Self-organization of axial polarity, inside-out layer pattern, and species-specific progenitor dynamics in human ES cell–derived neocortex

Cerebral organoids model human brain development and microcephaly

Lancaster, M. A., Renner, M., Martin, C.-A., Wenzel, D., Bicknell, L. S., Hurles, M. E., Homfray, T., Penninger, J. M., Jackson, A. P., & Knoblich, J. A. (2013) · Nature · doi:10.1038/nature12517 · Cited by 5,372

A ROCK inhibitor permits survival of dissociated human embryonic stem cells

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Self-Organized Formation of Polarized Cortical Tissues from ESCs and Its Active Manipulation by Extrinsic Signals

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Self-formation of optic cups and storable stratified neural retina from human ESCs

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Neurogenic radial glia in the outer subventricular zone of human neocortex

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Neurons in Rhesus Monkey Visual Cortex: Systematic Relation between Time of Origin and Eventual Disposition

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Development of the human cerebral cortex: Boulder Committee revisited

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Developmental history of the transient subplate zone in the visual and somatosensory cortex of the macaque monkey and human brain

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OSVZ progenitors of human and ferret neocortex are epithelial-like and expand by integrin signaling

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iPSC-derived neural organoids in dementia research: Recent advances and future directions

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Longitudinal, label-free, high-resolution imaging of glioblastoma spheroid response to therapy: a translational tool for preclinical evaluation of chemotherapy, radiation, and immunotherapy

Serafini, C. E., Davarzani, A., Cappabianca, D., Li, Z., Collazos, D. B., Mamaghani, D., Mao, L., Saha, K., Karumbaiah, L., & Robles, F. E. (2025) · Optica · doi:10.1364/optica.574860 · Cited by 7

Reconstructing human corticogenesis: Insights from cerebral organoids into neurodevelopment and disease modeling

Tenreiro, M. F., & Muotri, A. R. (2026) · Developmental Cell · doi:10.1016/j.devcel.2026.02.018 · Cited by 6

Autism mutations rewire protein interaction networks to drive neurodevelopmental pathology

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Organoids Modeling Stroke in a Petri Dish

Giorgi, C., Castelli, V., d’Angelo, M., & Cimini, A. (2024) · Biomedicines · doi:10.3390/biomedicines12040877 · Cited by 7

Recent advances and applications of human brain models

Nishimura, K., Osaki, H., Tezuka, K., Nakashima, D., Numata, S., & Masamizu, Y. (2024) · Frontiers in Neural Circuits · doi:10.3389/fncir.2024.1453958 · Cited by 5

Advances in human pluripotent stem cell reporter systems

Puspita, L., Juwono, V. B., & Shim, J. (2024) · iScience · doi:10.1016/j.isci.2024.110856 · Cited by 6

Multi-omic analysis of guided and unguided forebrain organoids reveals differences in cellular composition and metabolic profiles

Øhlenschlæger, M. S., Jensen, P., Havelund, J. F., Schmidt, S. I., Mohamed, F. A., Sutcliffe, M. A., Elmkvist, S. B., Criscuolo, L., Wingett, S., Chiaradia, I., et al. (2026) · Cell Reports Methods · doi:10.1016/j.crmeth.2025.101295 · Cited by 5

IGF-2/IGF-1R signaling has distinct effects on Sox1, Irx3, and Six3 expressions during ES cell derived-neuroectoderm development in vitro

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Development of a 3-D Organoid System Using Human Induced Pluripotent Stem Cells to Model Idiopathic Autism

Lunden, J. W., Durens, M., Nestor, J. E., Niescier, R. F., Herold, K., Brandenburg, C., Lin, Y., Blatt, G. J., & Nestor, M. W. (2020) · Advances in neurobiology · doi:10.1007/978-3-030-45493-7_10 · Cited by 5

2D and 3D Human Induced Pluripotent Stem Cell-Based Models to Dissect Primary Cilium Involvement during Neocortical Development

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Diseased, differentiated and difficult: Strategies for improved engineering of in vitro neurological systems

