{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T17:38:09Z","timestamp":1792604289350,"version":"4.5.0"},"reference-count":50,"publisher":"National Academy of Sciences","issue":"43","license":[{"start":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T00:00:00Z","timestamp":1792540800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100020302","name":"UC | UCSD | Materials Research Science and Engineering Center, University of California, San Diego","doi-asserted-by":"publisher","award":["DMR-2614051"],"award-info":[{"award-number":["DMR-2614051"]}],"id":[{"id":"10.13039\/100020302","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100006221","name":"United States - Israel Binational Science Foundation","doi-asserted-by":"publisher","award":["BSF 2023269"],"award-info":[{"award-number":["BSF 2023269"]}],"id":[{"id":"10.13039\/100006221","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,10,27]]},"abstract":"<jats:p>\n                    Cyanobacteria provide a promising platform for light-driven heterologous protein production. In heterologous production systems, cells are typically lysed for product purification or to facilitate interaction with external substrates, unless the product is secreted. Fusion to molecular \u201cshipping labels\u201d known as signal peptides can target chimeric proteins to native membrane translocation pathways and result in secretion. In this work, we identified signal peptides from the model cyanobacterium\n                    <jats:italic toggle=\"yes\">Synechococcus elongatus<\/jats:italic>\n                    PCC 7942 (\n                    <jats:italic toggle=\"yes\">S. elongatus<\/jats:italic>\n                    ) and genetically fused to two heterologous enzymes: the reporter nanoluciferase (NLuc) and an oxidative laccase (CotA). Increased extracellular activity was observed for one or more signal-protein variants for both enzymes. Furthermore, cells of\n                    <jats:italic toggle=\"yes\">S. elongatus<\/jats:italic>\n                    producing a signal-laccase chimera were encapsulated within an alginate hydrogel, and the resulting engineered living material released sufficient enzyme to decolorize a dye pollutant in its surroundings. These findings streamline recombinant protein production in cyanobacteria and facilitate environmental interactions with cell products, expanding the potential applications of both free-living and material-encapsulated cyanobacteria.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2624792123","type":"journal-article","created":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T16:44:30Z","timestamp":1792601070000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["Signal peptides for engineered protein secretion by free and hydrogel-encapsulated cyanobacteria"],"prefix":"10.1073","volume":"123","author":[{"given":"Nathan T.","family":"Soulier","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Biology, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Materials Science and Engineering Program, University of California San Diego","place":["La Jolla"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7146-7095","authenticated-orcid":false,"given":"Debika","family":"Datta","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Materials Science and Engineering Program, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Aiiso Yufeng Li Family Department of Chemical and Nano Engineering, University of California San Diego","place":["La Jolla"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0521-1881","authenticated-orcid":false,"given":"Rakefet","family":"Schwarz","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03kgsv495","id-type":"ROR","asserted-by":"publisher"}],"name":"The Mina and Everard Goodman Faculty of Life Sciences, Bar-Ilan University","place":["Ramat-Gan, Israel"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5869-6942","authenticated-orcid":false,"given":"Jonathan K.","family":"Pokorski","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Materials Science and Engineering Program, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Aiiso Yufeng Li Family Department of Chemical and Nano Engineering, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Institute for Materials Discovery and Design, University of California San Diego","place":["La Jolla"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5463-3207","authenticated-orcid":false,"given":"James W.","family":"Golden","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Biology, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Materials Science and Engineering Program, University of California San Diego","place":["La Jolla"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4264-7019","authenticated-orcid":false,"given":"Susan S.","family":"Golden","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Biology, University of California San Diego","place":["La Jolla"]},{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Materials Science and Engineering Program, University of California San Diego","place":["La