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These strains were engineered by modulating intracellular levels of dinucleotide second messengers. We systematically investigated the capacity of biofilms formed by four strains to reduce different Fe(III)-containing minerals including ferrihydrite, goethite, and lepidocrocite. By growing the\n              <jats:italic toggle=\"yes\">G. sulfurreducens<\/jats:italic>\n              biofilm on the Fe(III) mineral-coated slides, our results showed that the strains forming thin biofilms on surfaces of Fe(III) minerals exhibited faster Fe(III) reduction rates compared to those with thick biofilms. Transcriptomic analyses revealed the upregulation of the genes encoding bacterial EET-involved proteins in the thin biofilms, highlighting the significant role of these proteins in reducing Fe(III)-containing minerals by\n              <jats:italic toggle=\"yes\">G. sulfurreducens<\/jats:italic>\n              biofilms. Furthermore, genetic characterization identified the participation of two novel\n              <jats:italic toggle=\"yes\">c<\/jats:italic>\n              -type cytochromes (\n              <jats:italic toggle=\"yes\">c<\/jats:italic>\n              -Cyts), GSU1996 and GSU2513, in the reduction of Fe(III)-containing minerals by\n              <jats:italic toggle=\"yes\">G. sulfurreducens<\/jats:italic>\n              biofilms. The results from this study provide an improved understanding of mineral\u2013microbe interaction.\n            <\/jats:p>\n            <jats:sec>\n              <jats:title>\n                <jats:bold>IMPORTANCE<\/jats:bold>\n              <\/jats:title>\n              <jats:p>\n                <jats:italic toggle=\"yes\">Geobacter<\/jats:italic>\n                is a predominant species within biofilm communities that facilitate iron reduction, a process essential for the biogeochemical cycling of iron and other elements. However, the specific properties of\n                <jats:italic toggle=\"yes\">Geobacter<\/jats:italic>\n                biofilms crucial for iron reduction remain unclear. By manipulating intracellular levels of dinucleotide second messengers to generate strains with varying biofilm properties, this research reveals that thinner biofilms exhibit superior rates of ferric iron [Fe(III)] mineral reduction compared to thicker biofilms. This finding highlights the vital role of proteins involved in extracellular electron transfer (EET) in enhancing the reduction of Fe(III)-containing minerals. The study further identifies two novel\n                <jats:italic toggle=\"yes\">c<\/jats:italic>\n                -type cytochromes, GSU1996 and GSU2513, as important contributors to this process. These discoveries not only advance our understanding of microbial iron reduction but also offer new perspectives on the interactions between biofilms and mineral surfaces, potentially informing future research and applications in biogeochemical cycling and bioenergy.\n              <\/jats:p>\n            <\/jats:sec>\n          <\/jats:sec>","DOI":"10.1128\/aem.00369-25","type":"journal-article","created":{"date-parts":[[2025,4,9]],"date-time":"2025-04-09T13:01:21Z","timestamp":1744203681000},"update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":15,"title":["Extracellular electron transfer proteins contribute to reduction of ferric minerals by\n            <i>Geobacter<\/i>\n            biofilms"],"prefix":"10.1128","volume":"91","author":[{"given":"Jiacheng","family":"Xu","sequence":"first","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Zhou","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xi","family":"Han","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jian","family":"Liu","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6155-2544","authenticated-orcid":true,"given":"Yiran","family":"Dong","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]},{"name":"State Key Laboratory of Geomicrobiology and Environmental Changes, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3829-7423","authenticated-orcid":true,"given":"Yongguang","family":"Jiang","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuhong","family":"Zhong","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4546-4741","authenticated-orcid":true,"given":"Liang","family":"Shi","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]},{"name":"State Key Laboratory of Geomicrobiology and Environmental Changes, China University of Geosciences","place":["Wuhan, China"]},{"name":"Hubei Key Laboratory of Yangtze Catchment Environmental Aquatic Science, China University of Geosciences","place":["Wuhan, China"]},{"name":"State Environmental Protection Key Laboratory of Source Apportionment and Control of Aquatic Pollution, Ministry of Ecology and Environment, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6440-5678","authenticated-orcid":true,"given":"Yidan","family":"Hu","sequence":"additional","affiliation":[{"name":"School of Environmental Studies, China University of Geosciences","place":["Wuhan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.2136\/sssabookser7"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0016-7037(03)00371-5"},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1021\/es1040249"},{"key":"e_1_3_4_5_2","first-page":"1","volume-title":"Microbial metal respiration: from geochemistry to potential applications","author":"Majzlan J","year":"2012","unstructured":"Majzlan J. 2012. 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