{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,8]],"date-time":"2026-07-08T16:24:42Z","timestamp":1783527882061,"version":"3.55.0"},"reference-count":76,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2026,7,8]],"date-time":"2026-07-08T00:00:00Z","timestamp":1783468800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Foods"],"abstract":"<jats:p>The role of lactic acid bacteria (LAB) in cocoa bean fermentation has long been considered central within the traditional yeast\u2013LAB\u2013acetic acid bacteria (AAB) succession model. However, their functional necessity for successful fermentation remains debated. While yeasts and AAB are consistently associated with key transformations such as pulp degradation, ethanol formation, ethanol oxidation, heat generation, and internal bean modification, increasing evidence indicates that cocoa fermentation can proceed effectively under minimal, transient, or reduced LAB activity. This review re-examines the presumed indispensability of LAB using a function-based framework that distinguishes microbial contributions according to their relationship with fermentation completion and process optimization. By integrating evidence from controlled fermentations, microbial suppression studies, starter-culture research, and recent ecological and multi-omics analyses, this review suggests that LAB are better interpreted, under the conditions reported to date, as context-dependent modulators rather than universal core drivers of fermentation completion. LAB contribute to acid balance, citrate metabolism, mannitol production, microbial interactions, ecological stabilization, and quality modulation; however, available evidence indicates that the principal biochemical transformations defining fermentation completion can occur when yeast and AAB activities remain preserved. These conclusions should be interpreted with caution, as direct experimental evidence remains limited and is largely derived from controlled or small-scale fermentation systems. This function-oriented perspective shifts the focus from taxonomic recurrence to biochemical necessity and provides a rational basis for designing starter cultures centered on pathway coverage, metabolic performance, process consistency, and cocoa quality.<\/jats:p>","DOI":"10.3390\/foods15142415","type":"journal-article","created":{"date-parts":[[2026,7,8]],"date-time":"2026-07-08T15:40:09Z","timestamp":1783525209000},"page":"2415","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Revisiting the Role of Lactic Acid Bacteria in Cacao Fermentation: From Traditional Paradigm to Functional Precision"],"prefix":"10.3390","volume":"15","author":[{"given":"Tania Mar\u00eda","family":"Guzm\u00e1n-Armenteros","sequence":"first","affiliation":[{"name":"Facultad de Ingenier\u00eda Mec\u00e1nica y Ciencias de la Producci\u00f3n, Carrera de Ingenier\u00eda en Alimentos, Escuela Superior Polit\u00e9cnica del Litoral, Campus Gustavo Galindo, km 30.5 V\u00eda Perimetral, Guayaquil 090902, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Armando","family":"Echeverr\u00eda","sequence":"additional","affiliation":[{"name":"Universidad Cat\u00f3lica Santiago de Guayaquil, Subsistema de Posgrados, Guayaquil 090508, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jenny","family":"Ruales","sequence":"additional","affiliation":[{"name":"Department of Food Science and Biotechnology, Escuela Polit\u00e9cnica Nacional, Quito P.O. Box 17-01-2759, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luis","family":"Ramos-Guerrero","sequence":"additional","affiliation":[{"name":"Facultad de Ingenier\u00eda y Ciencias Aplicadas, Carrera de Ingenier\u00eda Agroindustrial, Grupo de Investigaci\u00f3n en Alimentos y Agroindustria (GIAA), Universidad de Las Am\u00e9ricas (UDLA), Quito 170124, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,8]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Korcari, D., Fanton, A., Ricci, G., Rabitti, N.S., Laureati, M., Hogenboom, J., Pellegrino, L., Emide, D., Barbiroli, A., and