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To date, there are no reports on engineered filamentous cyanobacteria for bioethanol or biohydrogen production. However, biofilm formation in filamentous cyanobacteria could provide an advantage for biotechnological applications. We used natural transformation and homologous recombination to establish recombinant expression at the\n                    <jats:italic>psaD<\/jats:italic>\n                    locus of the filamentous cyanobacterium\n                    <jats:italic>Phormidium lacuna<\/jats:italic>\n                    . A similar approach has already been successful in the unicellular cyanobacterium\n                    <jats:italic>Synechocystis<\/jats:italic>\n                    sp. PCC 6803 (Appel et al. 2020). Fusion of the NiFe hydrogenase subunits HoxY\/HoxH from\n                    <jats:italic>Microcystis aeruginosa<\/jats:italic>\n                    to the PSI subunit PsaD increased hydrogen production five-fold under light in the presence of DCMU, although dark fermentative hydrogen levels remained higher. Structural modeling indicates that electron transfer may be limited by the FeS cluster distance, suggesting opportunities for further optimization. Using sfGFP fluorescence, we showed that the\n                    <jats:italic>psaD<\/jats:italic>\n                    promoter drives strong expression when the gene is positioned 3\u2032 of the\n                    <jats:italic>psaD<\/jats:italic>\n                    gene or expressed as a PsaD\u2013sfGFP fusion protein. The\n                    <jats:italic>psaD<\/jats:italic>\n                    promoter was then used for expression of pyruvate decarboxylase and alcohol dehydrogenase, resulting in ethanol production of 0.2\u2030 (v\/v) and outperforming the kanamycin promoter used in previous unsuccessful trials. These results demonstrate that\n                    <jats:italic>P. lacuna<\/jats:italic>\n                    is a robust host for biohydrogen and bioethanol production and validate the\n                    <jats:italic>psaD<\/jats:italic>\n                    promoter as a strong regulatory element for filamentous cyanobacteria in biotechnological applications.\n                  <\/jats:p>","DOI":"10.1007\/s00203-026-04932-4","type":"journal-article","created":{"date-parts":[[2026,5,30]],"date-time":"2026-05-30T08:59:58Z","timestamp":1780131598000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Increased hydrogen and ethanol production in transformants of the filamentous cyanobacterium Phormidium lacuna"],"prefix":"10.1007","volume":"208","author":[{"given":"Nina","family":"Spohrer","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Florian","family":"Reinicke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefan","family":"Malik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Racha","family":"Tarakji","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nora","family":"Weber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4327-9737","authenticated-orcid":false,"given":"Tilman","family":"Lamparter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,30]]},"reference":[{"key":"4932_CR1","doi-asserted-by":"publisher","unstructured":"Abreu AP, Martins R, Nunes J (2023) Emerging Applications of Chlorella sp. and Spirulina (Arthrospira) sp. 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