{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,27]],"date-time":"2026-10-27T22:50:35Z","timestamp":1793141435627,"version":"4.5.0"},"reference-count":14,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,8,30]],"date-time":"2021-08-30T00:00:00Z","timestamp":1630281600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,8,30]],"date-time":"2021-08-30T00:00:00Z","timestamp":1630281600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["Natural Sciences and Engineering Research Council of Canada"],"award-info":[{"award-number":["Natural Sciences and Engineering Research Council of Canada"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100010785","name":"Canada First Research Excellence Fund","doi-asserted-by":"publisher","award":["Canada First Research Excellence Fund"],"award-info":[{"award-number":["Canada First Research Excellence Fund"]}],"id":[{"id":"10.13039\/501100010785","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002784","name":"Canada Excellence Research Chairs, Government of Canada","doi-asserted-by":"publisher","award":["Canada Excellence Research Chairs, Government of Canada"],"award-info":[{"award-number":["Canada Excellence Research Chairs, Government of Canada"]}],"id":[{"id":"10.13039\/501100002784","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000893","name":"Simons Foundation","doi-asserted-by":"publisher","award":["Simons Foundation"],"award-info":[{"award-number":["Simons Foundation"]}],"id":[{"id":"10.13039\/100000893","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Can-GARD","award":["Can-GARD"],"award-info":[{"award-number":["Can-GARD"]}]},{"DOI":"10.13039\/501100008453","name":"Ontario Institute for Regenerative Medicine","doi-asserted-by":"publisher","award":["Ontario Institute for Regenerative Medicine"],"award-info":[{"award-number":["Ontario Institute for Regenerative Medicine"]}],"id":[{"id":"10.13039\/501100008453","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Sharon Francis Institute for Regenerative Medicine","award":["Sharon Francis Institute for Regenerative Medicine"],"award-info":[{"award-number":["Sharon Francis Institute for Regenerative Medicine"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Brain"],"published-print":{"date-parts":[[2021,12]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The phosphatase and tensin homolog (PTEN) protein, encoded by the <jats:italic>PTEN<\/jats:italic> gene on chromosome 10, is a negative regulator of the phosphoinositide 3-kinase (PI3K) signaling pathway. Loss of <jats:italic>PTEN<\/jats:italic> has been linked to an array of human diseases, including neurodevelopmental disorders such as macrocephaly and autism. However, it remains unknown whether increased dosage of <jats:italic>PTEN<\/jats:italic> can lead to human disease. A recent human genetics study identifies chromosome 10 microduplication encompassing <jats:italic>PTEN<\/jats:italic> in patients with microcephaly. Here we generated a human brain organoid model of increased <jats:italic>PTEN<\/jats:italic> dosage. We showed that mild <jats:italic>PTEN<\/jats:italic> overexpression led to reduced neural precursor proliferation, premature neuronal differentiation, and the formation of significantly smaller brain organoids. <jats:italic>PTEN<\/jats:italic> overexpression resulted in decreased AKT activation, and treatment of wild-type organoids with an AKT inhibitor recapitulated the reduced brain organoid growth phenotypes. Together, our findings provide functional evidence that <jats:italic>PTEN<\/jats:italic> is a dosage-sensitive gene that regulates human neurodevelopment, and that increased <jats:italic>PTEN<\/jats:italic> dosage in brain organoids results in microcephaly-like phenotypes.<\/jats:p>","DOI":"10.1186\/s13041-021-00841-3","type":"journal-article","created":{"date-parts":[[2021,8,30]],"date-time":"2021-08-30T08:16:54Z","timestamp":1630311414000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":38,"title":["Modeling PTEN overexpression-induced microcephaly in human brain organoids"],"prefix":"10.1186","volume":"14","author":[{"given":"Navroop","family":"Dhaliwal","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wendy W.Y.","family":"Choi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julien","family":"Muffat","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5849-5910","authenticated-orcid":false,"given":"Yun","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,8,30]]},"reference":[{"key":"841_CR1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1146\/annurev-neuro-080317-061747","volume":"41","author":"KD Winden","year":"2018","unstructured":"Winden KD, Ebrahimi-Fakhari D, Sahin M. Abnormal mTOR Activation in Autism. Annu Rev Neurosci. 2018;41:1\u201323.","journal-title":"Annu Rev Neurosci"},{"issue":"8","key":"841_CR2","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1136\/jmedgenet-2018-105471","volume":"56","author":"D Oliveira","year":"2019","unstructured":"Oliveira D, et al. 10q23.31 microduplication encompassing PTEN decreases mTOR signalling activity and is associated with autosomal dominant primary microcephaly. J Med Genet. 