{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T09:22:19Z","timestamp":1791969739123,"version":"4.3.4"},"reference-count":104,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T00:00:00Z","timestamp":1789516800000},"content-version":"vor","delay-in-days":258,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,14]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Siderophores are classically viewed as shared iron-scavenging public goods, yet their ecological roles in multispecies communities remain incompletely understood. Here, we establish a synthetic microbial community to dissect how different siderophores, their uptake compatibility and spatial structure shape iron competition. Using Corynebacterium glutamicum as a model, we show that this siderophore non-producer accesses diverse xenosiderophores, including enterobactin secreted by Escherichia coli. However, exploitation was constrained and co-cultures converged to stable compositions. Dose\u2013response experiments combined with mathematical modelling indicated that the producer retains more effective access to enterobactin than the exploiter. The presence of Pseudomonas putida altered this interaction, as it exploited enterobactin while producing pyoverdine, a siderophore inaccessible to the other community members that restricted their iron access. Across different cultivation scales, community dynamics was strongly influenced by spatial organization and initial composition. These findings identify siderophores as context-dependent iron-allocation agents that can promote microbial coexistence or exclusion.<\/jats:p>","DOI":"10.1093\/ismejo\/wrag247","type":"journal-article","created":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T14:34:41Z","timestamp":1789569281000},"source":"Crossref","is-referenced-by-count":1,"title":["Context-dependent siderophore exploitability shapes microbial community structure"],"prefix":"10.1093","volume":"20","author":[{"given":"Aileen","family":"Kr\u00fcger","sequence":"first","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]},{"name":"Institute of Microbial Interactions, Faculty of Mathematics and Natural Sciences, Heinrich-Heine-University D\u00fcsseldorf , Universit\u00e4tsstra\u00dfe 1, 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seung-Hyun","family":"Paik","sequence":"additional","affiliation":[{"name":"Institute of Molecular Enzyme Technology, IMET, Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf , Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michelle","family":"Bund","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthias","family":"Pesch","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stephan","family":"Krueger","sequence":"additional","affiliation":[{"name":"CEPLAS Metabolism and Metabolomics Laboratory, Cluster of Excellence on Plant Sciences (CEPLAS), Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf , Universit\u00e4tsstra\u00dfe 1, 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Benita","family":"L\u00fcckel","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Athanasios","family":"Papadopoulos","sequence":"additional","affiliation":[{"name":"Center for Structural Studies, Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf, Universit\u00e4tsstra\u00dfe 1 , 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johanna","family":"Wiechert","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tanvir","family":"Hassan","sequence":"additional","affiliation":[{"name":"Computational Life Science, Department of Biology, RWTH Aachen University , 52074 Worringerweg 3, Aachen ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ulrike","family":"Weber","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]},{"name":"Institute of Microbial Interactions, Faculty of Mathematics and Natural Sciences, Heinrich-Heine-University D\u00fcsseldorf , Universit\u00e4tsstra\u00dfe 1, 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Romain","family":"Avellan","sequence":"additional","affiliation":[{"name":"Institute of Biology, Leiden University , Sylviusweg 72, 2333 Leiden ,","place":["Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sander H J","family":"Smits","sequence":"additional","affiliation":[{"name":"Center for Structural Studies, Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf, Universit\u00e4tsstra\u00dfe 1 , 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Bott","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c1kos T","family":"Kov\u00e1cs","sequence":"additional","affiliation":[{"name":"Institute of Biology, Leiden University , Sylviusweg 72, 2333 Leiden ,","place":["Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3494-9420","authenticated-orcid":false,"given":"Philipp","family":"Westhoff","sequence":"additional","affiliation":[{"name":"CEPLAS Metabolism