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We explored the use of extracellular electron transfer for energy conservation in diverse lactic acid bacteria (LAB), microorganisms that mainly rely on fermentative metabolism and are important in food fermentations. The LAB\n                    <jats:italic>Lactiplantibacillus plantarum<\/jats:italic>\n                    uses extracellular electron transfer to increase its NAD\n                    <jats:sup>+<\/jats:sup>\n                    \/NADH ratio, generate more ATP through substrate-level phosphorylation, and accumulate biomass more rapidly. This novel, hybrid metabolism is dependent on a type-II NADH dehydrogenase (Ndh2) and conditionally requires a flavin-binding extracellular lipoprotein (PplA) under laboratory conditions. It confers increased fermentation product yield, metabolic flux, and environmental acidification in laboratory media and during kale juice fermentation. The discovery of a single pathway that simultaneously blends features of fermentation and respiration in a primarily fermentative microorganism expands our knowledge of energy conservation and provides immediate biotechnology applications.\n                  <\/jats:p>","DOI":"10.7554\/elife.70684","type":"journal-article","created":{"date-parts":[[2021,12,29]],"date-time":"2021-12-29T08:16:02Z","timestamp":1640765762000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.70684","source":"Crossref","is-referenced-by-count":116,"title":["Extracellular electron transfer increases fermentation in lactic acid bacteria via a hybrid metabolism"],"prefix":"10.7554","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1192-8256","authenticated-orcid":true,"given":"Sara","family":"Tejedor-Sanz","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/008zs3103","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of BioSciences, Rice University","place":["Houston, United States"]},{"id":[{"id":"https:\/\/ror.org\/02jbv0t02","id-type":"ROR","asserted-by":"publisher"}],"name":"Biological Nanostructures Facility, The Molecular Foundry, Lawrence Berkeley National Laboratory","place":["Berkeley, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0648-8281","authenticated-orcid":true,"given":"Eric T","family":"Stevens","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05rrcem69","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Food Science & Technology, University of California\u2010Davis","place":["Davis, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Siliang","family":"Li","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/008zs3103","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of BioSciences, Rice University","place":["Houston, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter","family":"Finnegan","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05rrcem69","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Food Science & Technology, University of California\u2010Davis","place":["Davis, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James","family":"Nelson","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05rrcem69","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Electrical and Computer Engineering, University of California\u2010Davis","place":["Davis, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andre","family":"Knoesen","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05rrcem69","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Electrical and Computer Engineering, University of California\u2010Davis","place":["Davis, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8074-1348","authenticated-orcid":true,"given":"Samuel H","family":"Light","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Microbiology, University of Chicago","place":["Chicago, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8909-6712","authenticated-orcid":true,"given":"Caroline M","family":"Ajo-Franklin","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/008zs3103","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of BioSciences, Rice University","place":["Houston, United States"]},{"id":[{"id":"https:\/\/ror.org\/02jbv0t02","id-type":"ROR","asserted-by":"publisher"}],"name":"Biological Nanostructures Facility, The Molecular Foundry, Lawrence Berkeley National Laboratory","place":["Berkeley, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3643-9766","authenticated-orcid":true,"given":"Maria L","family":"Marco","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05rrcem69","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Food Science & Technology, University of California\u2010Davis","place":["Davis, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4374","published-online":{"date-parts":[[2022,2,11]]},"reference":[{"key":"bib1","article-title":"FastQC: a quality control tool for high throughput sequence data","volume-title":"Babraham Bioinformatics","author":"Andrews","year":"2010","unstructured":"Andrews S. 2010. 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