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Article History

Received: 13 March 2025

Accepted: 3 July 2025

First Online: 24 September 2025

Acknowledgements

: We are grateful to T. Davey and R. Laws at Electron Microscopy Research Services, Newcastle University, UK, for their assistance with tissue processing for transmission electron microscopy. We also wish to acknowledge the support of M. Palmer, University of Exeter, UK, for his support developing the SRFG analysis pipeline.

: The datasets generated during and/or analysed during the current study are available from the corresponding author upon reasonable request.

: The study funder was not involved in the design of the study; the collection, analysis, and interpretation of data; writing the report; and did not impose any restrictions regarding the publication of the report. This work was supported by a Steve Morgan Foundation Grand Challenge Senior Research Fellowship awarded to SJR (22/0006504), and a Multimodal Multiscale Initiative awarded by the Medical Research Council (MR/W031515/1). This research is supported by the National Institute for Health and Care Research (NIHR) Exeter Biomedical Research Centre (BRC). The views expressed are those of the authors and not necessarily those of the NIHR or the Department of Health and Social Care.

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

: NJD processed human donor pancreas tissue, performed EM tissue processing, sectioning and imaging, analysed the data and wrote the manuscript first draft. NK and YA-S analysed the data, and reviewed and edited the manuscript. MH-S, MFS and CAB processed human donor pancreas tissue and reviewed and edited the manuscript. RC performed FFPE tissue embedding, sectioning and staining and reviewed and edited the manuscript. CSF and SJR carried out HALO data generation/analysis and reviewed/edited the manuscript. JAMS acquired funding, advised on data analysis and wrote, reviewed and edited the manuscript. All authors approved the final version of the manuscript. JAMS is the guarantor of this work and, as such, had full access to all the data in the study and takes responsibility for the integrity of the data and the accuracy of the data analysis.