Fortina, M. (2023). Fine Cocoa Fermentation with Selected Lactic Acid Bacteria: Fermentation Performance and Impact on Chocolate Composition and Sensory Properties. Foods, 12.","DOI":"10.3390\/foods12020340"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.ijfoodmicro.2017.05.008","article-title":"Lactic Acid Bacteria Involved in Cocoa Beans Fermentation from Ivory Coast: Species Diversity and Citrate Lyase Production","volume":"256","author":"Ouattara","year":"2017","journal-title":"Int. J. Food Microbiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1016\/j.fm.2010.10.010","article-title":"Dynamics and Species Diversity of Communities of Lactic Acid Bacteria and Acetic Acid Bacteria during Spontaneous Cocoa Bean Fermentation in Vessels","volume":"28","author":"Lefeber","year":"2011","journal-title":"Food Microbiol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1093\/femsre\/fuaa014","article-title":"Functional Role of Yeasts, Lactic Acid Bacteria, and Acetic Acid Bacteria in Cocoa Fermentation Processes","volume":"44","author":"Leroy","year":"2020","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_5","first-page":"1","article-title":"Analysis of Cocoa Fermentation with the Addition of Yeast and Lactic Acid Bacteria","volume":"2","author":"Analianasari","year":"2023","journal-title":"J. Pengemb. Agroindustri Terap."},{"key":"ref_6","first-page":"825","article-title":"Selection of Lactic Acid Bacteria as Starter Culture for Cocoa Fermentation (Theobroma cacao L.)","volume":"9","author":"Tsaaqifah","year":"2023","journal-title":"J. Penelit. Pendidik. IPA"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2130","DOI":"10.1038\/s41564-025-02077-6","article-title":"A Defined Microbial Community Reproduces Attributes of Fine Flavour Chocolate Fermentation","volume":"10","author":"Gopaulchan","year":"2025","journal-title":"Nat. Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Campos, S.D.M., Mart\u00ednez-Burgos, W., Dos Reis, G., Oc\u00e1n-Torres, D., Costa, G.D.S., Vega, F.R., Badel, B.A., Coronado, L.S., Serra, J.L., and Soccol, C. (2025). The Role of Microbial Dynamics, Sensorial Compounds, and Producing Regions in Cocoa Fermentation. Microbiol. Res., 16.","DOI":"10.3390\/microbiolres16040075"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"5841","DOI":"10.1080\/10408398.2021.2025035","article-title":"Ecological Succession and Functional Characteristics of Lactic Acid Bacteria in Traditional Fermented Foods","volume":"63","author":"Wang","year":"2022","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e24927","DOI":"10.1016\/j.heliyon.2024.e24927","article-title":"Electromagnetic Fields Effects on Microbial Growth in Cocoa Fermentation: A Controlled Experimental Approach Using Established Growth Models","volume":"10","author":"Guerra","year":"2024","journal-title":"Heliyon"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"109478","DOI":"10.1016\/j.foodres.2020.109478","article-title":"Exploring the Contribution of Fructophilic Lactic Acid Bacteria to Cocoa Beans Fermentation: Isolation, Selection and Evaluation","volume":"136","author":"Viesser","year":"2020","journal-title":"Food Res. Int."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"3401","DOI":"10.17576\/jsm-2022-5110-23","article-title":"Isolation of Lactic Acid Bacteria from Cocoa Bean Fermentation as Potential Antibacterial Agent against ESKAPE Pathogens","volume":"51","author":"Zin","year":"2022","journal-title":"Sains Malays."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/1750-3841.17612","article-title":"Screening and Characterizing Indigenous Yeasts, Lactic Acid Bacteria, and Acetic Acid Bacteria from Cocoa Fermentation in Hainan for Aroma Development","volume":"90","author":"Chang","year":"2025","journal-title":"J. Food Sci."