{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,19]],"date-time":"2026-10-19T23:42:25Z","timestamp":1792453345591,"version":"4.5.0"},"reference-count":65,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T00:00:00Z","timestamp":1792454400000},"content-version":"vor","delay-in-days":89,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Appl Microbiol Biotechnol"],"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Fungal pigments hold significant food industrial value due to their safe and sustainable properties. Here we identified a naturally occurring coculture pair\n                      <jats:italic>Epicoccum nigrum<\/jats:italic>\n                      SPDX618 and\n                      <jats:italic>Mucor racemosus<\/jats:italic>\n                      SPDX522 in vinegar mash that produced red pigments, which was not observed in either monoculture. Our results showed that\n                      <jats:italic>E. nigrum<\/jats:italic>\n                      acted as the pigment producer while\n                      <jats:italic>M. racemosus<\/jats:italic>\n                      served as the elicitor. Response surface methodology and pH stabilization strategy improved the yield of red pigment into 328.7 U\/g. Through bioactivity-guided fractionation coupled with UHPLC-MS\/MS and in silico structure reconstitution, four red pigments were preliminarily assigned as anthracycline-like compounds. RNA-seq analysis revealed a putative dual-PKS gene cluster (SPDX618009169\u2013SPDX618009178) with exclusively upregulation in cocultured\n                      <jats:italic>E. nigrum<\/jats:italic>\n                      that is potentially involved in red pigment biosynthesis. Besides, the production of the yellow pigment epipyrone was reduced in cocultured\n                      <jats:italic>E. nigrum<\/jats:italic>\n                      , consistent with the downregulated transcription level of its biosynthesis genes. Comparative metabolomics analysis further revealed a broad downregulation in both of primary metabolism and secondary metabolism in coculture system, indicating that a competition interaction relationship also existed between\n                      <jats:italic>E. nigrum<\/jats:italic>\n                      and\n                      <jats:italic>M. racemosus<\/jats:italic>\n                      . Altogether, this study reports a fungal coculture system that exhibited red pigments production. These findings contribute to the understanding of microbial interaction-driven metabolite biosynthesis and may provide a foundation for future exploration of their potential application as a colorant.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      \u2022\u00a0\n                      <jats:italic>An Epicoccum nigrum\u2013Mucor racemosus coculture system was identified to produce red pigment for the first time.<\/jats:italic>\n                    <\/jats:p>\n                    <jats:p>\n                      \u2022\n                      <jats:italic>E. nigrum acts as the red pigment producer and M. racemosus serves as the elicitor.<\/jats:italic>\n                    <\/jats:p>\n                    <jats:p>\n                      \u2022\n                      <jats:italic>RNA-seq analysis tentatively identified a putative dual-PKS gene cluster upregulated in coculture that is putatively involved in red pigment biosynthesis.<\/jats:italic>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1007\/s00253-026-13956-1","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T13:34:09Z","timestamp":1784813649000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A coculture system of Epicoccum nigrum with Mucor racemosus induces red pigment biosynthesis in vinegar mash"],"prefix":"10.1007","volume":"110","author":[{"given":"Fengli","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fangyuan","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinle","family":"Liang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4725-6962","authenticated-orcid":false,"given":"Wenjie","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,23]]},"reference":[{"issue":"5","key":"13956_CR1","doi-asserted-by":"publisher","first-page":"2159","DOI":"10.1093\/nar\/gky066","volume":"46","author":"S