and Metabolomics Laboratory, Cluster of Excellence on Plant Sciences (CEPLAS), Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf , Universit\u00e4tsstra\u00dfe 1, 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna","family":"Matuszy\u0144ska","sequence":"additional","affiliation":[{"name":"Computational Life Science, Department of Biology, RWTH Aachen University , 52074 Worringerweg 3, Aachen ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6215-1857","authenticated-orcid":false,"given":"Dietrich","family":"Kohlheyer","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0096-8084","authenticated-orcid":false,"given":"Thomas","family":"Drepper","sequence":"additional","affiliation":[{"name":"Institute of Molecular Enzyme Technology, IMET, Faculty of Mathematics and Natural Sciences, Heinrich Heine University D\u00fcsseldorf , Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6209-7950","authenticated-orcid":false,"given":"Julia","family":"Frunzke","sequence":"additional","affiliation":[{"name":"Institute of Bio- and Geosciences 1 , IBG-1: Biotechnology, Forschungszentrum J\u00fclich, Wilhelm-Johnen-Stra\u00dfe, 52428 J\u00fclich ,","place":["Germany"]},{"name":"Institute of Microbial Interactions, Faculty of Mathematics and Natural Sciences, Heinrich-Heine-University D\u00fcsseldorf , Universit\u00e4tsstra\u00dfe 1, 40225 D\u00fcsseldorf ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,16]]},"reference":[{"key":"2026100408265083500_ref1","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/s0168-6445(03)00055-x","article-title":"Bacterial iron homeostasis","volume":"27","author":"Andrews","year":"2003","journal-title":"FEMS Microbiol Rev"},{"key":"2026100408265083500_ref2","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1038\/s41579-019-0284-4","article-title":"Bacterial siderophores in community and host interactions","volume":"18","author":"Kramer","year":"2020","journal-title":"Nat Rev Microbiol"},{"key":"2026100408265083500_ref3","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1023\/a:1009227206890","article-title":"Salicylic acid is not a bacterial siderophore: a theoretical study","volume":"13","author":"Chipperfield","year":"2000","journal-title":"Biometals"},{"key":"2026100408265083500_ref4","doi-asserted-by":"publisher","first-page":"413","DOI":"10.1128\/mmbr.00012-07","article-title":"Siderophore-based iron acquisition and pathogen control","volume":"71","author":"Miethke","year":"2007","journal-title":"Microbiol Mol Biol Rev"},{"key":"2026100408265083500_ref5","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1146\/annurev.nu.01.070181.000331","article-title":"Iron absorption and transport in microorganisms","volume":"1","author":"Neilands","year":"1981","journal-title":"Annu Rev Nutr"},{"key":"2026100408265083500_ref6","doi-asserted-by":"publisher","first-page":"2496","DOI":"10.1021\/acs.accounts.5b00301","article-title":"Coordination chemistry of microbial iron transport","volume":"48","author":"Raymond","year":"2015","journal-title":"Acc Chem Res"},{"key":"2026100408265083500_ref7","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1111\/1751-7915.12117","article-title":"Siderophores in environmental research: roles and applications","volume":"7","author":"Ahmed","year":"2014","journal-title":"Microb Biotechnol"},{"key":"2026100408265083500_ref8","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1039\/b906679a","article-title":"Chemistry and biology of siderophores","volume":"27","author":"Hider","year":"2010","journal-title":"Nat Prod Rep"},{"key":"2026100408265083500_ref9","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1002\/chin.198003104","article-title":"Coordination chemistry of microbial iron transport compounds. 19. Stability constants and electrochemical behavior of ferric enterobactin and model complexes","volume":"11","author":"Harris","year":"1979","journal-title":"ChemInform"},{"key":"2026100408265083500_ref10","doi-asserted-by":"publisher","DOI":"10.1099\/mic.0.001688","article-title":"Microbial primer: bacterial metallophores \u2013 their role for metal homeostasis and social interactions","volume":"172","author":"Laffont","year":"2026","journal-title":"Microbiology"},{"key":"2026100408265083500_ref11","doi-asserted-by":"publisher","first-page":"14536","DOI":"10.1016\/S0021-9258(18)77336-5","article-title":"Molecular analysis of the Escherichia coli ferric enterobactin receptor FepA","volume":"265","author":"Armstrong","year":"1990","journal-title":"J Biol Chem"},{"key":"2026100408265083500_ref12","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1023\/a:1009270302536","article-title":"Sequence heterogeneity of the ferripyoverdine uptake (fpvA), but not the ferric uptake regulator (fur), genes among strains of the fluorescent pseudomonads Pseudomonas