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Article History

Received: 18 October 2024

Accepted: 4 March 2025

First Online: 28 April 2025

Acknowledgements

: We thank B. Peers for the insulin:gfp zebrafish (University of Liège, Liège, Belgium), the National Institute of Genetics (NIG) scientist for pT2-L200R150-MCS1, S. J. Du for the pT2EF1a xbp1-gfp plasmid (University of Maryland School of Medicine, Baltimore, MD, USA) and T. van Ham for the 3.5ubb:secAnnexinV-mVenus plasmid (University Medical Center Rotterdam, Rotterdam, the Netherlands). We also thank E. Carroll (Delft University of Technology, Delft, the Netherlands) and all laboratory members for input, the Central Animal Facility (CDP) for facilitating and supporting experimental animal research, and the UMCG Microscopy and Imaging Center (UMIC) for supporting access to advanced microscopes and image processing.

: Original light microscopy images and raw electron microscopy data are open access via the BioImage Archive (S-BIAD1479, DOI: ). All raw EM data in zoomable format and the ESM Videos are available via .

: Part of the work was performed at the UMCG Microscopy and Imaging Center (UMIC), sponsored by ZonMW grant 91111.006, the Netherlands Electron Microscopy Infrastructure (NEMI), NWO National Roadmap for Large-Scale Research Infrastructure of the Dutch Research Council (NWO 184.034.014), the Network for Pancreatic Organ donors with Diabetes (nPOD; RRID:SCR_014641), a collaborative type 1 diabetes research project sponsored by Breakthrough T1D (nPOD: 5-SRA-2018-557-Q-R), and The Leona M. & Harry B. Helmsley Charitable Trust (grant no. 2018PG-T1D053). The content and views expressed are the responsibility of the authors and do not necessarily reflect the official view of nPOD. Organ Procurement Organizations (OPO) partnering with nPOD to provide research resources are listed at jdrfnpod.org/for-partners/npod-partners. BNGG and AZ are supported by ZonMW OPEN: 09120012010107; BHPD, BNGG NWO-ENPPS.LIFT.019.030. NF is supported by De Cock-Hadders Foundation and the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No. 754425.

: The authors declare that there are no relationships or activities that might bias, or be perceived to bias, their work.

: The authorship is based on the ICMJE uniform requirements. NF, WMHH, BHPD and AHGW generated plasmids and zebrafish lines. All authors conceived and designed the experiments. NF, WMHH, BHPD, AHGW and KK performed the experiments and acquired and analysed the data. NF wrote the manuscript. All authors contributed to interpretation of data, performed critical review of the manuscript and approved of the final version to be published. BNGG supervised this project. BNGG is the guarantor of this work.