Ardui","year":"2018","unstructured":"Ardui S, Ameur A, Vermeesch JR, Hestand MS (2018) Single molecule real-time (SMRT) sequencing comes of age: applications and utilities for medical diagnostics. Nucleic Acids Res 46(5):2159\u20132168. https:\/\/doi.org\/10.1093\/nar\/gky066","journal-title":"Nucleic Acids Res"},{"issue":"1","key":"13956_CR2","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1038\/75556","volume":"25","author":"M Ashburner","year":"2000","unstructured":"Ashburner M, Ball CA, Blake JA, Botstein D, Butler H, Cherry JM, Davis AP, Dolinski K, Dwight SS, Eppig JT, Harris MA, Hill DP, Issel-Tarver L, Kasarskis A, Lewis S, Matese JC, Richardson JE, Ringwald M, Rubin GM, Sherlock G (2000) Gene ontology: tool for the unification of biology. Nat Genet 25(1):25\u201329. https:\/\/doi.org\/10.1038\/75556","journal-title":"Nat Genet"},{"issue":"9","key":"13956_CR3","doi-asserted-by":"publisher","first-page":"1640","DOI":"10.1016\/j.tetlet.2014.01.090","volume":"55","author":"B Balakrishnan","year":"2014","unstructured":"Balakrishnan B, Che C-C, Pan T-M, Kwon H-J (2014) Mpp7 controls regioselective Knoevenagel condensation during the biosynthesis of Monascus azaphilone pigments. Tetrahedron Lett 55(9):1640\u20131643. https:\/\/doi.org\/10.1016\/j.tetlet.2014.01.090","journal-title":"Tetrahedron Lett"},{"key":"13956_CR4","doi-asserted-by":"crossref","unstructured":"Blin K, Shaw S, Augustijn HE, Reitz ZL, Biermann F, Alanjary M, Fetter A, Terlouw BR, Metcalf WW, Helfrich EJN, Wezel GPV, Medema MH, Weber T (2023) AntiSMASH 7.0: new and improved predictions for detection, regulation, chemical structures and visualisation. Nucleic Acids Res 51(W1):W46-W50 ARTN gkad34410.1093\/nar\/gkad344","DOI":"10.1093\/nar\/gkad344"},{"issue":"15","key":"13956_CR5","doi-asserted-by":"publisher","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","volume":"30","author":"AM Bolger","year":"2014","unstructured":"Bolger AM, MarcLohse UB (2014) Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30(15):2114\u20132120. https:\/\/doi.org\/10.1093\/bioinformatics\/btu170","journal-title":"Bioinformatics"},{"key":"13956_CR6","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/bs.ircmb.2015.10.001","volume":"321","author":"E Calzada","year":"2016","unstructured":"Calzada E, Onguka O, Claypool SM (2016) Phosphatidylethanolamine metabolism in health and disease. Int Rev Cell Mol Biol 321:29\u201388. https:\/\/doi.org\/10.1016\/bs.ircmb.2015.10.001","journal-title":"Int Rev Cell Mol Biol"},{"key":"13956_CR7","doi-asserted-by":"publisher","DOI":"10.1016\/j.fbio.2025.106453","volume":"68","author":"Z Cheng","year":"2025","unstructured":"Cheng Z, Yang J, Yan R, Wang B, Bai Y, Miao Z, Sun J, Li H, Wang X, Sun B (2025) Interactive mechanism-guided microbial interaction dynamics in food fermentations: lactic acid bacteria and yeasts as a case example. Food Biosci 68:106453. https:\/\/doi.org\/10.1016\/j.fbio.2025.106453","journal-title":"Food Biosci"},{"issue":"8","key":"13956_CR8","doi-asserted-by":"publisher","first-page":"2965","DOI":"10.1021\/ja8088185","volume":"131","author":"Y-M Chiang","year":"2009","unstructured":"Chiang Y-M, Szewczyk E, Davidson AD, Keller N, Oakley BR, Wang CCC (2009) A gene cluster containing two fungal polyketide synthases encodes the biosynthetic pathway for a polyketide, Asperfuranone, in Aspergillus nidulans. J Am Chem Soc 131(8):2965\u20132970. https:\/\/doi.org\/10.1021\/ja8088185","journal-title":"J Am Chem Soc"},{"issue":"W1","key":"13956_CR9","doi-asserted-by":"publisher","first-page":"W486","DOI":"10.1093\/nar\/gky310","volume":"46","author":"J Chong","year":"2018","unstructured":"Chong J, Soufan O, Li C, Caraus I, Li S, Bourque G, Wishart DS, Xia J (2018) MetaboAnalyst 4.0: towards more transparent and integrative metabolomics analysis. Nucleic Acids Res 46(W1):W486\u2013W494. https:\/\/doi.org\/10.1093\/nar\/gky310","journal-title":"Nucleic Acids Res"},{"key":"13956_CR10","doi-asserted-by":"publisher","DOI":"10.1016\/j.rbc.2025.100052","volume":"12","author":"MJ Davies","year":"2025","unstructured":"Davies MJ (2025) Methionine oxidation products as biomarkers of oxidative damage to proteins and modulators of cellular metabolism and toxicity. Redox Biol 