aeruginosa, pseudomonas aureofaciens, Pseudomonas fluorescens and pseudomonas putida","volume":"12","author":"Thupvong","year":"1999","journal-title":"Biometals"},{"key":"2026100408265083500_ref13","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1111\/j.1365-2958.2007.05905.x","article-title":"Substrate induction of siderophore transport in Bacillus subtilis mediated by a novel one-component regulator","volume":"66","author":"Gaballa","year":"2007","journal-title":"Mol Microbiol"},{"key":"2026100408265083500_ref14","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1016\/j.febslet.2012.11.027","article-title":"Uptake of xenosiderophores in Bacillus subtilis occurs with high affinity and enhances the folding stabilities of substrate binding proteins","volume":"587","author":"Miethke","year":"2013","journal-title":"FEBS Lett"},{"key":"2026100408265083500_ref15","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1111\/j.1758-2229.2009.00041.x","article-title":"A survey of TonB-dependent receptors in fluorescent pseudomonads","volume":"1","author":"Cornelis","year":"2009","journal-title":"Environ Microbiol Rep"},{"key":"2026100408265083500_ref16","doi-asserted-by":"publisher","first-page":"75","DOI":"10.3389\/fcimb.2013.00075","article-title":"Pseudomonas aeruginosa adapts its iron uptake strategies in function of the type of infections","volume":"3","author":"Cornelis","year":"2013","journal-title":"Front Cell Infect Microbiol"},{"key":"2026100408265083500_ref17","doi-asserted-by":"publisher","first-page":"254","DOI":"10.1016\/j.chembiol.2010.02.010","article-title":"Siderophores from neighboring organisms promote the growth of uncultured bacteria","volume":"17","author":"D'Onofrio","year":"2010","journal-title":"Chem Biol"},{"key":"2026100408265083500_ref18","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1146\/annurev.mi.36.100182.001441","article-title":"Microbial envelope proteins related to iron","volume":"36","author":"Neilands","year":"1982","journal-title":"Ann Rev Microbiol"},{"key":"2026100408265083500_ref19","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1016\/0006-291x(75)90375-7","article-title":"In vitro competition between ferrichrome and phage for the outer membrane T5 receptor complex of Escherichia coli","volume":"64","author":"Luckey","year":"1975","journal-title":"Biochem Biophys Res Commun"},{"key":"2026100408265083500_ref20","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1111\/j.1365-2958.2007.06061.x","article-title":"Characterization of five novel Pseudomonas aeruginosa cell-surface signalling systems","volume":"67","author":"Llamas","year":"2008","journal-title":"Mol Microbiol"},{"key":"2026100408265083500_ref21","doi-asserted-by":"publisher","first-page":"1024","DOI":"10.1038\/nature02744","article-title":"Cooperation and competition in pathogenic bacteria","volume":"430","author":"Griffin","year":"2004","journal-title":"Nature"},{"key":"2026100408265083500_ref22","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1098\/rspb.2002.2209","article-title":"Cooperation, virulence and siderophore production in bacterial parasites","volume":"270","author":"West","year":"2003","journal-title":"Proc Biol Sci"},{"key":"2026100408265083500_ref23","doi-asserted-by":"publisher","first-page":"1443","DOI":"10.1111\/evo.13230","article-title":"The evolution of siderophore production as a competitive trait","volume":"71","author":"Niehus","year":"2017","journal-title":"Evolution"},{"key":"2026100408265083500_ref24","doi-asserted-by":"publisher","first-page":"414","DOI":"10.1038\/s41467-017-00509-4","article-title":"Siderophore cheating and cheating resistance shape competition for iron in soil and freshwater pseudomonas communities","volume":"8","author":"Butait\u0117","year":"2017","journal-title":"Nat Commun"},{"key":"2026100408265083500_ref25","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1007\/s11274-025-04287-w","article-title":"Xenosiderophores: bridging the gap in microbial iron acquisition strategies","volume":"41","author":"Kumar","year":"2025","journal-title":"World J Microbiol Biotechnol"},{"key":"2026100408265083500_ref26","doi-asserted-by":"publisher","first-page":"3132","DOI":"10.1128\/aem.03520-14","article-title":"Pseudomonas fluorescens pirates both ferrioxamine and ferricoelichelin siderophores from Streptomyces ambofaciens","volume":"81","author":"Galet","year":"2015","journal-title":"Appl Environ Microbiol"},{"key":"2026100408265083500_ref27","doi-asserted-by":"publisher","first-page":"wrae123","DOI":"10.1093\/ismejo\/wrae123","article-title":"Nasal commensals reduce Staphylococcus aureus proliferation by restricting siderophore availability","volume":"18","author":"Zhao","year":"2024","journal-title":"ISME