2019;56(8):543\u20137.","journal-title":"J Med Genet"},{"issue":"2","key":"841_CR3","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1016\/j.celrep.2018.09.032","volume":"25","author":"A Omer Javed","year":"2018","unstructured":"Omer Javed A, et al. Microcephaly modeling of kinetochore mutation reveals a brain-specific phenotype. Cell Rep. 2018;25(2):368\u201382. e365.","journal-title":"Cell Rep"},{"issue":"7467","key":"841_CR4","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1038\/nature12517","volume":"501","author":"MA, Lancaster","year":"2013","unstructured":"Lancaster MA,, et al. Cerebral organoids model human brain development and microcephaly. Nature. 2013;501(7467):373\u20139.","journal-title":"Nature"},{"issue":"11","key":"841_CR5","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1007\/s13238-017-0479-2","volume":"8","author":"R Li","year":"2017","unstructured":"Li R, et al. Recapitulating cortical development with organoid culture in vitro and modeling abnormal spindle-like (ASPM related primary) microcephaly disease. Protein Cell. 2017;8(11):823\u201333.","journal-title":"Protein Cell"},{"issue":"6519","key":"841_CR6","doi-asserted-by":"publisher","first-page":"935","DOI":"10.1126\/science.abb5390","volume":"370","author":"C Esk","year":"2020","unstructured":"Esk C, et al. A human tissue screen identifies a regulator of ER secretion as a brain-size determinant. Science. 2020;370(6519):935\u201341.","journal-title":"Science"},{"issue":"27","key":"841_CR7","doi-asserted-by":"publisher","first-page":"7117","DOI":"10.1073\/pnas.1719266115","volume":"115","author":"J Muffat","year":"2018","unstructured":"Muffat J, et al. Human induced pluripotent stem cell-derived glial cells and neural progenitors display divergent responses to Zika and dengue infections. Proc Natl Acad Sci U S A. 2018;115(27):7117\u201322.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5","key":"841_CR8","doi-asserted-by":"publisher","first-page":"1238","DOI":"10.1016\/j.cell.2016.04.032","volume":"165","author":"X Qian","year":"2016","unstructured":"Qian X, et al. Brain-region-specific organoids using mini-bioreactors for modeling ZIKV exposure. Cell. 2016;165(5):1238\u201354.","journal-title":"Cell"},{"issue":"7606","key":"841_CR9","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1038\/nature18296","volume":"534","author":"FR, Cugola","year":"2016","unstructured":"Cugola FR,, et al. The Brazilian Zika virus strain causes birth defects in experimental models. Nature. 2016;534(7606):267\u201371.","journal-title":"Nature"},{"issue":"3","key":"841_CR10","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1016\/j.stem.2016.11.017","volume":"20","author":"Y Li","year":"2017","unstructured":"Li Y, et al. Induction of expansion and folding in human cerebral organoids. Cell stem cell. 2017;20(3):385\u201396 e383.","journal-title":"Cell stem cell"},{"issue":"2","key":"841_CR11","doi-asserted-by":"publisher","first-page":"292","DOI":"10.1086\/519999","volume":"81","author":"E Boland","year":"2007","unstructured":"Boland E, et al. Mapping of deletion and translocation breakpoints in 1q44 implicates the serine\/threonine kinase AKT3 in postnatal microcephaly and agenesis of the corpus callosum. Am J Hum Genet. 2007;81(2):292\u2013303.","journal-title":"Am J Hum Genet"},{"issue":"8","key":"841_CR12","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1038\/ng.2329","volume":"44","author":"JH, Lee","year":"2012","unstructured":"Lee JH,, et al. De novo somatic mutations in components of the PI3K-AKT3-mTOR pathway cause hemimegalencephaly. Nat Genet. 2012;44(8):941\u20135.","journal-title":"Nat Genet"},{"issue":"8","key":"841_CR13","doi-asserted-by":"publisher","first-page":"934","DOI":"10.1038\/ng.2331","volume":"44","author":"JB, Riviere","year":"2012","unstructured":"Riviere JB,, et al. De novo germline and postzygotic mutations in AKT3, PIK3R2 and PIK3CA cause a spectrum of related megalencephaly syndromes. Nat Genet. 2012;44(8):934\u201340.","journal-title":"Nat Genet"},{"issue":"1","key":"841_CR14","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.neuron.2012.03.010","volume":"74","author":"A Poduri","year":"2012","unstructured":"Poduri A, et al. Somatic activation of AKT3 causes hemispheric developmental brain malformations. Neuron. 2012;74(1):41\u20138.","journal-title":"Neuron"}],"container-title":["Molecular Brain"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13041-021-00841-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13041-021-00841-3\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13041-021-00841-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,8,30]],"date-time":"2021-08-30T08:25:48Z","timestamp":1630311948000},"score":1,"resource":{"primary":{"URL":"https:\/\/molecularbrain.biomedcentral.com\/articles\/10.1186\/s13041-021-00841-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,8,30]]},"references-count":14,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2021,12]]}},"alternative-id":["841"],"URL":"https:\/\/doi.org\/10.1186\/s13041-021-00841-3","relation":{},"ISSN":["1756-6606"],"issn-type":[{"value":"1756-6606","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,8,30]]},"assertion":[{"value":"3 June 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"19 August 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 August 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The WIBR3 human embryonic stem cell line was approved for use by the\nStem Cell Oversight Committee of the Canadian Institutes of Health Research,\nand the Research Ethics Board of the Hospital for Sick Children.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"131"}}