Article(id=1321170164104679426, tenantId=1146029695717560320, journalId=1260987677001138203, issueId=1321170163563614210, articleNumber=null, orderNo=null, doi=10.13386/j.issn1002-0306.2025010133, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1736784000000, receivedDateStr=2025-01-14, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1792921113276, onlineDateStr=2026-10-25, pubDate=1769875200000, pubDateStr=2026-02-01, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1792921113276, onlineIssueDateStr=2026-10-25, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1792921113276, creator=13701087609, updateTime=1792921113276, updator=13701087609, issue=Issue{id=1321170163563614210, tenantId=1146029695717560320, journalId=1260987677001138203, year='2026', volume='47', issue='3', pageStart='1', pageEnd='497', issueExtLink='null', onlineDate='null', pubDate='1769875200000', pubDateStr='2026-02-01', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1792921113148, creator='13701087609', updateTime=1792921113148, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext=null, issueFiles=null, downloadFileDto=null}, startPage=12, endPage=21, ext={EN=ArticleExt(id=1321170164276645891, articleId=1321170164104679426, tenantId=1146029695717560320, journalId=1260987677001138203, language=EN, title=Research Progress on Relationship between Microbial and Flavor Development in Coffee Fermentation, columnId=1261336279195726298, journalTitle=Science and Technology of Food Industry, columnName=Papers Invited by Guest Editors, runingTitle=null, highlight=null, articleAbstract=

Fermentation serves as a critical process in coffee fruit processing for flavor development. Microbial metabolites produced during this biochemical transformation substantially contribute to the aromatic profile and flavor characteristics of roasted coffee beans. Microorganisms degrade coffee fruit mucilage and produce metabolites such as esters, ketones, alcohols, acids, and aldehydes. The interactions between these compounds contribute to the complexity of the final flavor profile of coffee. This review systematically summarizes the predominant microbial community structures and succession of coffee fermentation processes (wet, dry, and semi-dry methods), elucidates the impacts of lactic acid bacteria, yeasts, other bacterial, and filamentous fungi on flavor development, and the underlying metabolic pathways involved in these biochemical interactions. This review aims to establish a theoretical framework for targeted flavor modulation in coffee and to advance the sustainable efficiency of coffee industry practices.

, authors=Jie NI1, 2, Huihua TANG1, Haoyuan TANG2, Yanping TANG2, Hong LI1, *, Yongjin HU1, *, authorsList=Jie NI, Huihua TANG, Haoyuan TANG, Yanping TANG, Hong LI, Yongjin HU, authorCompany=null, correspAuthors=Hong LI, Yongjin HU, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright © 2026 Science and Technology of Food Industry. All rights reserved., copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1321170165262307339, articleId=1321170164104679426, tenantId=1146029695717560320, journalId=1260987677001138203, language=CN, title=咖啡发酵过程中的微生物及其与风味形成关系的研究进展, columnId=1321170164360531972, journalTitle=食品工业科技, columnName=特邀主编专栏—咖啡、可可、茶等特色饮料作物加工(客座主编:董文江、许勇泉、付才力), runingTitle=null, highlight=null, articleAbstract=

发酵是咖啡果实加工中形成风味的重要工序,发酵过程中微生物产生的代谢物对烘焙咖啡豆的香气和风味有显著贡献。微生物降解咖啡果粘液的同时产生酯类、酮类、醇类、酸类和醛类代谢产物,这些化合物之间的相互作用形成了咖啡最终风味特征的复杂性。本文综述了咖啡湿法、干法和半干法发酵过程中的主要微生物群落组成和变化;论述了乳酸菌、酵母菌、其他细菌和丝状真菌对咖啡风味形成的影响及相关的代谢机制,旨在为咖啡风味定向调控提供理论基础,并进一步促进咖啡产业高效发展。

, authors=倪婕1, 2, 汤回花1, 唐浩源2, 唐艳平2, 李宏1, *, 胡永金1, *, authorsList=倪婕, 汤回花, 唐浩源, 唐艳平, 李宏, 胡永金, authorCompany=null, correspAuthors=李宏, 胡永金, authorNote=

倪婕(1996−)(ORCID:0009−0007−1568−5283),女,博士,讲师,研究方向:食品微生物资源利用,E-mail:

, correspAuthorsNote=
李宏(1974−),男,博士,研究员,研究方向:农产品加工,E-mail:。
胡永金(1972−)(ORCID:0000−0001−5149−0079),男,博士,教授,研究方向:食品微生物资源与利用,E-mail:。
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Science and Technology of Food Industry, 2025, 46(7): 1−10.], articleTitle=null, refAbstract=null), Reference(id=1321170178692468897, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=84, rfOrder=92, authorNames=null, journalName=null, refType=null, unstructuredReference=MIAO Xingyu, WANG Qinghua, CHAI Chunrong, et al. Effects of enhanced fermentation of Saccharomyces cerevisiae and Lactobacillus plantarum on volatile flavor of coffee[J]. Science and Technology of Food Industry, 2025, 46(7): 1−10., articleTitle=null, refAbstract=null), Reference(id=1321170178776354978, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=85, rfOrder=93, authorNames=null, journalName=null, refType=null, unstructuredReference=MARTINEZ S J, RABELO M H S, BRESSANI A P P, et al. Novel stainless steel tanks enhances coffee fermentation quality[J]. 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Journal of Food Composition and Analysis, 2025, 139: 107073., articleTitle=null, refAbstract=null), Reference(id=1321170178914767012, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=87, rfOrder=95, authorNames=null, journalName=null, refType=null, unstructuredReference=ASWATHI K N, SHIRKE A, PRAVEEN A, et al. Functioning of Saccharomyces cerevisiae in honey coffee (Coffea canephora) and their effect on metabolites, volatiles and flavor profiles[J]. Food Research International, 2024, 180: 114092., articleTitle=null, refAbstract=null), Reference(id=1321170178990264485, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=88, rfOrder=96, authorNames=null, journalName=null, refType=null, unstructuredReference=LEE L W, CHEONG M W, CURRAN P, et al. Coffee fermentation and flavor–An intricate and delicate relationship[J]. Food Chemistry, 2015, 185: 182−191., articleTitle=null, refAbstract=null), Reference(id=1321170179061567654, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=89, rfOrder=97, authorNames=null, journalName=null, refType=null, unstructuredReference=SENINDE D R, CHAMBERS E IV. Coffee flavor: A review[J]. Beverages, 2020, 6(3): 44., articleTitle=null, refAbstract=null), Reference(id=1321170179145453735, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=90, rfOrder=98, authorNames=null, journalName=null, refType=null, unstructuredReference=WALKER G M, STEWART G G. Saccharomyces cerevisiae in the production of fermented beverages[J]. Beverages, 2016, 2(4): 30., articleTitle=null, refAbstract=null), Reference(id=1321170179233534120, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=91, rfOrder=99, authorNames=null, journalName=null, refType=null, unstructuredReference=CHIN X H, HO S, CHAN G, et al. Aromatic yeasts: Interactions and implications in coffee fermentation aroma profiles[J]. 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Main microbial community in different processing methods

, figureFileSmall=null, figureFileBig=null, tableContent=
主要菌群湿法干法半干法文献
乳酸菌明串珠菌属(Leuconostoc),乳杆菌属(Lactobacillus),乳球菌属(Lactococcus)乳植杆菌属(Lactiplantibacillus),乳杆菌属(Lactobacillus),乳球菌属(Lactococcus),魏斯氏菌属(Weissella)和片球菌属(Pediococcus)乳植杆菌属(Lactiplantibacillus),乳杆菌属(Lactobacillus),明串珠菌属(Leuconostoc),乳球菌属(Lactococcus)[4,9]
酵母菌酿酒酵母属(Saccharomyces),汉逊酵母属(Hanseniaspora),假丝酵母属(Candida),毕赤酵母属(Pichia),红酵母属(Rhodotorula)酿酒酵母属(Saccharomyces),德巴利酵母属(Debaryomyces),毕赤酵母属(Pichia),有孢圆酵母属(Torulaspora),隐球酵母属(Cryptococcus)毕赤酵母属(Pichia),酿酒酵母属(Saccharomyces),有孢圆酵母属(Torulaspora),德巴利酵母属(Debaryomyces)[8,10]
其他细菌不动杆菌属(Acinetobacter),克雷伯菌属(Klebsiella),肠杆菌属(Enterobacter),芽孢杆菌属(Bacillus),葡糖杆菌属(Gluconobacter)肠杆菌属(Enterobacter),不动杆菌属(Acinetobacter),哈夫尼菌属(Hafnia),克雷伯菌属(Klebsiella),假单胞菌属(Pseudomonas),变形杆菌属(Proteus), 芽孢杆菌属(Bacillus)肠杆菌属(Enterobacter),芽孢杆菌属(Bacillus),克雷伯菌属 (Klebsiella) ,不动杆菌属(Acinetobacter),芽孢杆菌属(Bacillus)[8−9]
丝状真菌曲霉属(Aspergillus),青霉菌属(Penicillium)青霉菌属(Penicillium),镰刀菌属(Fusarium),枝孢属(Cladosporium),曲霉属(Aspergillus)曲霉属(Aspergillus),枝孢属(Cladosporium),青霉菌属(Penicillium)[11]
), ArticleFig(id=1321170168424812610, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, language=CN, label=表1, caption=