J"},{"key":"2026100408265083500_ref28","doi-asserted-by":"publisher","first-page":"92493","DOI":"10.7554\/eLife.92493","article-title":"Antimicrobial activity of iron-depriving pyoverdines against human opportunistic pathogens","volume":"13","author":"Vollenweider","year":"2024","journal-title":"elife"},{"key":"2026100408265083500_ref29","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1038\/s41579-024-01090-6","article-title":"Bacterial siderophores: diversity, uptake pathways and applications","volume":"23","author":"Schalk","year":"2025","journal-title":"Nat Rev Microbiol"},{"key":"2026100408265083500_ref30","doi-asserted-by":"publisher","first-page":"1435","DOI":"10.1021\/acschembio.5c00286","article-title":"Identification and characterization of the biosynthesis of the hybrid NRPS-NIS siderophore Nocardichelin","volume":"20","author":"Fisk","year":"2025","journal-title":"ACS Chem Biol"},{"key":"2026100408265083500_ref31","doi-asserted-by":"publisher","first-page":"1150","DOI":"10.1007\/s43393-024-00257-9","article-title":"Characterization of siderophore from probiotic bacillus spp. strain isolated from traditional fermented food of the Himalaya","volume":"4","author":"Goswami","year":"2024","journal-title":"Syst Microbiol Biomanuf"},{"key":"2026100408265083500_ref32","doi-asserted-by":"publisher","first-page":"1277176","DOI":"10.3389\/fcimb.2023.1277176","article-title":"Microbial interaction-induced siderophore dynamics lead to phenotypic differentiation of Staphylococcus aureus. Original research","volume":"13","author":"Rajapitamahuni","year":"2023","journal-title":"Front Cell Infect Microbiol"},{"key":"2026100408265083500_ref33","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1186\/s40168-024-01913-x","article-title":"The Staphylococcus aureus-antagonizing human nasal commensal staphylococcus lugdunensis depends on siderophore piracy","volume":"12","author":"Rosenstein","year":"2024","journal-title":"Microbiome"},{"key":"2026100408265083500_ref34","doi-asserted-by":"publisher","DOI":"10.1093\/ismejo\/wrad001","article-title":"Competition for iron shapes metabolic antagonism between Bacillus subtilis and Pseudomonas marginalis","volume":"18","author":"Lyng","year":"2024","journal-title":"ISME J"},{"key":"2026100408265083500_ref35","doi-asserted-by":"publisher","first-page":"2025.05.12.653243","DOI":"10.1101\/2025.05.12.653243","article-title":"Siderophores can alter the population dynamics of fungal-bacterial communities by inhibiting specialized metabolism","author":"Pham","year":"2025"},{"key":"2026100408265083500_ref36","doi-asserted-by":"publisher","first-page":"2026.02.24.707706","DOI":"10.64898\/2026.02.24.707706","article-title":"Metabolic exchange and siderophore sharing underlie emergent biofilm synergism","author":"Xie","year":"2026"},{"key":"2026100408265083500_ref37","doi-asserted-by":"publisher","first-page":"1087","DOI":"10.1007\/s00775-006-0151-4","article-title":"Siderophore-mediated iron transport in Bacillus subtilis and Corynebacterium glutamicum","volume":"11","author":"Dertz","year":"2006","journal-title":"J Biol Inorg Chem"},{"key":"2026100408265083500_ref38","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1021\/acssynbio.7b00261","article-title":"Corynebacterium glutamicum chassis C1*: building and testing a novel platform host for synthetic biology and industrial biotechnology","volume":"7","author":"Baumgart","year":"2018","journal-title":"ACS Synth Biol"},{"key":"2026100408265083500_ref39","doi-asserted-by":"publisher","DOI":"10.1128\/aem.00065-20","article-title":"The iron deficiency response of Corynebacterium glutamicum and a link to thiamine biosynthesis","volume":"86","author":"K\u00fcberl","year":"2020","journal-title":"Appl Environ Microbiol"},{"key":"2026100408265083500_ref40","doi-asserted-by":"publisher","first-page":"5595","DOI":"10.1128\/jb.175.17.5595-5603.1993","article-title":"Isoleucine synthesis in Corynebacterium glutamicum: molecular analysis of the ilvB-ilvN-ilvC operon","volume":"175","author":"Keilhauer","year":"1993","journal-title":"J Bacteriol"},{"key":"2026100408265083500_ref41","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1007\/BF00165889","article-title":"Requirement of chelating compounds for the growth of Corynebacterium glutamicum in synthetic media","volume":"32","author":"Liebl","year":"1989","journal-title":"Appl Microbiol Biotechnol"},{"key":"2026100408265083500_ref42","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1186\/1475-2859-8-31","article-title":"Validation of a high-throughput fermentation system based on online monitoring of biomass and fluorescence in continuously shaken microtiter plates","volume":"8","author":"Kensy","year":"2009","journal-title":"Microb Cell Factories"},{"key":"2026100408265083500_ref43","doi-asserted-by":"publisher","first-page":"6949","DOI":"10.1128\/iai.00866-06","article-title":"Identification