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"103767","DOI":"10.1016\/j.fm.2021.103767","article-title":"Mapping the Functional and Strain Diversity of the Main Microbiota Involved in Cocoa Fermentation from Cote d\u2019Ivoire","volume":"98","author":"Ouattara","year":"2021","journal-title":"Food Microbiol."},{"key":"ref_15","first-page":"236","article-title":"Isolation and Identification of Microbial Species Found in Cocoa Fermentation as Microbial Starter Culture Candidates for Cocoa Bean Fermentation in Colombia","volume":"36","author":"Tovar","year":"2020","journal-title":"Pelita Perkeb. (A Coffee Cocoa Res. J.)"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"012061","DOI":"10.1088\/1755-1315\/439\/1\/012061","article-title":"Identification of Lactic Acid Bacteria and Acetic Acid Bacteria of Lampung Cocoa Beans Fermentation","volume":"439","author":"Fahrurrozi","year":"2020","journal-title":"IOP Conf. Ser. Earth Environ. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1007\/s00284-008-9097-9","article-title":"Diversity of Predominant Lactic Acid Bacteria Associated with Cocoa Fermentation in Nigeria","volume":"56","author":"Kostinek","year":"2008","journal-title":"Curr. Microbiol."},{"key":"ref_18","first-page":"21","article-title":"Sucesi\u00f3n Microbiana Durante La Fermentaci\u00f3n Espont\u00e1nea de Cacao En Unidades Productivas","volume":"12","year":"2021","journal-title":"Cienc. En. Desarro."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.ijfoodmicro.2015.03.031","article-title":"The Effect of Lactic Acid Bacteria on Cocoa Bean Fermentation","volume":"205","author":"Ho","year":"2015","journal-title":"Int. J. Food Microbiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.ijfoodmicro.2018.04.040","article-title":"Unravelling the Contribution of Lactic Acid Bacteria and Acetic Acid Bacteria to Cocoa Fermentation Using Inoculated Organisms","volume":"279","author":"Ho","year":"2018","journal-title":"Int. J. Food Microbiol."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Verce, M., Schoonejans, J., Aguirre, C.H., Molina-Bravo, R., De Vuyst, L., and Weckx, S. (2021). A Combined Metagenomics and Metatranscriptomics Approach to Unravel Costa Rican Cocoa Box Fermentation Processes Reveals Yet Unreported Microbial Species and Functionalities. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.641185"},{"key":"ref_22","first-page":"38","article-title":"Microbial Dynamics Associated with Spontaneous Fermentation of Cocoa (Theobroma cacao L.) in Cameroon and Evaluation of the Quality of Marketable Beans","volume":"11","author":"Minyaka","year":"2022","journal-title":"Int. J. Nutr. Food Sci."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Schwan, R.F., and Fleet, G.H. (2015). The Functional Role of Lactic Acid Bacteria in Cocoa Bean Fermentation. Cocoa and Coffee Fermentations, Taylor & Francis.","DOI":"10.1201\/b17536"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1809","DOI":"10.1128\/AEM.02189-06","article-title":"Dynamics and Biodiversity of Populations of Lactic Acid Bacteria and Acetic Acid Bacteria Involved in Spontaneous Heap Fermentation of Cocoa Beans in Ghana","volume":"73","author":"Camu","year":"2007","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"109015","DOI":"10.1016\/j.ijfoodmicro.2020.109015","article-title":"Co-Culturing Fructophilic Lactic Acid Bacteria and Yeast Enhanced Sugar Metabolism and Aroma Formation during Cocoa Beans Fermentation","volume":"339","author":"Viesser","year":"2020","journal-title":"Int. J. Food Microbiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.ijfoodmicro.2016.07.004","article-title":"Detailed Analyses of the Bacterial Populations in Processed Cocoa Beans of Different Geographic Origin, Subject to Varied Fermentation Conditions","volume":"236","author":"Bortolini","year":"2016","journal-title":"Int. J. Food Microbiol."