Elder, N. H., Fattahi, F., McDevitt, T. C., & Zholudeva, L. V. (2022) · Frontiers in Cellular Neuroscience · doi:10.3389/fncel.2022.962103 · Cited by 6

Targeted Microinjection and Electroporation of Primate Cerebral Organoids for Genetic Modification

Tynianskaia, L., Eşiyok, N., Huttner, W. Β., & Heide, M. (2023) · Journal of Visualized Experiments · doi:10.3791/65176 · Cited by 7

Organoids containing neural-like cells derived from chicken iPSCs respond to poly:IC through the RLR family

Katayama, M., Onuma, M., Kato, N., Nakajima, N., & Fukuda, T. (2023) · PLoS ONE · doi:10.1371/journal.pone.0285356 · Cited by 5

An Exploration of Organoid Technology: Present Advancements, Applications, and Obstacles

Mishra, I., Gupta, K., Mishra, R., Chaudhary, K., & Sharma, V. (2023) · Current Pharmaceutical Biotechnology · doi:10.2174/0113892010273024230925075231 · Cited by 5

Causes of microcephaly in human—theoretical considerations

Heide, M., & Huttner, W. Β. (2023) · Frontiers in Neuroscience · doi:10.3389/fnins.2023.1306166 · Cited by 5

Recapitulating Hypothalamus and Pituitary Development Using Embryonic Stem/Induced Pluripotent Stem Cells

Suga, H. (2016) · Research and perspectives in endocrine interactions · doi:10.1007/978-3-319-41603-8_4 · Cited by 5

Lab-grown mini-brains upgraded

Yang, L., & Ng, H. (2017) · Nature Cell Biology · doi:10.1038/ncb3601 · Cited by 5

Stem Cells to Inform the Neurobiology of Mental Illness

Johnstone, M., Hillary, R. F., & Clair, D. S. (2018) · Current topics in behavioral neurosciences · doi:10.1007/7854_2018_57 · Cited by 5

A Static Self-Directed Method for Generating Brain Organoids from Human Embryonic Stem Cells

Boisvert, E. M., Means, R. E., Michaud, M., Thomson, J. J., Madri, J. A., & Katz, S. G. (2020) · Journal of Visualized Experiments · doi:10.3791/60379 · Cited by 5

Energy Metabolism and Intracellular pH Alteration in Neural Spheroids Carrying Down Syndrome

Kashirina, A. S., Gavrina, A. I., Kryukov, E., Elagin, V. V., Kolesova, Y. S., Artyuhov, A., Momotyuk, E., Abdyev, V. K., Meshcheryakova, N., Вадимовна, З. Е., et al. (2021) · Biomedicines · doi:10.3390/biomedicines9111741 · Cited by 5

Recent Development of Brain Organoids for Biomedical Application

Kim, H., Jang, E. J., Sankpal, N. V., Patel, M., & Patel, R. (2022) · Macromolecular Bioscience · doi:10.1002/mabi.202200346 · Cited by 5

Making pituitary hormone-producing cells in a dish [Review]

Suga, H. (2016) · Endocrine Journal · doi:10.1507/endocrj.ej16-0232 · Cited by 6

Alzheimer’s Disease: Insights from Genetic Mouse Models and Current Advances in Human IPSC-Derived Neurons

Harasta, A. E., & Ittner, L. M. (2017) · Advances in neurobiology · doi:10.1007/978-3-319-57193-5_1 · Cited by 5

Self-organizing cortex generated from human iPSCs with combination of FGF2 and ambient oxygen

Eguchi, N., Sora, I., & Muguruma, K. (2018) · Biochemical and Biophysical Research Communications · doi:10.1016/j.bbrc.2018.03.049 · Cited by 5

Generation of Various Telencephalic Regions from Human Embryonic Stem Cells in Three-Dimensional Culture

Kadoshima, T., Sakaguchi, H., & Eiraku, M. (2017) · Methods in molecular biology · doi:10.1007/978-1-4939-6949-4_1 · Cited by 4

Transcriptional evaluation of the developmental accuracy, reproducibility and robustness of kidney organoids derived from human pluripotent stem cellsPreprint