and characterization of a novel ABC iron transport system, fit. Escherichia coli","volume":"74","author":"Ouyang","year":"2006","journal-title":"Infect Immun"},{"key":"2026100408265083500_ref44","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1007\/s11274-013-1425-y","article-title":"Incorporation of L-lysine and 2,2\u2032-dipyridyl in the growth media promotes desferrioxamine E production by an actinobacterium","volume":"30","author":"Nakouti","year":"2014","journal-title":"World J Microbiol Biotechnol"},{"key":"2026100408265083500_ref45","doi-asserted-by":"publisher","first-page":"1143861","DOI":"10.3389\/fmicb.2023.1143861","article-title":"Two new siderophores produced by pseudomonas sp. NCIMB 10586: the anti-oomycete non-ribosomal peptide synthetase-dependent mupirochelin and the NRPS-independent triabactin","volume":"14","author":"Grosse","year":"2023","journal-title":"Front Microbiol"},{"key":"2026100408265083500_ref46","doi-asserted-by":"publisher","first-page":"545","DOI":"10.1038\/s42003-022-03493-8","article-title":"Coordination of siderophore gene expression among clonal cells of the bacterium Pseudomonas aeruginosa","volume":"5","author":"Mridha","year":"2022","journal-title":"Commun Biol"},{"key":"2026100408265083500_ref47","volume-title":"Molecular Cloning: A Laboratory Manual","author":"Green","year":"2012","edition":"4"},{"key":"2026100408265083500_ref48","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1038\/nmeth.1318","article-title":"Enzymatic assembly of DNA molecules up to several hundred kilobases","volume":"6","author":"Gibson","year":"2009","journal-title":"Nat Methods"},{"key":"2026100408265083500_ref49","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1007\/s002530051557","article-title":"A heat shock following electroporation induces highly efficient transformation of Corynebacterium glutamicum with xenogeneic plasmid DNA","volume":"52","author":"van der Rest","year":"1999","journal-title":"Appl Microbiol Biotechnol"},{"key":"2026100408265083500_ref50","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/0378-1119(94)90324-7","article-title":"Small mobilizable multi-purpose cloning vectors derived from the Escherichia coli plasmids pK18 and pK19: selection of defined deletions in the chromosome of Corynebacterium glutamicum","volume":"145","author":"Sch\u00e4fer","year":"1994","journal-title":"Gene"},{"key":"2026100408265083500_ref51","doi-asserted-by":"publisher","first-page":"282","DOI":"10.1007\/s002030100262","article-title":"Molecular analysis of the cytochrome bc1-aa3 branch of the Corynebacterium glutamicum respiratory chain containing an unusual diheme cytochrome c1","volume":"175","author":"Niebisch","year":"2001","journal-title":"Arch Microbiol"},{"key":"2026100408265083500_ref52","doi-asserted-by":"publisher","first-page":"2172","DOI":"10.1073\/pnas.86.7.2172","article-title":"One-step preparation of competent Escherichia coli: transformation and storage of bacterial cells in the same solution","volume":"86","author":"Chung","year":"1989","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026100408265083500_ref53","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/0076-6879(91)04006-a","article-title":"Plasmid transformation of Escherichia coli and other bacteria","volume":"204","author":"Hanahan","year":"1991","journal-title":"Methods Enzymol"},{"key":"2026100408265083500_ref54","doi-asserted-by":"publisher","first-page":"6640","DOI":"10.1073\/pnas.120163297","article-title":"One-step inactivation of chromosomal genes in Escherichia coli K-12 using PCR products","volume":"97","author":"Datsenko","year":"2000","journal-title":"Proc Natl Acad Sci"},{"key":"2026100408265083500_ref55","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat Methods"},{"key":"2026100408265083500_ref56","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1002\/mrd.22489","article-title":"The ImageJ ecosystem: an open platform for biomedical image analysis","volume":"82","author":"Schindelin","year":"2015","journal-title":"Mol Reprod Dev"},{"key":"2026100408265083500_ref57","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1021\/acssensors.5c02481","article-title":"A novel biosensor for ferrous iron developed via CoBiSe: a computational method for rapid biosensor design","volume":"11","author":"Papadopoulos","year":"2026","journal-title":"ACS Sens"},{"key":"2026100408265083500_ref58","doi-asserted-by":"crossref","first-page":"6167","DOI":"10.1021\/ac070470c","article-title":"Acidic acetonitrile for cellular metabolome extraction from Escherichia coli","volume":"79","author":"Rabinowitz","year":"2007","journal-title":"Anal Chem"},{"key":"2026100408265083500_ref59","article-title":"Acia-workflows: automated single-cell imaging analysis for scalable and deep learning-based live-cell imaging