{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T04:55:50Z","timestamp":1790916950495,"version":"4.1.0"},"reference-count":45,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T00:00:00Z","timestamp":1790899200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:p>\n                    Bulk fermentation of cigar tobacco leaves couples microbial succession and chemical transformations; however, it remains unclear whether commercially distinct filler and wrapper leaves follow different ecological and biochemical trajectories under identical industrial conditions. Using 16S rRNA and ITS amplicon sequencing, together with conventional chemical indices and volatile profiling, we tracked filler and wrapper leaves (two commercial cultivars, variety 204 and variety 108) through 0, 10, 18, 49, and 89\u202fdays of industrial bulk fermentation. The results revealed that reducing sugars declined by 63% in filler leaves (0.57 to 0.21%) and 66% in wrapper leaves (0.59 to 0.20%), whereas alkaloid dynamics differed between materials: filler leaves showed a transient mid-fermentation rise before decline, while wrapper leaves declined from the outset. Bacterial communities shifted from Proteobacteria-dominated early assemblages toward increased Firmicutes during later fermentation, with\n                    <jats:italic>Acinetobacter, Staphylococcus<\/jats:italic>\n                    and\n                    <jats:italic>Pseudomonas<\/jats:italic>\n                    as major genera. Fungal communities were dominated primarily by\n                    <jats:italic>Ascomycota<\/jats:italic>\n                    , with\n                    <jats:italic>Aspergillus<\/jats:italic>\n                    prevalent across fermentation and\n                    <jats:italic>Papiliotrema<\/jats:italic>\n                    becoming more prevalent at later stages. Bray-Curtis based ordination showed structured, directional succession in both bacterial and fungal communities, and fermentation time explained substantially more community variation than material type (bacteria: R\n                    <jats:sup>2<\/jats:sup>\n                    \u202f=\u202f0.57 vs. 0.04; fungi: R\n                    <jats:sup>2<\/jats:sup>\n                    \u202f=\u202f0.42 vs. 0.08). Correlation, redundancy and network analyses linked both bacterial and fungal community variation to sugars, nitrogenous compounds, alkaloids, and volatile classes, although these associations remain correlative rather than causal. By directly comparing filler and wrapper leaves under identical industrial fermentation environment, this study highlights that commercially distinct cigar materials follow divergent ecological and biochemical trajectories, providing a process-resolved basis for fermentation monitoring and material-specific quality management.\n                  <\/jats:p>","DOI":"10.3389\/fmicb.2026.1922631","type":"journal-article","created":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T04:39:59Z","timestamp":1790915999000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Comparative analysis of microbial succession and biochemical changes in cigar tobacco filler and wrapper leaves during fermentation"],"prefix":"10.3389","volume":"17","author":[{"given":"Zhaoliang","family":"Geng","sequence":"first","affiliation":[{"name":"Hainan Provincial Branch of China National Tobacco Corporation","place":["Haikou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bin","family":"Cai","sequence":"additional","affiliation":[{"name":"Hainan Provincial Branch of China National Tobacco Corporation","place":["Haikou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanqi","family":"Hu","sequence":"additional","affiliation":[{"name":"China Tobacco Shandong Industrial Co., Ltd.","place":["Jinan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Beisen","family":"Lin","sequence":"additional","affiliation":[{"name":"Hainan Provincial Branch of China National Tobacco Corporation","place":["Haikou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tongjing","family":"Yan","sequence":"additional","affiliation":[{"name":"Hainan Provincial Branch of China National Tobacco Corporation","place":["Haikou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huajun","family":"Gao","sequence":"additional","affiliation":[{"name":"Hainan Provincial Branch of China National Tobacco Corporation","place":["Haikou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abdullah","family":"Khan","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tobacco