Phipson, B., Er, P. X., Hale, L. J., Yen, D., Lawlor, K. T., Takasato, M., Sun, J. Q. J., Wolvetang, E. J., Oshlack, A., & Little, M. H. (2017) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/238428 · Cited by 5

Protocol for minimizing necrotic core formation in iPSC-derived cortical organoids

Mamun, M. M. A., & Chetty, S. (2026) · STAR Protocols · doi:10.1016/j.xpro.2026.104609 · Cited by 4

Patient-derived organoids predict personalized drug response and reveal alternative therapeutic options in glioblastoma

Park, J., Qiu, M., Cho, E. J., Seo, S., Moon, J.-H., Kim, M. U., Mun, H., Yun, H., Park, H., 임아림, et al. (2026) · Cell Reports Medicine · doi:10.1016/j.xcrm.2026.102850 · Cited by 5

Revealing the Impact of Mitochondrial Fitness During Early Neural Development Using Human Brain Organoids

Romero-Morales, A. I., & Gama, V. (2022) · Frontiers in Molecular Neuroscience · doi:10.3389/fnmol.2022.840265 · Cited by 5

Space Renaissance and Neurodegeneration

Marotta, D., Stoudemire, J., Clements, T., Loring, J. F., Grisanti, P., Noggle, S., & Fossati, V. (2022) · Springer eBooks · doi:10.1007/978-3-031-18440-6_9 · Cited by 3

Innovative explorations: unveiling the potential of organoids for investigating environmental pollutant exposure

Zhang, Y., Liu, K., He, H., Xiao, H., Fang, Z., Chen, X., & Li, H. (2024) · Environmental Science and Pollution Research · doi:10.1007/s11356-024-32256-7 · Cited by 6

Human stem cell models to unravel brain cancer

Dave, B. M., & Tailor, J. K. (2024) · BMC Cancer · doi:10.1186/s12885-024-13187-5 · Cited by 4

Advancements in brain organoid models for neurodegenerative disease research

Park, S., & Park, J. (2024) · Organoid · doi:10.51335/organoid.2024.4.e12 · Cited by 4

Advances in Human Brain Organoids: Methodological Innovations and Future Directions for Drug Discovery

Kanupriya, K., Verma, S., Sharma, V., Mishra, I., & Mishra, R. (2025) · Current Drug Research Reviews · doi:10.2174/0125899775369286250206050006 · Cited by 5

Self-organization and applications of neural organoids

Sakaguchi, H. (2025) · European Journal of Cell Biology · doi:10.1016/j.ejcb.2025.151496 · Cited by 5

Silicate Microfiber Scaffolds Support the Formation and Expansion of the Cortical Neuronal Layer of Cerebral Organoids With a Sheet-Like Configuration

Terada, E., Bamba, Y., Takagaki, M., Kawabata, S., Tachi, T., Nakamura, H., Nishida, T., & Kishima, H. (2023) · Stem Cells Translational Medicine · doi:10.1093/stcltm/szad066 · Cited by 5

Recapitulation and investigation of human brain development with neural organoids

Tamada, A., & Muguruma, K. (2023) · IBRO Neuroscience Reports · doi:10.1016/j.ibneur.2023.12.006 · Cited by 7

Patient-derived induced pluripotent stem cells: Tools to advance the understanding and drug discovery in Major Depressive Disorder

Vaz, A., Salgado, A. J., Patrício, P., & Pinto, L. (2024) · Psychiatry Research · doi:10.1016/j.psychres.2024.116033 · Cited by 4

Role of cell metabolism in the pathophysiology of brain size-associated neurodevelopmental disorders

Xing, L., Huttner, W. Β., & Namba, T. (2024) · Neurobiology of Disease · doi:10.1016/j.nbd.2024.106607 · Cited by 4

Modeling Alzheimer's disease using cerebral organoids: Current challenges and prospects

Abijo, A. Z., Olatunji, S., Adelodun, S. T., Asamu, M. O., & Omeiza, N. A. (2024) · Brain Organoid and Systems Neuroscience Journal · doi:10.1016/j.bosn.2024.09.001 · Cited by 6

Cell type specification and diversity in subpallial organoids

Pavon, N., Sun, Y., & Pak, C. (2024) · Frontiers in Genetics · doi:10.3389/fgene.2024.1440583 · Cited by 4