12:100052. https:\/\/doi.org\/10.1016\/j.rbc.2025.100052","journal-title":"Redox Biol"},{"issue":"4","key":"13956_CR11","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1038\/s41592-019-0344-8","volume":"16","author":"K D\u00fchrkop","year":"2019","unstructured":"D\u00fchrkop K, Fleischauer M, Melnik AV, Ludwig M, Aksenov AA, Meusel M, Dorrestein PC, Rousu J, B\u00f6cker S (2019) SIRIUS 4: a rapid tool for turning tandem mass spectra into metabolite structure information. Nat Methods 16(4):299\u2013302. https:\/\/doi.org\/10.1038\/s41592-019-0344-8","journal-title":"Nat Methods"},{"issue":"6","key":"13956_CR12","doi-asserted-by":"publisher","first-page":"1160","DOI":"10.1021\/acs.orglett.3c04259","volume":"26","author":"A Fan","year":"2024","unstructured":"Fan A, Zhong B, Liu D, Lu Y, Wu M, Jin H, Shi XM, Ren J, Zhang B, Su XD, Ma M, Li SM, Lin W (2024) Biosynthesis of epipyrone A reveals a highly specific membrane-bound fungal C-glycosyltransferase for pyrone galactosylation. Org Lett 26(6):1160\u20131165. https:\/\/doi.org\/10.1021\/acs.orglett.3c04259","journal-title":"Org Lett"},{"key":"13956_CR13","doi-asserted-by":"publisher","DOI":"10.1016\/j.ifset.2022.103259","volume":"83","author":"G-Y Fang","year":"2023","unstructured":"Fang G-Y, Mu X-J, Huang B-W, Wud G-Z, Jiang Y-J (2023) Fungal biodiversity and interaction complexity were the important drivers of multifunctionality for flavor production in a spontaneously fermented vinegar. Innov Food Sci Emerg Technol 83:103259. https:\/\/doi.org\/10.1016\/j.ifset.2022.103259","journal-title":"Innov Food Sci Emerg Technol"},{"issue":"12","key":"13956_CR14","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0084102","volume":"8","author":"JC Frisvad","year":"2013","unstructured":"Frisvad JC, Yilmaz N, Thrane U, Rasmussen KB, Houbraken J, Samson RA (2013) Talaromyces atroroseus, a new species efficiently producing industrially relevant red pigments. PLoS ONE 8(12):e84102. https:\/\/doi.org\/10.1371\/journal.pone.0084102","journal-title":"PLoS ONE"},{"key":"13956_CR15","doi-asserted-by":"publisher","unstructured":"Garc\u00ed\u00ada-Alcalde F, Okonechnikov K, Carbonell J, M.Cruz L, G\u00f6tz S, Tarazona S, Dopazo J, Meyer TF, Conesa A (2012) Qualimap: evaluating next-generation sequencing alignment data. Bioinformatics 28(20):2678\u20132679. https:\/\/doi.org\/10.1093\/bioinformatics\/bts503","DOI":"10.1093\/bioinformatics\/bts503"},{"issue":"6","key":"13956_CR16","doi-asserted-by":"publisher","first-page":"907","DOI":"10.1016\/S0031-9422(00)86041-0","volume":"6","author":"O Gribanovski-Sassu","year":"1967","unstructured":"Gribanovski-Sassu O, Foppen FH (1967) The carotenoids of the fungus Epicoccum nigrum link. Phytochemistry 6(6):907\u2013909. https:\/\/doi.org\/10.1016\/S0031-9422(00)86041-0","journal-title":"Phytochemistry"},{"key":"13956_CR17","doi-asserted-by":"publisher","DOI":"10.1016\/j.fitote.2020.104698","volume":"146","author":"H Harwoko","year":"2020","unstructured":"Harwoko H, Leec J, Hartmanne R, M\u00e1ndif A, Kurt\u00e1nf T, M\u00fcllerg WEG, Feldbr\u00fcgge M, Kalscheuera R, Ancheevaa E, Daletosa G, Franka M, Liua Z, Proksch P (2020) Azacoccones F-H, new flavipin-derived alkaloids from an endophytic fungus Epicoccumnigrum MK214079. Fitoterapia 146:104698. https:\/\/doi.org\/10.1016\/j.fitote.2020.104698","journal-title":"Fitoterapia"},{"issue":"2","key":"13956_CR18","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1007\/s00253-019-10216-3","volume":"104","author":"T Isbrandt","year":"2019","unstructured":"Isbrandt T, Tolborg G, \u00d8dum A, Workman M, Larsen TO (2019) Atrorosins: a new subgroup of Monascus pigments from Talaromyces atroroseus. Appl Microbiol Biotechnol 104(2):615\u2013622. https:\/\/doi.org\/10.1007\/s00253-019-10216-3","journal-title":"Appl Microbiol Biotechnol"},{"key":"13956_CR19","doi-asserted-by":"publisher","DOI":"10.3390\/plants11182381","author":"X Ji","year":"2022","unstructured":"Ji X, Tang J, Zhang J (2022) Effects of salt stress on the morphology, growth and physiological parameters of Juglans microcarpa L. seedlings. Plants (Basel). https:\/\/doi.org\/10.3390\/plants11182381","journal-title":"Plants (Basel)"},{"key":"13956_CR20","doi-asserted-by":"publisher","DOI":"10.3389\/fchem.2020.00369","volume":"8","author":"R