Biology and Processing, Ministry of Agriculture and Rural Affairs, Tobacco Research Institute of Chinese Academy of Agricultural Sciences","place":["Qingdao, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinghua","family":"Ma","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tobacco Biology and Processing, Ministry of Agriculture and Rural Affairs, Tobacco Research Institute of Chinese Academy of Agricultural Sciences","place":["Qingdao, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,10,2]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"927","DOI":"10.3390\/molecules25040927","article-title":"Impact of fermentation on the phenolic compounds and antioxidant activity of whole cereal grains: a mini review","volume":"25","author":"Adebo","year":"2020","journal-title":"Molecules"},{"key":"ref2","doi-asserted-by":"publisher","first-page":"554051","DOI":"10.3389\/fmicb.2020.554051","article-title":"Fungal composition and diversity of the tobacco leaf phyllosphere during curing of leaves","volume":"11","author":"Chen","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref3","doi-asserted-by":"publisher","DOI":"10.1101\/081257","article-title":"UNOISE2: improved error-correction for Illumina 16S and ITS amplicon sequencing","author":"Edgar","year":"2016","journal-title":"bioRxiv"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1002\/9780470122556.ch8","article-title":"Chemical changes in the harvested tobacco leaf: part II. Chemical and enzymic conversions during fermentation and aging","volume":"10","author":"Frankenburg","year":"1950","journal-title":"Adv. Enzymol. Relat. Areas Mol. Biol."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"25556","DOI":"10.1038\/s41598-025-07310-0","article-title":"Characterization of aroma profiles and microbial communities of cigar tobacco leaves from different varieties and origins and their correlations analysis","volume":"15","author":"Geng","year":"2025","journal-title":"Sci. Rep."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"1415","DOI":"10.3390\/agronomy11071415","article-title":"The significance of microbial transformation of nitrogen compounds in the light of integrated crop management","volume":"11","author":"Grzyb","year":"2021","journal-title":"Agronomy"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"103190","DOI":"10.1016\/j.ejsobi.2020.103190","article-title":"Litter composition has stronger influence on the structure of soil fungal than bacterial communities","volume":"98","author":"Habtewold","year":"2020","journal-title":"Eur. J. Soil Biol."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"e68122","DOI":"10.1590\/fst.68122","article-title":"Exogenous additives facilitate the fermentation of cigar tobacco leaves: improving sensory quality and contents of aroma components","volume":"42","author":"Hu","year":"2022","journal-title":"Food Sci. Technol."},{"key":"ref9","author":"Huang","year":"2021"},{"key":"ref10","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1007\/s00203-022-03347-1","article-title":"Effects of Bacillus subtilis subsp. on the microbial community and aroma components of flue-cured tobacco leaves based on metagenome analysis","volume":"204","author":"Huang","year":"2022","journal-title":"Arch. Microbiol."},{"key":"ref11","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1007\/s00253-010-2763-4","article-title":"Bacterial diversities on unaged and aging flue-cured tobacco leaves estimated by 16S rRNA sequence analysis","volume":"88","author":"Huang","year":"2010","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref12","doi-asserted-by":"publisher","first-page":"1550383","DOI":"10.3389\/fbioe.2025.1550383","article-title":"Effects of different fermentation temperatures on microbiomes of cigar tobacco leaves","volume":"13","author":"Jia","year":"2025","journal-title":"Front. Bioeng. Biotechnol."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"103189","DOI":"10.1016\/j.fochx.2025.103189","article-title":"Untargeted metabolomics reveals phenolic compound dynamics during mung bean fermentation","volume":"31","author":"Lang","year":"2025","journal-title":"Food Chem"},{"key":"ref14","doi-asserted-by":"publisher","first-page":"e0226066","DOI":"10.1371\/journal.pone.0226066","article-title":"Classification of flavors in cigarillos and little cigars and their variable cellular and acellular oxidative and cytotoxic responses","volume":"14","author":"Lawyer","year":"2019","journal-title":"PLoS One"},{"key":"ref15","doi-asserted-by":"publisher","first-page":"1438566","DOI":"10.3389\/fmicb.2024.1438566","article-title":"The dynamics of microbial community structure and metabolic function in different parts of cigar tobacco leaves during air-curing","volume":"15","author":"Li","year":"2024","journal-title":"Front. Microbiol."