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Pacheco-Montealegre, M., D\u00e1vila-Mora, L., Botero-Rute, L., Reyes, A., and Caro-Quintero, A. (2020). Fine Resolution Analysis of Microbial Communities Provides Insights into the Variability of Cocoa Bean Fermentation. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.00650"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"964","DOI":"10.1016\/j.fm.2011.01.010","article-title":"Comparison of the Bacterial Species Diversity of Spontaneous Cocoa Bean Fermentations Carried out at Selected Farms in Ivory Coast and Brazil","volume":"28","author":"Papalexandratou","year":"2011","journal-title":"Food Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1326","DOI":"10.1016\/j.fm.2011.06.003","article-title":"Spontaneous Organic Cocoa Bean Box Fermentations in Brazil Are Characterized by a Restricted Species Diversity of Lactic Acid Bacteria and Acetic Acid Bacteria","volume":"28","author":"Papalexandratou","year":"2011","journal-title":"Food Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Tigrero-Vaca, J., Villavicencio-V\u00e1squez, M., Coronel, J., and Cevallos-Cevallos, J. (2025). Multi-Platform Metagenomic Characterization of the Microbial Community during Spontaneous Cacao Fermentation. Front. Bioeng. Biotechnol., 13.","DOI":"10.3389\/fbioe.2025.1630515"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1007\/s43393-023-00160-9","article-title":"Role of Fermentation and Microbes in Cacao Fermentation and Their Impact on Cacao Quality","volume":"3","author":"Hirko","year":"2023","journal-title":"Syst. Microbiol. Biomanufacturing"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Tigrero-Vaca, J., Maridue\u00f1a-Zavala, M.G., Liao, H.-L., Prado-Lince, M., Zambrano-Vera, C.S., Monserrate-Maggi, B., and Cevallos-Cevallos, J.M. (2022). Microbial Diversity and Contribution to the Formation of Volatile Compounds during Fine-Flavor Cacao Bean Fermentation. Foods, 11.","DOI":"10.3390\/foods11070915"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e0058421","DOI":"10.1128\/AEM.00584-21","article-title":"Metagenome-Assembled Genomes Contribute to Unraveling of the Microbiome of Cocoa Fermentation","volume":"87","author":"Almeida","year":"2021","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"113849","DOI":"10.1016\/j.foodres.2023.113849","article-title":"Integrating Shotgun Metagenomics and Metabolomics to Elucidate the Dynamics of Microbial Communities and Metabolites in Fine Flavor Cocoa Fermentation in Hainan","volume":"177","author":"Chang","year":"2023","journal-title":"Food Res. Int."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"e40542","DOI":"10.1016\/j.heliyon.2024.e40542","article-title":"Multiomics Analysis Reveals Microbial Diversity and Activity through Spontaneous Fermentation of Theobroma cacao","volume":"10","author":"Bahule","year":"2024","journal-title":"Heliyon"},{"key":"ref_36","first-page":"46","article-title":"Establishing Geographical Indicator of Fermented Cacao Beans Using Microbiome Fingerprinting","volume":"4","author":"Nugroho","year":"2024","journal-title":"J. Biotechnol. Nat. Sci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"28","DOI":"10.9734\/arrb\/2024\/v39i42072","article-title":"Assessment of Microbial Interactions between Potential Lactic Acid Bacteria Starters and Major Microorganisms During in vitro Fermentation in a Medium Similar to Cocoa Pulp in C\u00f4te d\u2019Ivoire","volume":"39","author":"Ahoussi","year":"2024","journal-title":"Annu. Res. Rev. Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"6694","DOI":"10.1128\/AEM.00594-11","article-title":"Interesting Starter Culture Strains for Controlled Cocoa Bean Fermentation Revealed by Simulated Cocoa Pulp Fermentations of Cocoa-Specific Lactic Acid Bacteria","volume":"77","author":"Lefeber","year":"2011","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Marwati, T., Kurniawan, F.I., Karisma, V.L.K., Hatmi, R., Fajariyah, A., Fitrotin, U., Djaafar, T., Wikandari, R., and Rahayu, E.S. (2024). Indigenous Lactic Acid Bacteria as a Biological Control Agent to Prevent Fungi Contamination in the Fermentation of Cocoa Beans. Food Sci. Technol., 44.","DOI":"10.5327\/fst.17923"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1016\/j.ijfoodmicro.2018.10.001","article-title":"Screening of Lactic Acid Bacteria and Yeast Strains to Select Adapted Anti-Fungal Co-Cultures for Cocoa Bean Fermentation","volume":"290","author":"Romanens","year":"2019","journal-title":"Int. J. Food Microbiol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"e0236824","DOI":"10.1128\/aem.02368-24","article-title":"Microbiome Metabolic Capacity Is Buffered against Phylotype Losses by Functional Redundancy","volume":"91","author":"Cross","year":"2025","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"7708","DOI":"10.1128\/AEM.01206-10","article-title":"Kinetic Analysis of Strains of Lactic Acid Bacteria and Acetic Acid Bacteria in Cocoa Pulp Simulation Media toward Development of a Starter Culture for Cocoa Bean Fermentation","volume":"76","author":"Lefeber","year":"2010","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1007\/s11274-021-03079-2","article-title":"Global Cocoa Fermentation Microbiome: Revealing New Taxa and Microbial Functions by next Generation Sequencing Technologies","volume":"37","author":"Viesser","year":"2021","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1111\/jam.13045","article-title":"The Cocoa Bean Fermentation Process: From Ecosystem Analysis to Starter Culture Development","volume":"121","author":"Vuyst","year":"2016","journal-title":"J. Appl. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"e02170","DOI":"10.1016\/j.heliyon.2019.e02170","article-title":"Metagenomics Analysis of Cocoa Bean Fermentation Microbiome Identifying Species Diversity and Putative Functional Capabilities","volume":"5","author":"Agyirifo","year":"2019","journal-title":"Heliyon"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Lima, C., De Castro, G., Solar, R., Vaz, A., Lobo, F., Pereira, G., Rodrigues, C., Vandenberghe, L., Pinto, L.R.M., and Da Costa, A.M. (2022). Unraveling Potential Enzymes and Their Functional Role in Fine Cocoa Beans Fermentation Using Temporal Shotgun Metagenomics. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.994524"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"103608","DOI":"10.1016\/j.fm.2020.103608","article-title":"Integrating Microbial Metagenomics and Physicochemical Parameters and a New Perspective on Starter Culture for Fine Cocoa Fermentation","volume":"93","author":"Carolina","year":"2021","journal-title":"Food Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Falcon\u00ed, C., Y\u00e1nez-Mendiz\u00e1bal, V., Haro, R., and Claudio, D. (2023). Inoculum of a Native Microbial Starter Cocktail to Optimize Fine-Aroma Cocoa (Theobroma cacao) Bean Fermentation. Agronomy, 13.","DOI":"10.20944\/preprints202307.0954.v1"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Illeghems, K., De Vuyst, L., and Weckx, S. (2015). Comparative Genome Analysis of the Candidate Functional Starter Culture Strains Lactobacillus fermentum 222 and Lactobacillus plantarum 80 for Controlled Cocoa Bean Fermentation Processes. BMC Genom., 16.","DOI":"10.1186\/s12864-015-1927-0"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.fm.2015.03.005","article-title":"Applying Meta-Pathway Analyses through Metagenomics to Identify the Functional Properties of the Major Bacterial Communities of a Single Spontaneous Cocoa Bean Fermentation Process Sample","volume":"50","author":"Illeghems","year":"2015","journal-title":"Food Microbiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"116410","DOI":"10.1016\/j.foodres.2025.116410","article-title":"Metataxonomic, Chemical, and Sensory Aspects of Cocoa Fermentation Using Single Hybrids and Combinations of Hybrids and Their Effects on Chocolate Quality","volume":"210","author":"Dias","year":"2025","journal-title":"Food Res. Int."