Kalra","year":"2020","unstructured":"Kalra R, Conlan XA, Goel M (2020) Fungi as a potential source of pigments: harnessing filamentous fungi. Front Chem 8:369. https:\/\/doi.org\/10.3389\/fchem.2020.00369","journal-title":"Front Chem"},{"key":"13956_CR21","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1093\/nar\/28.1.27","volume":"28","author":"M Kanehisa","year":"2000","unstructured":"Kanehisa M, Goto S (2000) KEGG: Kyoto encyclopedia of genes and genomes. Nucleic Acids Res 28:27\u201330","journal-title":"Nucleic Acids Res"},{"issue":"4","key":"13956_CR22","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1038\/nmeth.3317","volume":"12","author":"D Kim","year":"2015","unstructured":"Kim D, Langmead B, Salzberg SL (2015) HISAT: a fast spliced aligner with low memory requirements. Nat Methods 12(4):357\u2013360. https:\/\/doi.org\/10.1038\/nmeth.3317","journal-title":"Nat Methods"},{"issue":"4","key":"13956_CR23","doi-asserted-by":"publisher","first-page":"821","DOI":"10.1016\/j.bbrc.2016.11.119","volume":"482","author":"Y-J Kim","year":"2017","unstructured":"Kim Y-J, Ryu H-M, Choi J-Y, Cho J-H, Kim C-D, Park S-H, Kim Y-L (2017) Hypoxanthine causes endothelial dysfunction through oxidative stress-induced apoptosis. Biochem Biophys Res Commun 482(4):821\u2013827. https:\/\/doi.org\/10.1016\/j.bbrc.2016.11.119","journal-title":"Biochem Biophys Res Commun"},{"issue":"9","key":"13956_CR24","doi-asserted-by":"publisher","first-page":"597","DOI":"10.3390\/jof10090597","volume":"10","author":"AJ Lee","year":"2024","unstructured":"Lee AJ, Hammond J, Sheridan J, Swift S, Villas-Boas SG (2024) Antifungal activity of disalt of epipyrone a from Epicoccum nigrum likely via disrupted fatty acid elongation and sphingolipid biosynthesis. J Fungi 10(9):597. https:\/\/doi.org\/10.3390\/jof10090597","journal-title":"J Fungi"},{"issue":"7","key":"13956_CR25","doi-asserted-by":"publisher","first-page":"1166","DOI":"10.3390\/molecules22071166","volume":"22","author":"C Li","year":"2017","unstructured":"Li C, Sarotti AM, Yang B, Turkson J, Cao S (2017) A new N-methoxypyridone from the co-cultivation of hawaiian endophytic fungi Camporesia sambuci FT1061 and Epicoccum sorghinum FT1062. Molecules 22(7):1166. https:\/\/doi.org\/10.3390\/molecules22071166","journal-title":"Molecules"},{"issue":"23","key":"13956_CR26","doi-asserted-by":"publisher","first-page":"3847","DOI":"10.1039\/C9QO01087D","volume":"6","author":"H-T Li","year":"2019","unstructured":"Li H-T, Tang L, Liu T, Yang R, Yang Y, Zhou H, Ding Z-T (2019) Polyoxygenated meroterpenoids and a bioactive illudalane derivative from a co-culture of Armillaria sp. and Epicoccum sp. Org Chem Front 6(23):3847\u20133853","journal-title":"Org Chem Front"},{"issue":"7","key":"13956_CR27","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1093\/bioinformatics\/btt656","volume":"30","author":"Y Liao","year":"2014","unstructured":"Liao Y, Smyth GK, Shi W (2014) FeatureCounts: an efficient general purpose program for assigning sequence reads to genomic features. Bioinformatics 30(7):923\u2013930. https:\/\/doi.org\/10.1093\/bioinformatics\/btt656","journal-title":"Bioinformatics"},{"issue":"14","key":"13956_CR28","doi-asserted-by":"publisher","DOI":"10.1016\/j.bmcl.2020.127242","volume":"30","author":"YJ Lima","year":"2020","unstructured":"Lima YJ, Choia E, Parkb S-H, Kwon H-J (2020) Genetic localization of the orevactaene\/epipyrone biosynthetic gene cluster in Epicoccum nigrum. Bioorg Med Chem Lett 30(14):127242. https:\/\/doi.org\/10.1016\/j.bmcl.2020.127242","journal-title":"Bioorg Med Chem Lett"},{"issue":"5","key":"13956_CR29","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1007\/s00449-011-0654-4","volume":"35","author":"X-J Liu","year":"2012","unstructured":"Liu X-J, Liu R-S, Li H-M, Tang Y-J (2012) Lycopene production from synthetic medium by Blakeslea trispora NRRL 2895 (+) and 2896 (-) in a stirred-tank fermenter. Bioprocess Biosyst Eng 35(5):739\u2013749. https:\/\/doi.org\/10.1007\/s00449-011-0654-4","journal-title":"Bioprocess Biosyst Eng"},{"issue":"1","key":"13956_CR30","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1128\/JB.181.1.305-318.1999","volume":"181","author":"N Lomovskaya","year":"1999","unstructured":"Lomovskaya N, Otten SL, Doi-Katayama