analysis workflows","author":"Seiffarth","year":"2025"},{"key":"2026100408265083500_ref60","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1016\/s0304-4165(98)00089-0","article-title":"The role of isochorismate hydroxymutase genes entC and menF in enterobactin and menaquinone biosynthesis in Escherichia coli","volume":"1425","author":"Dahm","year":"1998","journal-title":"Biochim Biophys Acta"},{"key":"2026100408265083500_ref61","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1007\/BF01026778","article-title":"Design of a defined medium for growth of Corynebacterium glutamicum in which citrate facilitates iron uptake","volume":"11","author":"von der Osten","year":"1989","journal-title":"Biotechnol Lett"},{"key":"2026100408265083500_ref62","doi-asserted-by":"publisher","first-page":"2069","DOI":"10.1038\/ismej.2014.52","article-title":"Density of founder cells affects spatial pattern formation and cooperation in Bacillus subtilis biofilms","volume":"8","author":"van Gestel","year":"2014","journal-title":"ISME J"},{"key":"2026100408265083500_ref63","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2025.0169","article-title":"Exploring negative frequency-dependent selection across levels: from genetics to ecology and back again","volume":"381","author":"Fraser","year":"2026","journal-title":"Philos Trans R Soc B: Biol Sci"},{"key":"2026100408265083500_ref64","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1146\/annurev.es.05.110174.000555","article-title":"Frequency-dependent selection","volume":"5","author":"Ayala","year":"1974","journal-title":"Annu Rev Ecol Evol Syst"},{"key":"2026100408265083500_ref65","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1126\/science.162.3859.1243","article-title":"The tragedy of the commons","volume":"162","author":"Hardin","year":"1968","journal-title":"Science"},{"key":"2026100408265083500_ref66","doi-asserted-by":"publisher","first-page":"128","DOI":"10.1002\/pst.426","article-title":"Guidelines for accurate EC50\/IC50 estimation","volume":"10","author":"Sebaugh","year":"2011","journal-title":"Pharm Stat"},{"key":"2026100408265083500_ref67","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1128\/jb.177.1.252-258.1995","article-title":"Nucleotide sequence of pvdD, a pyoverdine biosynthetic gene from Pseudomonas aeruginosa: PvdD has similarity to peptide synthetases","volume":"177","author":"Merriman","year":"1995","journal-title":"J Bacteriol"},{"key":"2026100408265083500_ref68","doi-asserted-by":"publisher","first-page":"3673","DOI":"10.1038\/s41467-019-11508-y","article-title":"The complex of ferric-enterobactin with its transporter from Pseudomonas aeruginosa suggests a two-site model","volume":"10","author":"Moyni\u00e9","year":"2019","journal-title":"Nat Commun"},{"key":"2026100408265083500_ref69","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1111\/j.1574-6941.2007.00388.x","article-title":"Siderophore-mediated cooperation and virulence in Pseudomonas aeruginosa","volume":"62","author":"Buckling","year":"2007","journal-title":"FEMS Microbiol Ecol"},{"key":"2026100408265083500_ref70","doi-asserted-by":"publisher","first-page":"R442","DOI":"10.1016\/j.cub.2019.03.001","article-title":"Public goods and cheating in microbes","volume":"29","author":"Smith","year":"2019","journal-title":"Curr Biol"},{"key":"2026100408265083500_ref71","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1038\/ismej.2007.96","article-title":"Interspecific competition and siderophore-mediated cooperation in Pseudomonas aeruginosa","volume":"2","author":"Harrison","year":"2008","journal-title":"ISME J"},{"key":"2026100408265083500_ref72","doi-asserted-by":"publisher","first-page":"1383","DOI":"10.1038\/s41467-018-03791-y","article-title":"Conditional privatization of a public siderophore enables Pseudomonas aeruginosa to resist cheater invasion","volume":"9","author":"Jin","year":"2018","journal-title":"Nat Commun"},{"key":"2026100408265083500_ref73","doi-asserted-by":"publisher","first-page":"2092","DOI":"10.1021\/cb500319n","article-title":"Direct evidence of iron uptake by the gram-positive siderophore-shuttle mechanism without iron reduction","volume":"9","author":"Fukushima","year":"2014","journal-title":"ACS Chem Biol"},{"key":"2026100408265083500_ref74","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00036-12","article-title":"The black queen hypothesis: evolution of dependencies through adaptive gene loss","volume":"3","author":"Morris","year":"2012","journal-title":"MBio"},{"key":"2026100408265083500_ref75","doi-asserted-by":"publisher","first-page":"2085","DOI":"10.1038\/ismej.2016.22","article-title":"Beyond the black queen hypothesis","volume":"10","author":"Mas","year":"2016","journal-title":"ISME J"},{"key":"2026100408265083500_ref76","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1038\/ismej.2016.195","article-title":"Cheating