},{"key":"ref16","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1186\/s12866-020-02035-8","article-title":"Influence of microbiota and metabolites on the quality of tobacco during fermentation","volume":"20","author":"Li","year":"2020","journal-title":"BMC Microbiol."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"6","DOI":"10.1186\/s40793-024-00548-7","article-title":"Bacterial wilt affects the structure and assembly of microbial communities along the soil-root continuum","volume":"19","author":"Liang","year":"2024","journal-title":"Environ. Microbiome"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"1025881","DOI":"10.3389\/fmicb.2022.1025881","article-title":"Phyllosphere microbial community of cigar tobacco and its corresponding metabolites","volume":"13","author":"Liu","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref19","first-page":"15","article-title":"Research progress on tobacco leaf fermentation","volume":"27","author":"Liu","year":"2011","journal-title":"Guangxi J Light Industry"},{"key":"ref20","doi-asserted-by":"publisher","first-page":"e1171","DOI":"10.1002\/mbo3.1171","article-title":"Microbial community and metabolic function analysis of cigar tobacco leaves during fermentation","volume":"10","author":"Liu","year":"2021","journal-title":"Microbiol. Open"},{"key":"ref21","doi-asserted-by":"publisher","first-page":"125895","DOI":"10.1016\/j.foodchem.2019.125895","article-title":"Untargeted and targeted metabolomics reveal the chemical characteristic of pu-erh tea (Camellia assamica) during pile-fermentation","volume":"311","author":"Long","year":"2020","journal-title":"Food Chem."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1146\/annurev-micro-032521-041956","article-title":"Microbiome assembly in fermented foods","volume":"77","author":"Louw","year":"2023","journal-title":"Ann. Rev. Microbiol."},{"key":"ref23","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1016\/j.copbio.2016.11.010","article-title":"Health benefits of fermented foods: microbiota and beyond","volume":"44","author":"Marco","year":"2017","journal-title":"Curr. Opin. Biotechnol."},{"key":"ref24","doi-asserted-by":"publisher","first-page":"308","DOI":"10.1186\/s13104-024-06951-0","article-title":"Sequence-matching adapter trimmers generate consistent quality and assembly metrics for Illumina sequencing of RNA viruses","volume":"17","author":"Nabakooza","year":"2024","journal-title":"BMC. Res. Notes"},{"key":"ref9001","year":"2005"},{"key":"ref25","doi-asserted-by":"publisher","DOI":"10.1007\/s00253-025-13598-9","article-title":"Bacteria dynamics and its correlation with chemical composition changes in tobacco leaves during flue curing","volume":"109","author":"Sha","year":"2025","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref26","doi-asserted-by":"publisher","first-page":"534","DOI":"10.1186\/s12866-024-03702-w","article-title":"Fermentation process of tobacco leaves drives the specific changes of microbial community","volume":"24","author":"Tao","year":"2024","journal-title":"BMC Microbiol."},{"key":"ref27","first-page":"1211","article-title":"Research advances in fermentation methods of cigar tobacco leaves","volume":"45","author":"Wan","year":"2017","journal-title":"J Shanxi Agric. Sci."},{"key":"ref28","doi-asserted-by":"publisher","first-page":"095012","DOI":"10.1088\/2515-7620\/ac9352","article-title":"The diversity, structure and function of microbial communities changes across aging process of tobacco leaves","volume":"4","author":"Wang","year":"2022","journal-title":"Environ. Res. Commun."},{"key":"ref29","doi-asserted-by":"publisher","first-page":"956","DOI":"10.3390\/microorganisms12050956","article-title":"The impact of aboveground Epichlo\u00eb endophytic fungi on the rhizosphere microbial functions of the host Melica transsilvanica","volume":"12","author":"Wang","year":"2024","journal-title":"Microorganisms"},{"key":"ref30","first-page":"1","article-title":"Research progress of air-cured tobacco fermentation","author":"Wang","year":"2014","journal-title":"J. Henan Agric. Sci."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"2815","DOI":"10.1038\/s41598-022-06593-x","article-title":"Soil fungal communities affect the chemical quality of flue-cured tobacco leaves in Bijie, Southwest China","volume":"12","author":"Wang","year":"2022","journal-title":"Sci. Rep."