Y, Fonstein L, Liu X, Takatsu T, Inventi-Solari A, Filippini S, Torti F, Colombo AL, Hutchinson CR (1999) Doxorubicin overproduction in Streptomyces peucetius: cloning and characterization of the dnru ketoreductase and dnrv genes and the doxa cytochrome p-450 hydroxylase gene. J Bacteriol 181(1):305\u2013318","journal-title":"J Bacteriol"},{"issue":"4","key":"13956_CR31","doi-asserted-by":"publisher","first-page":"842","DOI":"10.1016\/j.synbio.2024.07.002","volume":"9","author":"H Long","year":"2024","unstructured":"Long H, Zhou J, Ren Y, Lu J, Wang N, Liu H, Zhou X, Cai M (2024) Comparative omics directed gene discovery and rewiring for normal temperature-adaptive red pigment synthesis by polar psychrotrophic fungus Geomyces sp. WNF-15A. Synth Syst Biotechnol 9(4):842\u2013852. https:\/\/doi.org\/10.1016\/j.synbio.2024.07.002","journal-title":"Synth Syst Biotechnol"},{"key":"13956_CR32","doi-asserted-by":"publisher","unstructured":"Longobardi C, Lotito D, Staropoli A, Iervolino V, Cacciola NA, Montagnaro S, Vinale F, Damiano S, Ciarcia R (2026) Natural product epipyrone A from Epicoccum nigrum exhibits antiproliferative activity on canine mammary tumor cells through PI3K\/Akt\/mTOR pathway modulation. Antioxidants-Basel 15(2):173 ARTN 173. https:\/\/doi.org\/10.3390\/antiox15020173","DOI":"10.3390\/antiox15020173"},{"key":"13956_CR33","doi-asserted-by":"publisher","unstructured":"Ma JW, Dai YF, Shen X, Zhong M, Gong Y, Chen HY, Wang WJ, Liang XL (2026) Farnesol modulated growth and pigmentation in Talaromyces sp. LRS407 via ROS-mediated UPR and RQC signaling pathways. Food Microbiol 139 ARTN 105095. https:\/\/doi.org\/10.1016\/j.fm.2026.105095","DOI":"10.1016\/j.fm.2026.105095"},{"issue":"5952","key":"13956_CR34","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1126\/science.1175602","volume":"326","author":"SM Ma","year":"2009","unstructured":"Ma SM, Li JW-H, Choi JW, Zhou H, Lee KKM, Moorthie VA, Xie X, Kealey JT, Silva NAD, Vederas JC, Tang Y (2009) Complete reconstitution of a highly reducing iterative polyketide synthase. Science 326(5952):589\u2013592. https:\/\/doi.org\/10.1126\/science.1175602","journal-title":"Science"},{"issue":"1","key":"13956_CR35","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1186\/1475-2859-8-24","volume":"8","author":"SA Mapari","year":"2009","unstructured":"Mapari SA, Meyer AS, Thrane U, Frisvad JC (2009) Identification of potentially safe promising fungal cell factories for the production of polyketide natural food colorants using chemotaxonomic rationale. Microb Cell Fact 8(1):24. https:\/\/doi.org\/10.1186\/1475-2859-8-24","journal-title":"Microb Cell Fact"},{"issue":"1","key":"13956_CR36","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1007\/s10565-006-0143-8","volume":"23","author":"P Menna","year":"2006","unstructured":"Menna P, Minotti G, Salvatorelli E (2006) In vitro modeling of the structure\u2013activity determinants of anthracycline cardiotoxicity. Cell Biol Toxicol 23(1):49\u201362. https:\/\/doi.org\/10.1007\/s10565-006-0143-8","journal-title":"Cell Biol Toxicol"},{"key":"13956_CR37","doi-asserted-by":"publisher","DOI":"10.1016\/j.bcab.2022.102501","volume":"45","author":"HD Moghadam","year":"2022","unstructured":"Moghadam HD, Yazdi FT, Shahidi F, Sarabi-Jamab M, Es\u2019haghi Z (2022) Co-culture of Monascuspurpureus with Saccharomyces cerevisiae: a strategy for pigments increment and citrinin reduction. Biocatal Agric Biotechnol 45:102501","journal-title":"Biocatal Agric Biotechnol"},{"key":"13956_CR38","doi-asserted-by":"publisher","unstructured":"Mohamed H, Awad MF, Shah AM, Sadaqat B, Nazir Y, Naz T, Yang W, Song YD (2022) Coculturing of Mucor plumbeus and Bacillus subtilis bacterium as an efficient fermentation strategy to enhance fungal lipid and gamma-linolenic acid (GLA) production. Sci Rep-Uk 12(1):13111 ARTN 13111. https:\/\/doi.org\/10.1038\/s41598-022-17442-2","DOI":"10.1038\/s41598-022-17442-2"},{"key":"13956_CR39","doi-asserted-by":"publisher","unstructured":"Naz T, Saeed MT, Umer Z, Safdar W, Mohamed H, Song YD, Zhao XY (2026) Yarrowia lipolytica as a potential chassis for value-added products using a co-culture system and lignocellulose biomass-derived xylose. Front Microbiol 16:1704596 ARTN 