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Authors
Skak, Nikolaj https://orcid.org/0009-0009-8124-1124
Mertz, Nina https://orcid.org/0000-0003-3543-0433
Piwon, Nils https://orcid.org/0000-0002-1225-5794
Olsen, Martin R.
Caffarel-Salvador, Ester https://orcid.org/0000-0002-2844-7598
Brayden, David J. https://orcid.org/0000-0002-8781-8344
Østergaard, Jesper https://orcid.org/0000-0003-3448-4237
Lindhardt, Karsten https://orcid.org/0000-0002-5718-4950
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Funding
Funding for this research was provided by:
Innovationsfonden (1146-00004)
Innovationsfonden (1045-00053B)
Innovationsfonden (1045-00053B)
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Article History
Received: 21 November 2025
Accepted: 15 May 2026
First Online: 28 May 2026
Declarations
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: The housing of the animals was in accordance with EU Directive 2012/63/EU of 22 September 2012 on the protection of animals for scientific purposes and the animal experimentation was carried out under a license provided by the National Animal Experiments Inspectorate under the Ministry of Food, Agriculture and Fisheries of Denmark (license No: 2021-15-0201-00875). The studies involving ex vivo pig stomach tissue (Supplementary Information Fig. S and S ) were carried out under a permit for handling animal byproducts provided by the Ministry of Food, Agriculture and Fisheries of Denmark (license: DK-123-oth-1425482).
: KL and NS are co-inventors of multiple published Biograil patents (WO2019/121686; WO2023/073176; WO2022/162149; WO2023/073176; WO2023/144159) relevant for this paper. Biograil has other granted and pending patents.
: KL and NS are founders of Biograil Aps a pre-clinical stage pharmaceutical company developing the BIONDD ® technology for oral delivery of biologics. NP is chief development officer at Biograil ApS. NM and MRO are employed by Biograil ApS. ECS and DB are members of Biograil ApS’ Scientific Advisory Board. DB declares consultancy work on oral peptide administration with other companies. The authors declare no other competing interests.