fosters species co-existence in well-mixed bacterial communities","volume":"11","author":"Leinweber","year":"2017","journal-title":"ISME J"},{"key":"2026100408265083500_ref77","doi-asserted-by":"publisher","first-page":"R661","DOI":"10.1016\/j.cub.2007.06.004","article-title":"Evolutionary explanations for cooperation","volume":"17","author":"West","year":"2007","journal-title":"Curr Biol"},{"key":"2026100408265083500_ref78","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1086\/519860","article-title":"Frequency dependence and cooperation: theory and a test with bacteria","volume":"170","author":"Ross-Gillespie","year":"2007","journal-title":"Am Nat"},{"key":"2026100408265083500_ref79","doi-asserted-by":"publisher","first-page":"12577","DOI":"10.1073\/pnas.1301428110","article-title":"Cell-cell contacts confine public goods diffusion inside Pseudomonas aeruginosa clonal microcolonies","volume":"110","author":"Julou","year":"2013","journal-title":"Proc Natl Acad Sci"},{"key":"2026100408265083500_ref80","doi-asserted-by":"publisher","first-page":"e01169","DOI":"10.7554\/eLife.01169","article-title":"Spatial dilemmas of diffusible public goods","volume":"2","author":"Allen","year":"2013","journal-title":"elife"},{"key":"2026100408265083500_ref81","doi-asserted-by":"publisher","first-page":"2122","DOI":"10.1128\/jb.02596-14","article-title":"Sociality in Escherichia coli: enterochelin is a private good at low cell density and can Be shared at high cell density","volume":"197","author":"Scholz","year":"2015","journal-title":"J Bacteriol"},{"key":"2026100408265083500_ref82","doi-asserted-by":"publisher","first-page":"2166","DOI":"10.1111\/j.1558-5646.2010.00959.x","article-title":"Novel cooperation experimentally evolved between species","volume":"64","author":"Harcombe","year":"2010","journal-title":"Evolution"},{"key":"2026100408265083500_ref83","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/j.chom.2023.03.016","article-title":"Cross-feeding in the gut microbiome: ecology and mechanisms","volume":"31","author":"Culp","year":"2023","journal-title":"Cell Host Microbe"},{"key":"2026100408265083500_ref84","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1038\/ismej.2016.141","article-title":"Microbial mutualism dynamics governed by dose-dependent toxicity of cross-fed nutrients","volume":"11","author":"LaSarre","year":"2017","journal-title":"ISME J"},{"key":"2026100408265083500_ref85","doi-asserted-by":"publisher","first-page":"906","DOI":"10.1021\/ic00005a008","article-title":"Solution equilibria of enterobactin and metal-enterobactin complexes","volume":"30","author":"Loomis","year":"1991","journal-title":"Inorg Chem"},{"key":"2026100408265083500_ref86","doi-asserted-by":"publisher","first-page":"6320","DOI":"10.1039\/C4DT03559C","article-title":"Beyond iron: non-classical biological functions of bacterial siderophores","volume":"44","author":"Johnstone","year":"2015","journal-title":"Dalton Trans"},{"key":"2026100408265083500_ref87","doi-asserted-by":"publisher","first-page":"6391","DOI":"10.1021\/ic00104a059","article-title":"Bacterial iron transport: coordination properties of Pyoverdin PaA, a peptidic siderophore of Pseudomonas aeruginosa","volume":"33","author":"Albrecht-Gary","year":"1994","journal-title":"Inorg Chem"},{"key":"2026100408265083500_ref88","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1007\/s00027-003-0690-5","article-title":"Iron oxide dissolution and solubility in the presence of siderophores","volume":"66","author":"Kraemer","year":"2004","journal-title":"Aquat Sci"},{"key":"2026100408265083500_ref89","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1111\/ele.13912","article-title":"Siderophores drive invasion dynamics in bacterial communities through their dual role as public good versus public bad","volume":"25","author":"Figueiredo","year":"2022","journal-title":"Ecol Lett"},{"key":"2026100408265083500_ref90","doi-asserted-by":"publisher","first-page":"20440","DOI":"10.1073\/pnas.0908760106","article-title":"Molecular basis of pyoverdine siderophore recycling in Pseudomonas aeruginosa","volume":"106","author":"Imperi","year":"2009","journal-title":"Proc Natl Acad Sci"},{"key":"2026100408265083500_ref91","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1007\/BF00140485","article-title":"HPLC separation of enterobactin and linear 2,3-dihydroxybenzoylserine derivatives: a study on mutants of Escherichia coli defective in regulation (fur), esterase (Fes) and transport (fepA)","volume":"7","author":"Winkelmann","year":"1994","journal-title":"Biometals"},{"key":"2026100408265083500_ref92","doi-asserted-by":"publisher","first-page":"fuaf067","DOI":"10.1093\/femsre\/fuaf067","article-title":"Spatial structure: shaping the ecology