Minibrain Storm : Cerebral Organoids Aren't Real Brains?But They Provide a Powerful Platform for Modeling Brain Diseases Like Zika Infection, Alzheimer's, and Even Autism

Fischer, S. (2017) · IEEE Pulse · doi:10.1109/mpul.2017.2680360 · Cited by 4

Reprogramming the brain with synthetic neurobiology

Gardner, E. C., & Ellington, A. D. (2018) · Current Opinion in Biotechnology · doi:10.1016/j.copbio.2018.10.013 · Cited by 4

Three-dimensional culture systems in central nervous system research

Onuwaje, I., & Phillips, J. B. (2020) · Elsevier eBooks · doi:10.1016/b978-0-12-818084-6.00016-7 · Cited by 4

From neural stem cells to glioblastoma: A natural history of GBM recapitulated in vitro

Almengló, C., Caamaño, P., Fraga, M., Devesa, J., Costoya, J. A., & Arce, V. M. (2021) · Journal of Cellular Physiology · doi:10.1002/jcp.30409 · Cited by 5

IPSC-derived models in Africa: An HIV perspective

Naidoo, J., Hurrell, T., & Scholefield, J. (2022) · Biochimie · doi:10.1016/j.biochi.2022.01.013 · Cited by 4

Human brain organoid transplantation: ethical implications of enhancing specific cerebral functions in small-animal models

Harary, P. M., Blue, R., Castellanos, M., Dedhia, M., Hamimi, S. A., Jgamadze, D., Rees, B. W., Seshadri, N., Singh, S., Moreno, J. D., et al. (2023) · Molecular Psychology Brain Behavior and Society · doi:10.12688/molpsychol.17544.1 · Cited by 4

Using Human Induced Neural Precursor Cells to Define Early Neurodevelopmental Defects in Syndromic and Idiopathic Autism

Connacher, R., DiCicco-Bloom, E. M., & Millonig, J. H. (2018) · Current Pharmacology Reports · doi:10.1007/s40495-018-0155-0 · Cited by 4

Highly efficient induction of functionally mature excitatory neurons from feeder-free human ES/iPS cellsPreprint

Zhou, Z., Kakegawa, W., Fujimori, K., Sho, M., Shimamura, R., Supakul, S., Yoshimatsu, S., Kohyama, J., Yuzaki, M., & Okano, H. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.02.10.528087 · Cited by 4

Morphogen-guided neocortical organoids recapitulate regional areal identity and model neurodevelopmental disorder pathologyPreprint

Tsai, Y., Ozaki, H., Wang, X., Almet, A. A., Fleming, I., Shiraiwa, K., Noh, M. J. M., Nie, C., Trejo, S., Sugita, B. K., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.09.02.672952 · Cited by 4

Functional loss of ERBB receptor feedback inhibitor 1 ( MIG6 ) promotes glioblastoma tumorigenesis by aberrant activation of epidermal growth factor receptor ( EGFR )

Yi, S. A., Cho, D., Kim, S.-J., Kim, H., Choi, M. K., Choi, H. S., Shin, S., Yun, S., Lim, A., Jeong, J. K., et al. (2024) · Molecular Oncology · doi:10.1002/1878-0261.13717 · Cited by 3

Application of brain organoids in neurodevelopmental disorders

Zhao, C., Qi, Y., Xia, L., Zhu, T., & Xia, K. (2025) · Biochemical and Biophysical Research Communications · doi:10.1016/j.bbrc.2025.152405 · Cited by 3

Engineering human neuronal diversity: Morphogens and stem cell technologies for neurodevelopmental biology

Scuderi, S., Khouri‐Farah, N., Rauthan, R., Natu, A., Wang, H., Nelson, A., Jourdon, A., & Vaccarino, F. M. (2025) · Stem Cell Reports · doi:10.1016/j.stemcr.2025.102615 · Cited by 4

Single rosette-based generation of uniform cortical assembloids recapitulating cellular interactions between neurons and glial cells