1704596. https:\/\/doi.org\/10.3389\/fmicb.2025.1704596","DOI":"10.3389\/fmicb.2025.1704596"},{"key":"13956_CR40","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodres.2024.114780","volume":"192","author":"O Nikoloudaki","year":"2024","unstructured":"Nikoloudaki O, Aheto F, Cagno RD, Gobbetti M (2024) Synthetic microbial communities: a gateway to understanding resistance, resilience, and functionality in spontaneously fermented food microbiomes. Food Res Int 192:114780. https:\/\/doi.org\/10.1016\/j.foodres.2024.114780","journal-title":"Food Res Int"},{"key":"13956_CR41","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodchem.2021.131214","volume":"372","author":"FD Oliveira","year":"2022","unstructured":"Oliveira FD, Rocha ILD, Pinto DCGA, Ventura SPM, Santos AGD, Crevelin EJ, Ebinuma VDCS (2022) Identification of azaphilone derivatives of Monascus colorants from Talaromyces amestolkiae and their halochromic properties. Food Chem 372:131214. https:\/\/doi.org\/10.1016\/j.foodchem.2021.131214","journal-title":"Food Chem"},{"issue":"2","key":"13956_CR42","doi-asserted-by":"publisher","DOI":"10.3390\/md18020073","volume":"18","author":"E Oppong-Danquah","year":"2020","unstructured":"Oppong-Danquah E, Budnicka P, Bl\u00fcmel M, Tasdemir D (2020) Design of fungal co-cultivation based on comparative metabolomics and bioactivity for discovery of marine fungal agrochemicals. Mar Drugs 18(2):73. https:\/\/doi.org\/10.3390\/md18020073","journal-title":"Mar Drugs"},{"issue":"16","key":"13956_CR43","doi-asserted-by":"publisher","first-page":"5121","DOI":"10.1128\/aem.00478-08","volume":"74","author":"CD Reeves","year":"2008","unstructured":"Reeves CD, Hu Z, Reid R, Kealey JT (2008) Genes for the biosynthesis of the fungal polyketides hypothemycin from Hypomyces subiculosus and radicicol from Pochonia chlamydosporia. Appl Environ Microbiol 74(16):5121\u20135129. https:\/\/doi.org\/10.1128\/aem.00478-08","journal-title":"Appl Environ Microbiol"},{"key":"13956_CR44","doi-asserted-by":"publisher","DOI":"10.1038\/s41525-017-0042-3","volume":"3","author":"J Reiner","year":"2018","unstructured":"Reiner J, Pisani L, Qiao W, Singh R, Yang Y, Shi L, Khan WA, Sebra R, Cohen N, Babu A, Edelmann L, Jabs EW, Scott SA (2018) Cytogenomic identification and long-read single molecule real-time (SMRT) sequencing of a Bardet\u2013Biedl Syndrome 9 (BBS9) deletion. NPJ Genom Med 3:3. https:\/\/doi.org\/10.1038\/s41525-017-0042-3","journal-title":"NPJ Genom Med"},{"issue":"3","key":"13956_CR45","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1038\/cdd.2014.219","volume":"22","author":"P Rockenfeller","year":"2015","unstructured":"Rockenfeller P, Koska M, Pietrocola F, Minois N, Knittelfelder O, Sica V, Franz J, Carmona-Gutierrez D, Kroemer G, Madeo F (2015) Phosphatidylethanolamine positively regulates autophagy and longevity. Cell Death Differ 22(3):499\u2013508. https:\/\/doi.org\/10.1038\/cdd.2014.219","journal-title":"Cell Death Differ"},{"issue":"11","key":"13956_CR46","first-page":"2673","volume":"47","author":"K Sakata","year":"2014","unstructured":"Sakata K, Kuwatsuka T, Sakurai A, Takahashi N, Tamura G (2014) Isolation of aspirochlorine (=antibiotic a30641) as a true antimicrobial constituent of the antibiotic, oryzachlorin, from Aspergillus oryzae. Agric Biol Chem 47(11):2673\u20132674","journal-title":"Agric Biol Chem"},{"key":"13956_CR47","unstructured":"Sardaryan E (2002) Strain of the microorganism penicillium oxalicum var. armeniaca and its application. U.S. Patent No. US6340586. United States Patent and Trademark Office"},{"issue":"4","key":"13956_CR48","doi-asserted-by":"publisher","first-page":"1167","DOI":"10.1021\/acs.jnatprod.7b00144","volume":"80","author":"Z Shang","year":"2017","unstructured":"Shang Z, Salim AA, Capon RJ (2017) Chaunopyran A: co-cultivation of marine mollusk-derived fungi activates a rare class of 2-alkenyl-tetrahydropyran. J Nat Prod 80(4):1167\u20131172. https:\/\/doi.org\/10.1021\/acs.jnatprod.7b00144","journal-title":"J Nat Prod"},{"issue":"11","key":"13956_CR49","doi-asserted-by":"publisher","first-page":"2498","DOI":"10.1101\/gr.1239303","volume":"13","author":"P Shannon","year":"2003","unstructured":"Shannon P, Markiel A, Ozier O, Baliga NS, Wang JT, Ramage D, Amin N, Schwikowski B, Ideker T (2003) Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res 13(11):2498\u20132504","journal-title":"Genome Res"},{"key":"13956_CR50","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodchem.2022.134422","volume":"403","author":"N Shen","year":"2023","unstructured":"Shen N, Ren J, Liu Y, Sun W, Li Y, Xin H, Cui Y (2023) Natural edible pigments: a comprehensive review of resource, chemical classification, biosynthesis pathway, separated methods and application. Food Chem 403:134422. https:\/\/doi.org\/10.1016\/j.foodchem.2022.134422","journal-title":"Food Chem"},{"issue":"30","key":"13956_CR51","doi-asserted-by":"publisher","first-page":"16825","DOI":"10.1021\/acs.jafc.4c04764","volume":"72","author":"X Shen","year":"2024","unstructured":"Shen X, Lei C, Zhang A, Wang L, Chen D, Qi L, Hu Y, Chen G, Ran H, Yin W-B (2024) Stimulating novel and bioactive metabolite production by cocultivation of two fungi\u2014Aspergillus oryzae and Epicoccum dendrobii. J Agric Food Chem 72(30):16825\u201316834. https:\/\/doi.org\/10.1021\/acs.jafc.4c04764","journal-title":"J Agric Food Chem"},{"issue":"2","key":"13956_CR52","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1007\/s00203-023-03414-1","volume":"205","author":"TF Sousa","year":"2023","unstructured":"Sousa TF, J\u00fanior MBdA, Peres EG, Souza MP, Silva FMAd, Medeiros LSd, Souza ADLd, Souza AQLd, Yamagishi MEB, Silva GFd, Koolen HHF, Queiroz MVD (2023) Discovery of dual PKS involved in sclerotiorin biosynthesis in Penicilliummeliponae using genome mining and gene knockout. Arch Microbiol 205(2):75. https:\/\/doi.org\/10.1007\/s00203-023-03414-1","journal-title":"Arch Microbiol"},{"issue":"5","key":"13956_CR53","doi-asserted-by":"publisher","first-page":"2765","DOI":"10.26599\/fshw.2022.9250224","volume":"13","author":"S Tong","year":"2024","unstructured":"Tong S, Li W, Guo W, L\u00fc X, Sund J, Rao Y, Xiao Y, Yan Y, Ai L, Ni L (2024) Microbiomics and metabolomics insights into the microbial regulation on the formation of flavor components in the traditional fermentation process of Chinese Hongqu aged vinegar. Food Sci Hum Wellness 13(5):2765\u20132778. https:\/\/doi.org\/10.26599\/fshw.2022.9250224","journal-title":"Food Sci Hum Wellness"},{"issue":"3","key":"13956_CR54","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1016\/j.tim.2007.01.005","volume":"15","author":"ID Tsitsigiannis","year":"2007","unstructured":"Tsitsigiannis ID, Keller NP (2007) Oxylipins as developmental and host\u2013fungal communication signals. Trends Microbiol 15(3):109\u2013118. https:\/\/doi.org\/10.1016\/j.tim.2007.01.005","journal-title":"Trends Microbiol"},{"key":"13956_CR55","doi-asserted-by":"publisher","unstructured":"Wang QY, Yang W, Pu SC, Li CY, Wang HL, Mu DD, Wu XF, Zheng Z, Jiang ST, Li XJ (2018) Physicochemical properties of Sufu produced by single and mixed fermentation with Mucorracemosus and Rhizopus oryzae. Curr Top Nutr Res 18(1):63\u201369. https:\/\/doi.org\/10.37290\/ctnr2641-452x.18:63-69","DOI":"10.37290\/ctnr2641-452x.18:63-69"},{"issue":"6","key":"13956_CR56","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1071\/CH21033","volume":"74","author":"MT Welling","year":"2021","unstructured":"Welling MT, Deseo MA, Bacic A, Doblin MS (2021) Untargeted metabolomic analyses reveal chemical complexity of dioecious cannabis flowers. Aust J Chem 74(6):463\u2013479","journal-title":"Aust J Chem"},{"issue":"9","key":"13956_CR57","doi-asserted-by":"publisher","first-page":"618","DOI":"10.1038\/nrmicro2852","volume":"10","author":"AJ Westermann","year":"2012","unstructured":"Westermann AJ, Gorski SA, Vogel J (2012) Dual RNA-seq of pathogen and host. Nat Rev Microbiol 10(9):618\u2013630. https:\/\/doi.org\/10.1038\/nrmicro2852","journal-title":"Nat Rev Microbiol"},{"issue":"15","key":"13956_CR58","doi-asserted-by":"publisher","first-page":"2619","DOI":"10.1080\/14786419.2023.2195176","volume":"38","author":"AK Whaley","year":"2023","unstructured":"Whaley AK, Ponkratova AO, Orlova AA, Volobuev SV, Shakhova NV, Serebryakov EB, Smirnov SN, Pustovit NV, Kraeva LA, Luzhanin VG (2023) New benzoquinone pigments from the hydnoid fungus Sarcodontia setosa and their biosynthetic relationship. Nat Prod Res 38(15):2619\u20132628","journal-title":"Nat Prod