{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,13]],"date-time":"2026-09-13T21:17:46Z","timestamp":1789334266299,"version":"build-2803163510"},"reference-count":167,"publisher":"Oxford University Press (OUP)","issue":"8","license":[{"start":{"date-parts":[[2026,4,25]],"date-time":"2026-04-25T00:00:00Z","timestamp":1777075200000},"content-version":"vor","delay-in-days":2,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/100013060","name":"EMBL Heidelberg","doi-asserted-by":"publisher","award":["PID2021-123030NB-I00 (AR-I)"],"award-info":[{"award-number":["PID2021-123030NB-I00 (AR-I)"]}],"id":[{"id":"10.13039\/100013060","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100010663","name":"European Research Council","doi-asserted-by":"publisher","award":["ENHPATHY H2020-MSCA-ITN-2019\u2013860002 (AR-I)"],"award-info":[{"award-number":["ENHPATHY H2020-MSCA-ITN-2019\u2013860002 (AR-I)"]}],"id":[{"id":"10.13039\/100010663","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000780","name":"European Commission","doi-asserted-by":"publisher","award":["Chromrare HORIZON- MSCA-2021-DN-01\u2013101073334 (AR-I)"],"award-info":[{"award-number":["Chromrare HORIZON- MSCA-2021-DN-01\u2013101073334 (AR-I)"]}],"id":[{"id":"10.13039\/501100000780","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,4,23]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Defects in ZIC2, a member of the Zinc Finger of the Cerebellum family of transcription factors (TFs), cause holoprosencephaly, a congenital brain malformation characterized by the defective cleavage of cerebral hemispheres. However, the gene regulatory network (GRN) controlled by ZIC2 during neural development remains largely unexplored. Here, we combined a mouse embryonic stem cell (mESC) in vitro differentiation model toward anterior neural progenitors with genome editing and genomic methods to elucidate the ZIC2 GRN. We found that ZIC2 is dispensable in mESC due to compensation by ZIC3. In contrast, during neural induction ZIC2 directly controls the expression of master regulators implicated in the patterning and morphogenesis of specific brain regions (e.g. midbrain and roof plate). Mechanistically, ZIC2 plays a dual role in neural differentiation: during pluripotency exit, ZIC2 binds de novo to distal enhancers and increases their chromatin accessibility; during neural induction, ZIC2 is essential for the activation of a subset of the previously primed enhancers, which in turn control the expression of neural patterning regulators and signalling pathways (i.e. WNT) that prevent premature neuronal differentiation. Therefore, by sequentially acting as a promiscuous priming TF and selective enhancer activator, ZIC2 canalizes pluripotent cells toward neural progenitors with rostro-dorsal identities.<\/jats:p>","DOI":"10.1093\/nar\/gkag374","type":"journal-article","created":{"date-parts":[[2026,4,11]],"date-time":"2026-04-11T11:38:58Z","timestamp":1775907538000},"source":"Crossref","is-referenced-by-count":1,"title":["Dual role of ZIC2 during neural induction: from priming transcription factor to enhancer activator"],"prefix":"10.1093","volume":"54","author":[{"given":"Mar\u00eda","family":"Mariner-Faul\u00ed","sequence":"first","affiliation":[{"name":"Institute of Biomedicine and Biotechnology of Cantabria, CSIC\/Universidad de Cantabria , Santander, 39011 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"V\u00edctor","family":"S\u00e1nchez-Gaya","sequence":"additional","affiliation":[{"name":"Institute of Biomedicine and Biotechnology of Cantabria, CSIC\/Universidad de Cantabria , Santander, 39011 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah Malika","family":"Robert","sequence":"additional","affiliation":[{"name":"Institute of Biomedicine and Biotechnology of Cantabria, CSIC\/Universidad de Cantabria , Santander, 39011 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patricia","family":"Respuela","sequence":"additional","affiliation":[{"name":"Institute of Biomedicine and Biotechnology of Cantabria, CSIC\/Universidad de Cantabria , Santander, 39011 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara","family":"de la Cruz-Molina","sequence":"additional","affiliation":[{"name":"Center for Molecular Medicine Cologne, University of Cologne , Cologne, 50931 ,","place":["Germany"]},{"name":"Miltenyi Biotec B.V. & Co. KG , Bergisch Gladbach, 51429 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara","family":"Lobato-Moreno","sequence":"additional","affiliation":[{"name":"European Molecular Biology Laboratory, Molecular Systems Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"European Molecular Biology Laboratory, Genome Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"Faculty of Biosciences, Collaboration for Joint PhD Degree between EMBL and Heidelberg University , Heidelberg, 69117 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matteo","family":"Trovato","sequence":"additional","affiliation":[{"name":"European Molecular Biology Laboratory, Molecular Systems Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"Faculty of Biosciences, Collaboration for Joint PhD Degree between EMBL and Heidelberg University , Heidelberg, 69117 ,","place":["Germany"]},{"name":"IFOM ETS\u2014the AIRC Institute of Molecular Oncology , Milan, 20139 ,","place":["Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karin D","family":"Prummel","sequence":"additional","affiliation":[{"name":"European Molecular Biology Laboratory, Molecular Systems Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"European Molecular Biology Laboratory, Genome Biology Unit , Heidelberg, 69117 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kyung-Min","family":"Noh","sequence":"additional","affiliation":[{"name":"European Molecular Biology Laboratory, Genome Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"Department of Biomedicine, Aarhus University , Aarhus, 8000 ,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Judith B","family":"Zaugg","sequence":"additional","affiliation":[{"name":"European Molecular Biology Laboratory, Molecular Systems Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"European Molecular Biology Laboratory, Genome Biology Unit , Heidelberg, 69117 ,","place":["Germany"]},{"name":"Department of Biomedicine, University of Basel, Basel University Hospital , Basel, 4031 ,","place":["Switzerland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7137-1341","authenticated-orcid":false,"given":"\u00c1lvaro","family":"Rada-Iglesias","sequence":"additional","affiliation":[{"name":"Institute of Biomedicine and Biotechnology of Cantabria, CSIC\/Universidad de Cantabria , Santander, 39011 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,4,25]]},"reference":[{"key":"2026042505234186300_B1","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1016\/j.tig.2012.02.008","article-title":"Enhancers as information integration hubs in development: lessons from 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