Jolla"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2026,10,21]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12934-018-0901-3"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1186\/s13568-017-0394-1"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.00504-21"},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1128\/AEM.00671-19"},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12934-022-01917-y"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1186\/s13068-019-1416-9"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.3390\/environments7020013"},{"key":"e_1_3_4_8_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00371-17"},{"key":"e_1_3_4_9_2","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aaz4354"},{"key":"e_1_3_4_10_2","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-024-06910-2"},{"key":"e_1_3_4_11_2","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.03674-20"},{"key":"e_1_3_4_12_2","doi-asserted-by":"publisher","DOI":"10.1038\/srep23080"},{"key":"e_1_3_4_13_2","doi-asserted-by":"publisher","DOI":"10.1515\/hsz-2020-0247"},{"key":"e_1_3_4_14_2","doi-asserted-by":"publisher","DOI":"10.1128\/ecosalplus.esp-0019-2015"},{"key":"e_1_3_4_15_2","doi-asserted-by":"publisher","DOI":"10.1111\/1462-2920.12516"},{"key":"e_1_3_4_16_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.176.7.1857-1864.1994"},{"key":"e_1_3_4_17_2","doi-asserted-by":"publisher","DOI":"10.1002\/pro.757"},{"key":"e_1_3_4_18_2","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2022.933153"},{"key":"e_1_3_4_19_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbabio.2015.08.010"},{"key":"e_1_3_4_20_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2006.05365.x"},{"key":"e_1_3_4_21_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.tim.2022.03.009"},{"key":"e_1_3_4_22_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2958.2003.03546.x"},{"key":"e_1_3_4_23_2","doi-asserted-by":"publisher","DOI":"10.1111\/mmi.12407"},{"key":"e_1_3_4_24_2","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.02338-22"},{"key":"e_1_3_4_25_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0378-1097(02)01197-7"},{"key":"e_1_3_4_26_2","doi-asserted-by":"publisher","DOI":"10.1039\/c2ee21124f"},{"key":"e_1_3_4_27_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.envpol.2018.01.071"},{"key":"e_1_3_4_28_2","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.RA118.001929"},{"key":"e_1_3_4_29_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12934-020-01339-8"},{"key":"e_1_3_4_30_2","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiaf186"},{"key":"e_1_3_4_31_2","doi-asserted-by":"publisher","DOI":"10.1111\/mmi.14461"},{"key":"e_1_3_4_32_2","doi-asserted-by":"publisher","DOI":"10.1007\/s00253-025-13532-z"},{"key":"e_1_3_4_33_2","doi-asserted-by":"publisher","DOI":"10.1021\/acssynbio.3c00157"},{"key":"e_1_3_4_34_2","doi-asserted-by":"publisher","DOI":"10.1128\/AEM.01146-10"},{"key":"e_1_3_4_35_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2006.07.034"},{"key":"e_1_3_4_36_2","doi-asserted-by":"publisher","DOI":"10.1002\/adfm.202108057"},{"key":"e_1_3_4_37_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-47132-8"},{"key":"e_1_3_4_38_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41563-020-00857-5"},{"key":"e_1_3_4_39_2","doi-asserted-by":"publisher","DOI":"10.1007\/s42242-021-00129-4"},{"key":"e_1_3_4_40_2","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2021.772324"},{"key":"e_1_3_4_41_2","doi-asserted-by":"crossref","unstructured":"L. Tang A responsive living material prepared by diffusion reveals extracellular enzyme activity of cyanobacteria. Proc. Natl. Acad. Sci. U.S.A. 122 e2424405122 (2025).","DOI":"10.1073\/pnas.2424405122"},{"key":"e_1_3_4_42_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-40265-2"},{"key":"e_1_3_4_43_2","doi-asserted-by":"publisher","DOI":"10.3389\/fbioe.2018.00095"},{"key":"e_1_3_4_44_2","doi-asserted-by":"publisher","DOI":"10.2217\/fmb-2019-0010"},{"key":"e_1_3_4_45_2","doi-asserted-by":"publisher","DOI":"10.1021\/cb3002478"},{"key":"e_1_3_4_46_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12934-018-1041-5"},{"key":"e_1_3_4_47_2","doi-asserted-by":"publisher","DOI":"10.1111\/1758-2229.13050"},{"key":"e_1_3_4_48_2","doi-asserted-by":"publisher","DOI":"10.1002\/bit.21898"},{"key":"e_1_3_4_49_2","doi-asserted-by":"publisher","DOI":"10.1002\/cctc.202301055"},{"key":"e_1_3_4_50_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.emboj.7600606"}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2624792123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T16:45:07Z","timestamp":1792601107000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2624792123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,21]]},"references-count":50,"journal-issue":{"issue":"43","published-print":{"date-parts":[[2026,10,27]]}},"alternative-id":["10.1073\/pnas.2624792123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2624792123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,21]]},"assertion":[{"value":"2026-07-14","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-17","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-21","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2624792123"}}