and evolution of microbial communities","volume":"50","author":"B\u00e4cker","year":"2026","journal-title":"FEMS Microbiol Rev"},{"key":"2026100408265083500_ref93","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1007\/s10482-025-02098-5","article-title":"Siderophores produced by marine microorganisms","volume":"118","author":"Chen","year":"2025","journal-title":"Antonie Van Leeuwenhoek"},{"key":"2026100408265083500_ref94","doi-asserted-by":"publisher","first-page":"1536","DOI":"10.1111\/ele.12371","article-title":"Habitat structure and the evolution of diffusible siderophores in bacteria","volume":"17","author":"K\u00fcmmerli","year":"2014","journal-title":"Ecol Lett"},{"key":"2026100408265083500_ref95","doi-asserted-by":"publisher","first-page":"1157","DOI":"10.1038\/nature03891","article-title":"The contribution of species richness and composition to bacterial services","volume":"436","author":"Bell","year":"2005","journal-title":"Nature"},{"key":"2026100408265083500_ref96","doi-asserted-by":"publisher","DOI":"10.1093\/ismejo\/wraf025","article-title":"Strain identity effects contribute more to pseudomonas community functioning than strain interactions","volume":"19","author":"Kramer","year":"2025","journal-title":"ISME J"},{"key":"2026100408265083500_ref97","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1016\/j.tig.2015.09.001","article-title":"Buried treasure: evolutionary perspectives on microbial iron piracy","volume":"31","author":"Barber","year":"2015","journal-title":"Trends Genet"},{"key":"2026100408265083500_ref98","doi-asserted-by":"publisher","first-page":"20059","DOI":"10.1073\/pnas.1213344109","article-title":"Public good dynamics drive evolution of iron acquisition strategies in natural bacterioplankton populations","volume":"109","author":"Cordero","year":"2012","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026100408265083500_ref99","doi-asserted-by":"publisher","first-page":"1002","DOI":"10.1038\/s41564-020-0719-8","article-title":"Competition for iron drives phytopathogen control by natural rhizosphere microbiomes","volume":"5","author":"Gu","year":"2020","journal-title":"Nat Microbiol"},{"key":"2026100408265083500_ref100","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1186\/s12862-023-02152-8","article-title":"Community composition drives siderophore dynamics in multispecies bacterial communities","volume":"23","author":"O\u2019Brien","year":"2023","journal-title":"BMC Ecol Evol"},{"key":"2026100408265083500_ref101","doi-asserted-by":"publisher","first-page":"898979","DOI":"10.3389\/fmicb.2022.898979","article-title":"Mechanistic insights and potential use of Siderophores producing microbes in rhizosphere for mitigation of stress in plants grown in degraded land","volume":"13","author":"Singh","year":"2022","journal-title":"Front Microbiol"},{"key":"2026100408265083500_ref102","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1016\/j.sajb.2023.12.031","article-title":"Siderophore producing bacteria as biocontrol agent against phytopathogens for a better environment: a review","volume":"165","author":"Deb","year":"2024","journal-title":"S Afr J Bot"},{"key":"2026100408265083500_ref103","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1016\/j.freeradbiomed.2016.10.489","article-title":"Siderophore-mediated iron acquisition and modulation of host-bacterial interactions","volume":"105","author":"Ellermann","year":"2017","journal-title":"Free Radic Biol Med"},{"key":"2026100408265083500_ref104","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1016\/j.tips.2014.06.007","article-title":"Siderophore-drug complexes: potential medicinal applications of the 'Trojan horse' strategy","volume":"35","author":"G\u00f3rska","year":"2014","journal-title":"Trends Pharmacol Sci"}],"container-title":["The ISME Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/ismej\/advance-article-pdf\/doi\/10.1093\/ismejo\/wrag247\/71123659\/wrag247.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismej\/article-pdf\/20\/1\/wrag247\/71123659\/wrag247.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/ismej\/article-pdf\/20\/1\/wrag247\/71123659\/wrag247.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,4]],"date-time":"2026-10-04T12:26:57Z","timestamp":1791116817000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/ismej\/article\/doi\/10.1093\/ismejo\/wrag247\/8802179"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1]]},"references-count":104,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,1,14]]}},"URL":"https:\/\/doi.org\/10.1093\/ismejo\/wrag247","relation":{},"ISSN":["1751-7362","1751-7370"],"issn-type":[{"value":"1751-7362","type":"print"},{"value":"1751-7370","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1]]},"published":{"date-parts":[[2026,1]]},"article-number":"wrag247"}}