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"102315","DOI":"10.1016\/j.fochx.2025.102315","article-title":"Investigation of Aromatic Compounds and Olfactory Profiles in Cocoa Pulp Fermentation Using Yeast-Based Starters: A Volatilomics and Machine Learning Approach","volume":"26","author":"Chang","year":"2025","journal-title":"Food Chem. X"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"109034","DOI":"10.1016\/j.foodres.2020.109034","article-title":"Does Quorum Sensing Play a Role in Microbial Shifts along Spontaneous Fermentation of Cocoa Beans? An in silico Perspective","volume":"131","author":"Almeida","year":"2020","journal-title":"Food Res. Int."},{"key":"ref_54","first-page":"101851","article-title":"Impact of Fermentation and Drying on the Bioactive Compounds and Physicochemical Properties of Four Cacao Genotypes in Northwestern Peru","volume":"21","author":"Yoplac","year":"2025","journal-title":"J. Agric. Food Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"28","DOI":"10.9734\/jamb\/2021\/v21i1130399","article-title":"Isolation and Identification of Lactic Acid Bacteria and Acetic Acid Bacteria Playing a Lead Role in the Fermentation of Cocoa in Fako Division of Cameroon","volume":"21","author":"Levai","year":"2021","journal-title":"J. Adv. Microbiol."},{"key":"ref_56","first-page":"1046","article-title":"Cross-Biome Microbial Networks Reveal Functional Redundancy and Suggest Genome Reduction through Functional Complementarity","volume":"7","author":"Bastolla","year":"2022","journal-title":"Commun. Biol."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Van de Voorde, D., D\u00edaz-Mu\u00f1oz, C., Hernandez, C.E., Weckx, S., and De Vuyst, L. (2023). Yeast Strains Do Have an Impact on the Production of Cured Cocoa Beans, as Assessed with Costa Rican Trinitario Cocoa Fermentation Processes and Chocolates Thereof. Front. Microbiol., 14.","DOI":"10.3389\/fmicb.2023.1232323"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Freim\u00fcller Leischtfeld, S., H\u00e4mmerli, A., Lehmann, A., T\u00f6nz, A., Beck, B.M., Wild, J., Weis, S., Neutsch, L., and Miescher Schwenninger, S. (2025). Wickerhamomyces pijperi: An Up-And-Coming Yeast with Pectinolytic Activity Suitable for Cocoa Bean Fermentation. Appl. Microbiol., 5.","DOI":"10.3390\/applmicrobiol5020043"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"21904","DOI":"10.1038\/s41598-021-01427-8","article-title":"Dissecting Fine-Flavor Cocoa Bean Fermentation through Metabolomics Analysis to Break down the Current Metabolic Paradigm","volume":"11","author":"Cala","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"D\u00edaz-Mu\u00f1oz, C., Van de Voorde, D., Comasio, A., Verce, M., Hernandez, C.E., Weckx, S., and De Vuyst, L. (2021). Curing of Cocoa Beans: Fine-Scale Monitoring of the Starter Cultures Applied and Metabolomics of the Fermentation and Drying Steps. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.616875"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Guti\u00e9rrez-R\u00edos, H.G., Su\u00e1rez-Quiroz, M.L., Hern\u00e1ndez-Estrada, Z.J., Castellanos-Onorio, O.P., Alonso-Villegas, R., Rayas-Duarte, P., Cano-Sarmiento, C., Figueroa-Hern\u00e1ndez, C.Y., and Gonz\u00e1lez-Rios, O. (2022). Yeasts as Producers of Flavor Precursors during Cocoa Bean Fermentation and Their Relevance as Starter Cultures: A Review. Fermentation, 8.","DOI":"10.3390\/fermentation8070331"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1848","DOI":"10.1128\/AEM.03344-13","article-title":"Oxidation of Metabolites Highlights the Microbial Interactions and Role of Acetobacter pasteurianus during Cocoa Bean Fermentation","volume":"80","author":"Moens","year":"2014","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"4702","DOI":"10.1128\/AEM.01048-14","article-title":"The Key to Acetate: Metabolic Fluxes of Acetic Acid Bacteria under Cocoa Pulp Fermentation-Simulating