},{"key":"ref32","doi-asserted-by":"publisher","first-page":"1627842","DOI":"10.3389\/fbioe.2025.1627842","article-title":"Microbial community succession during tobacco fermentation reveals a flavor-improving mechanism","volume":"13","author":"Wang","year":"2025","journal-title":"Front. Bioeng. Biotechnol."},{"key":"ref33","doi-asserted-by":"publisher","first-page":"e32","DOI":"10.1002\/imt2.32","article-title":"GgClusterNet: an R package for microbiome network analysis and modularity-based multiple network layouts","volume":"1","author":"Wen","year":"2022","journal-title":"iMeta"},{"key":"ref34","doi-asserted-by":"publisher","first-page":"1264","DOI":"10.1080\/10498850.2021.1988016","article-title":"Isolation and identification of aroma-producing yeast from mackerel fermentation broth and its fermentation characteristics","volume":"30","author":"Wu","year":"2021","journal-title":"J. Aquat. Food Prod. Technol."},{"key":"ref35","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1007\/s00248-004-0171-y","article-title":"Bacterial colonization of the phyllosphere of Mediterranean perennial species as influenced by leaf structural and chemical features","volume":"50","author":"Yadav","year":"2005","journal-title":"Microb. Ecol."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"105","DOI":"10.4236\/jbm.2018.66008","article-title":"Research progress in tobacco fermentation","volume":"6","author":"Yang","year":"2018","journal-title":"J. Biosci. Med."},{"key":"ref37","doi-asserted-by":"publisher","first-page":"D643","DOI":"10.1093\/nar\/gkt1209","article-title":"The SILVA and \u201call-species living tree project (LTP)\u201d taxonomic frameworks","volume":"42","author":"Yilmaz","year":"2014","journal-title":"Nucleic Acids Res."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"1632731","DOI":"10.3389\/fmicb.2025.1632731","article-title":"Application of starch-degradation bacteria in cigar tobacco leaf fermentation: effects on starch degradation, microbial communities and metabolic pathways","volume":"16","author":"Yin","year":"2025","journal-title":"Front. Microbiol."},{"key":"ref39","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1007\/s00253-024-13043-3","article-title":"Analyzing microbial community and volatile compound profiles in the fermentation of cigar tobacco leaves","volume":"108","author":"Zhang","year":"2024","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"e01029-24","DOI":"10.1128\/spectrum.01029-24","article-title":"Integrated microbiology and metabolomics analysis reveal the fermentation process and the flavor development in cigar tobacco leaf","volume":"13","author":"Zhang","year":"2025","journal-title":"Microbiol. Spectrum"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"117889","DOI":"10.1016\/j.indcrop.2023.117889","article-title":"Study on tobacco quality improvement and bacterial community succession during microbial co-fermentation","volume":"208","author":"Zhang","year":"2024","journal-title":"Ind. Crop. Prod."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1186\/s13568-023-01642-8","article-title":"Changes in physicochemical properties and microbial community succession during leaf stacking fermentation","volume":"13","author":"Zhang","year":"2023","journal-title":"AMB Express"},{"key":"ref43","doi-asserted-by":"publisher","first-page":"907270","DOI":"10.3389\/fmicb.2022.907270","article-title":"Analysis of microbial community, volatile flavor compounds, and flavor of cigar tobacco leaves from different regions","volume":"13","author":"Zheng","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref44","doi-asserted-by":"publisher","first-page":"5266","DOI":"10.3390\/molecules29225266","article-title":"Optimisation of the ethanol fermentation process using hydrothermal pretreatment of cellulose waste-effect of fermentation pattern and strain","volume":"29","author":"Zhou","year":"2024","journal-title":"Molecules"}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1922631\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T04:40:06Z","timestamp":1790916006000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1922631\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,2]]},"references-count":45,"alternative-id":["10.3389\/fmicb.2026.1922631"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2026.1922631","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,2]]},"article-number":"1922631"}}