Kim, E. H., Kim, Y., Hong, S.-J., Kim, I. S., Lee, J.-H., Yoo, J.-Y., Kim, J., Yoo, K., Lee, H., Kim, J., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-66440-1 · Cited by 8

Variability vs. phenotype: Multimodal analysis of Dravet syndrome brain organoids powered by deep learning

Turpín, I., Modrego, A., Martí-Sarrias, A., García‐González, L., Ortega-Gascó, A., Haeb, A.-C., Pareja, R. I., Soriano, J., Ruíz, N., Peñuelas‐Haro, I., et al. (2025) · iScience · doi:10.1016/j.isci.2025.113831 · Cited by 6

Pluripotent stem cells-based neural organoids for modelling human brain development and diseases

Tong, L., Tian, P., Wang, R., Niu, S., Wang, R., Ye, Y., Wu, Y., Zhang, W., Wang, Y., Foggetti, A., et al. (2025) · Cell & Bioscience · doi:10.1186/s13578-025-01515-6 · Cited by 3

A modular platform for automated organoid culture and longitudinal imaging

Torres, S., Hernandez, S., Seiler, S. T., Schweiger, H., Vera-Choqqueccota, S., Kaurala, G. A., Sharf, T., Haussler, D. H., Mostajo-Radji, M. A., & Teodorescu, M. (2026) · Scientific Reports · doi:10.1038/s41598-026-40231-0 · Cited by 6

Advances in cellular and structural engineering of brain organoids for disease modeling: A Comprehensive Review

Kim, H., Vasquez, F. J. A., Bello, A. B., & Lee, S.-H. (2026) · Tissue and Cell · doi:10.1016/j.tice.2026.103401 · Cited by 3

The sodium channel SCN2A regulates cortical excitatory and inhibitory neurogenesis

Uy, J. A., Dargaei, Z., Geahchan, S., Botler, L., Pan, Y., Brown, C. O., Howe, J., Scherer, S. W., Bains, J. S., & Singh, K. K. (2026) · Nature Communications · doi:10.1038/s41467-026-75751-w · Cited by 2

Brain organoids and genome editing: A new era in understanding human brain development and disorders

Zhou, M., Cao, Y., Yue, K., Wu, W., Xie, Y., Hu, D., Zhao, J., Xu, F., Guo, J., Li, Z., et al. (2025) · Neural Regeneration Research · doi:10.4103/nrr.nrr-d-24-01546 · Cited by 3

Mini-Brains Make Big Leaps for Studying Human Neural Development and Disease

Du, X., & Parent, J. M. (2016) · Epiliepsy currents/Epilepsy currents · doi:10.5698/1535-7511-16.6.402 · Cited by 2

Modeling neurodegenerative diseases by human pluripotent stem cell-induced brain organoid

Luo, X., Sun, R., Tian, F., Hao, J., & Hu, L. (2018) · Translational Surgery · doi:10.4103/ts.ts_4_18 · Cited by 3

Bioengineering of brain organoids: Advancements and challenges

Gopurappilly, R., & Pal, R. (2022) · Elsevier eBooks · doi:10.1016/b978-0-12-824064-9.00002-2 · Cited by 3

Brain organoids

Glass, M. R., Kyere, F. A., French, D. L., Stein, J. L., & Waxman, E. A. (2022) · Elsevier eBooks · doi:10.1016/b978-0-12-822277-5.00005-5 · Cited by 4

Shedding light on latent pathogenesis and pathophysiology of mental disorders: The potential of iPS cell technology

Arioka, Y., Okumura, H., Sakaguchi, H., & Ozaki, N. (2023) · Psychiatry and Clinical Neurosciences · doi:10.1111/pcn.13545 · Cited by 3

Publisher's note

Li, K., Gu, L., Cai, H., Lu, H., Mackie, K., & Guo, F. (2024) · Drug Metabolism and Pharmacokinetics · doi:10.1016/j.dmpk.2024.101031 · Cited by 3

Systematic time-dependent visualization and quantitation of the neurogenic rate in brain organoids