Res"},{"issue":"21","key":"13956_CR59","doi-asserted-by":"publisher","first-page":"3027","DOI":"10.1002\/cbic.202100240","volume":"22","author":"K Williams","year":"2021","unstructured":"Williams K, Greco C, Bailey AM, Willis CL (2021) Core steps to the azaphilone family of fungal natural products. ChemBioChem 22(21):3027\u20133036. https:\/\/doi.org\/10.1002\/cbic.202100240","journal-title":"ChemBioChem"},{"issue":"12","key":"13956_CR60","doi-asserted-by":"publisher","first-page":"5577","DOI":"10.1002\/jsfa.9831","volume":"99","author":"W Xiang","year":"2019","unstructured":"Xiang W, Xu Q, Zhang N, Rao Y, Zhu L, Zhang Q (2019) Mucor indicus and Rhizopus oryzae co-culture to improve the flavor of Chinese turbid rice wine. J Sci Food Agric 99(12):5577\u20135585","journal-title":"J Sci Food Agric"},{"issue":"2","key":"13956_CR61","doi-asserted-by":"publisher","first-page":"464","DOI":"10.3390\/microorganisms11020464","volume":"11","author":"S Xu","year":"2023","unstructured":"Xu S, Li M, Hu Z, Shao Y, Ying J, Zhang H (2023) The potential use of fungal co-culture strategy for discovery of new secondary metabolites. Microorganisms 11(2):464. https:\/\/doi.org\/10.3390\/microorganisms11020464","journal-title":"Microorganisms"},{"issue":"8","key":"13956_CR62","doi-asserted-by":"publisher","first-page":"1049","DOI":"10.1016\/j.chembiol.2012.07.004","volume":"19","author":"AO Zabala","year":"2012","unstructured":"Zabala AO, Xu W, Chooi Y-H, Tang Y (2012) Characterization of a silent azaphilone gene cluster from Aspergillus niger ATCC 1015 reveals a hydroxylation-mediated pyran-ring formation. Chem Biol 19(8):1049\u20131059. https:\/\/doi.org\/10.1016\/j.chembiol.2012.07.004","journal-title":"Chem Biol"},{"key":"13956_CR63","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodchem.2022.134702","volume":"404","author":"L Zhang","year":"2023","unstructured":"Zhang L, Qin Z, Zhang L, Jiang Y, Zhu J (2023) Dynamic changes of quality and flavor characterization of Zhejiang rosy vinegar during fermentation and aging based on untargeted metabolomics. Food Chem 404:134702. https:\/\/doi.org\/10.1016\/j.foodchem.2022.134702","journal-title":"Food Chem"},{"issue":"13","key":"13956_CR64","doi-asserted-by":"publisher","first-page":"4530","DOI":"10.1021\/ja100060k","volume":"132","author":"H Zhou","year":"2010","unstructured":"Zhou H, Qiao K, Gao Z, Meehan MJ, Li JW-H, Zhao X, Dorrestein PC, Vederas JC, Tang Y (2010) Enzymatic synthesis of resorcylic acid lactones by cooperation of fungal iterative polyketide synthases involved in hypothemycin biosynthesis. J Am Chem Soc 132(13):4530\u20134531. https:\/\/doi.org\/10.1021\/ja100060k","journal-title":"J Am Chem Soc"},{"issue":"1","key":"13956_CR65","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1080\/19476337.2023.2207613","volume":"21","author":"M Zhou","year":"2023","unstructured":"Zhou M, Chen Y, Fang X, Wu L, Zhang Y (2023) Isolation and identification of pigment-producing filamentous fungus DBFL05 and its pigment characteristics and chemical structure. CyTA - J Food 21(1):374\u2013385. https:\/\/doi.org\/10.1080\/19476337.2023.2207613","journal-title":"CyTA - J Food"}],"container-title":["Applied Microbiology and Biotechnology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00253-026-13956-1","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00253-026-13956-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00253-026-13956-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,19]],"date-time":"2026-10-19T23:11:26Z","timestamp":1792451486000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00253-026-13956-1"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,23]]},"references-count":65,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["13956"],"URL":"https:\/\/doi.org\/10.1007\/s00253-026-13956-1","relation":{},"ISSN":["1432-0614"],"issn-type":[{"value":"1432-0614","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,23]]},"assertion":[{"value":"16 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 June 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 July 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 July 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethical approval","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"311"}}