84:laboratory 6767:BioEssays 6582:: 23–28. 6552:246488596 5810:: 16831. 5473:: 30742. 5118:BioEssays 4592:246069868 4102:: 17185. 3808:201868491 3571:Nanoscale 3503:203377662 2712:205388154 2614:Front Sci 2276:210922708 1861:Nanoscale 961:glomeruli 790:Myelinoid 396:sentience 299:excretion 295:endocrine 138:teratomas 130:stem cell 44:produced 6840:26223610 6797:26666846 6754:25891906 6696:33619405 6647:29326547 6596:25033469 6544:35110738 6481:37880227 6472:10600233 6432:24315439 6391:19462535 6383:27334259 6336:23727931 6273:30465765 6232:30174315 6180:29706353 6124:29622803 6075:29444177 6035:PLoS One 6003:18035408 5958:28905451 5904:28967916 5842:26582215 5785:35366791 5736:38086335 5695:29750582 5646:25957691 5597:29224780 5556:34252023 5505:27471043 5423:18333774 5388:35118399 5339:13749109 5331:29720634 5270:The Node 5251:52915553 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