Kosodo, Y., Suetsugu, T., Kobayashi, T., & Matsuzaki, F. (2017) · Biochemical and Biophysical Research Communications · doi:10.1016/j.bbrc.2016.12.187 · Cited by 4

hiPSC-Based Tissue Organoid Regeneration

Qu, Y., Yucer, N., Garcia, V. J., Giuliano, A. E., & Cui, X. (2018) · InTech eBooks · doi:10.5772/intechopen.76997 · Cited by 3

A Guide to Understanding “State-of-the-Art” Basic Research Techniques in Anesthesiology

Obal, D., Wu, S., McKinstry-Wu, A. R., & Tawfik, V. L. (2020) · Anesthesia & Analgesia · doi:10.1213/ane.0000000000004801 · Cited by 3

A new technique for modeling neuronal connectivity using human pluripotent stem cells

Lee, C., Bendriem, R. M., & Freed, W. J. (2015) · Restorative Neurology and Neuroscience · doi:10.3233/rnn-140488 · Cited by 3

Development of Brain Organoids with Genome-Edited iPSC-Derived Brain Cells

Zagha, N., & Winner, B. (2022) · Advances in neuroethics · doi:10.1007/978-3-030-97641-5_2 · Cited by 2

Neural induction of embryonic stem/induced pluripotent stem cells

Sarkar, A., Marchetto, M. C., & Gage, F. H. (2020) · Elsevier eBooks · doi:10.1016/b978-0-12-814405-3.00009-6 · Cited by 3

Single Cell transcriptional analysis of ex vivo models of cortical and hippocampal development identifies unique longitudinal trendsPreprint

Krizay, D. K., Goldstein, D. B., & Boland, M. J. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.12.11.519959 · Cited by 2

Brain organoid-on-a-chip to create multiple domains in forebrain organoidsPreprint

Tsai, Y., Ozaki, H., Morikawa, A., Shiraiwa, K., Pin, A. P., Salem, A. G., Phommahasay, K. A., Sugita, B. K., Vu, C. H., Hammad, S. M., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.09.18.558278 · Cited by 3

Spatiotemporal expression of thyroid hormone transporter MCT8 and THRA mRNA in human cerebral organoids recapitulating first trimester cortex developmentPreprint

Graffunder, A. S., Bresser, A. A. J., Vallone, V. B. F., Megges, M., Stachelscheid, H., Kühnen, P., & Opitz, R. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.11.16.567497 · Cited by 3

Two parallel lineage-committed progenitors contribute to the developing brainPreprint

Dundes, C. E., Jokhai, R. T., Ahsan, H. S., Kang, R. S., Salomon, R. E. A., Rajan, A., Kim, Y. S., Stanton, L. J., Xu, C. L., Do, S., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.07.02.662771 · Cited by 3

Unified Generation of Regionalized Neural Organoids from Single-Lumen NeuroepitheliumPreprint

Rao, J., He, Z., Loskarn, S., Bender, A., Jo, Y., Lückel, J., Curcio, M., Cubela, I., Camp, J. G., Treutlein, B., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.11.18.689013 · Cited by 4

Drug Discovery in Induced Pluripotent Stem Cell Models

Bourque, K., Mnasri, N., Jones-Tabah, J., Martins-Cannavino, K., & Hébert, T. E. (2021) · Elsevier eBooks · doi:10.1016/b978-0-12-820472-6.00049-9 · Cited by 1

Neocortical neurogenesis: a proneural gene perspective

Vasan, L., Moffat, A., Mattar, P. A., & Schuurmans, C. (2025) · FEBS Journal · doi:10.1111/febs.70158 · Cited by 3

Brain organoids as models of extracellular vesicle–mediated human neural communication

Rosa, G. L., Pascale, E., Iazzetta, M. R., Sozzi, E., Fico, A., Parrotta, E. I., & Fiorenzano, A. (2026) · Neural Regeneration Research · doi:10.4103/nrr.nrr-d-25-01806 · Cited by 2

Contribution of tissue clearing and 3D image analysis to in vitro modeling of human cortical development

Retho, A., Govindan, A. D., Bonnet, M., Brot, S., & Gaillard, A. (2026) · Scientific Reports · doi:10.1038/s41598-026-41741-7 · Cited by 2

Protocol for quality control screening of brain organoid morphology

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