Conditions","volume":"80","author":"Adler","year":"2014","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"205","DOI":"10.22146\/agritech.86796","article-title":"Pulp Reduction and Addition of Indigenous Microorganisms as Starter: Effects on Fermented Cocoa Bean Characteristics","volume":"45","author":"Fajariyah","year":"2025","journal-title":"agriTECH"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/j.ijfoodmicro.2015.09.004","article-title":"Molecular Identification and Physiological Characterization of Yeasts, Lactic Acid Bacteria and Acetic Acid Bacteria Isolated from Heap and Box Cocoa Bean Fermentations in West Africa","volume":"216","author":"Visintin","year":"2016","journal-title":"Int. J. Food Microbiol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Nguyen, T.C., Hoang, T.T., and Nguyen, H.H. (2025). Evaluation of Anti-Aspergillus Flavus Activity of Lactic Acid Bacteria Isolated from Vietnamese Fermented Cocoa Beans. Microbiol. Res., 16.","DOI":"10.3390\/microbiolres16060111"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/j.foodres.2018.10.002","article-title":"Antifungal Activity of Yeasts and Lactic Acid Bacteria Isolated from Cocoa Bean Fermentations","volume":"115","author":"Ruggirello","year":"2018","journal-title":"Food Res. Int."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1407","DOI":"10.25083\/rbl\/25.2\/1407.1417","article-title":"Antifungal Properties of Lactic Acid Bacteria Isolated from Cocoa Beans Fermentation in the Centre Region of Cameroon","volume":"25","author":"Fanche","year":"2020","journal-title":"Rom. Biotechnol. Lett."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Santos, J.P.N., Rodrigues, G.V.P., Ferreira, L.Y.M., Monteiro, G.P., Fonseca, P.L.C., Lopes, \u00cd.S., Flor\u00eancio, B.S., da Silva Junior, A.B., Ambr\u00f3sio, P.E., and Pirovani, C.P. (2024). The Virome of Cocoa Fermentation-Associated Microorganisms. Viruses, 16.","DOI":"10.3390\/v16081226"},{"key":"ref_70","first-page":"3","article-title":"Bacteriophage-Host Interactions as a Platform to Establish the Role of Phages in Modulating the Microbial Composition of Fermented Foods","volume":"1","author":"White","year":"2022","journal-title":"Microbiome Res. Rep."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Alseth, E.O., Custodio, R., Sundius, S.A., Kuske, R.A., Brown, S.P., and Westra, E.R. (2024). The Impact of Phage and Phage Resistance on Microbial Community Dynamics. PLoS Biol., 22.","DOI":"10.1371\/journal.pbio.3002346"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Ledormand, P., Desmasures, N., Schlusselhuber, M., Sesbo\u00fc\u00e9, A., Ledauphin, J., and Dalmasso, M. (2022). Phages Shape Microbial Dynamics and Metabolism of a Model Community Mimicking Cider, a Fermented Beverage. Viruses, 14.","DOI":"10.3390\/v14102283"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"e0042521","DOI":"10.1128\/AEM.00425-21","article-title":"Influence of Taxonomic and Functional Content of Microbial Communities on the Quality of Fermented Cocoa Pulp-Bean Mass","volume":"87","author":"Ferrocino","year":"2021","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1904","DOI":"10.1007\/s13197-019-04226-2","article-title":"Acetic Acid Bacteria (AAB) Involved in Cocoa Fermentation from Ivory Coast: Species Diversity and Performance in Acetic Acid Production","volume":"57","author":"Soumahoro","year":"2020","journal-title":"J. Food Sci. Technol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"210274","DOI":"10.1098\/rsos.210274","article-title":"Exploring Cocoa Bean Fermentation Mechanisms by Kinetic Modelling","volume":"9","author":"Ullrich","year":"2022","journal-title":"R. Soc. Open Sci."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Ghisolfi, R., Bandini, F., Vaccari, F., Bellotti, G., Bortolini, C., Patrone, V., Puglisi, E., and Morelli, L. (2023). Bacterial and Fungal Communities Are Specifically Modulated by the Cocoa Bean Fermentation Method. 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