不同加工方式存在的主要微生物

, figureFileSmall=null, figureFileBig=null, tableContent=
主要菌群湿法干法半干法文献
乳酸菌明串珠菌属(Leuconostoc),乳杆菌属(Lactobacillus),乳球菌属(Lactococcus)乳植杆菌属(Lactiplantibacillus),乳杆菌属(Lactobacillus),乳球菌属(Lactococcus),魏斯氏菌属(Weissella)和片球菌属(Pediococcus)乳植杆菌属(Lactiplantibacillus),乳杆菌属(Lactobacillus),明串珠菌属(Leuconostoc),乳球菌属(Lactococcus)[4,9]
酵母菌酿酒酵母属(Saccharomyces),汉逊酵母属(Hanseniaspora),假丝酵母属(Candida),毕赤酵母属(Pichia),红酵母属(Rhodotorula)酿酒酵母属(Saccharomyces),德巴利酵母属(Debaryomyces),毕赤酵母属(Pichia),有孢圆酵母属(Torulaspora),隐球酵母属(Cryptococcus)毕赤酵母属(Pichia),酿酒酵母属(Saccharomyces),有孢圆酵母属(Torulaspora),德巴利酵母属(Debaryomyces)[8,10]
其他细菌不动杆菌属(Acinetobacter),克雷伯菌属(Klebsiella),肠杆菌属(Enterobacter),芽孢杆菌属(Bacillus),葡糖杆菌属(Gluconobacter)肠杆菌属(Enterobacter),不动杆菌属(Acinetobacter),哈夫尼菌属(Hafnia),克雷伯菌属(Klebsiella),假单胞菌属(Pseudomonas),变形杆菌属(Proteus), 芽孢杆菌属(Bacillus)肠杆菌属(Enterobacter),芽孢杆菌属(Bacillus),克雷伯菌属 (Klebsiella) ,不动杆菌属(Acinetobacter),芽孢杆菌属(Bacillus)[8−9]
丝状真菌曲霉属(Aspergillus),青霉菌属(Penicillium)青霉菌属(Penicillium),镰刀菌属(Fusarium),枝孢属(Cladosporium),曲霉属(Aspergillus)曲霉属(Aspergillus),枝孢属(Cladosporium),青霉菌属(Penicillium)[11]
), ArticleFig(id=1321170168491921475, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, language=EN, label=Table 2, caption=

Main volatile compounds in fermented coffee inoculated with different yeast strains

, figureFileSmall=null, figureFileBig=null, tableContent=
酵母菌挥发性物质特征香味文献
酿酒酵母
Saccharomyces cerevisiae
醛(乙醛;己醛)、乙酯(琥珀酸二乙酯;乙酸乙酯) 、高级醇(1-辛醇;2-甲基-1丁醇) ;酮(2-己酮;3,4 -二甲基- 2 -戊酮);萜烯(2 -甲基 - 1,3 -丁二烯)柑橘、焦糖、香蕉、腰果、红糖、苹果酸[41,43]
德氏圆酵母
Torulaspora delbrueckii
醛(2-苯基-2-丁烯醛; 2 -甲基- 2 -戊烯醛)、醇(乙醇;硫醇)、酯(果味酯)、萜烯、酮(3,4-二甲基-2-戊酮;1-羟基-2-丙烷酮)可可香、奶香、甜味[44]
发酵毕赤酵母
Pichia fermentans
醛(乙醛;己醛)、酯(乙酸己酯;乙酸异丁酯;乙酸异戊酯)、高级醇(2-甲基-1-丁醇;3-甲基-1-丁醇;1-戊醇;2-己醇;1-辛醇;1-癸醇)、二酮(2,3-丁二酮)果香、黄油、香草、花香[45]
平滑念珠菌
Candida parapsilosis
醛(乙醛;糠醛;癸醛)、 酮(乙偶姻)、高级醇(2-甲基丁醇;1-戊醇;2-庚醇;糠醇;2-苯乙醇) 、萜烯(芳樟醇;香茅醇;薄荷醇)焦糖、柠檬酸、苹果、樱桃、青草味[46]
解脂耶氏酵母
Yarrowia lipolytica
醛(辛醛)、酯(2-甲基丁酸己酯)、吡啶( 2-甲氧基吡啶)、萜烯(1-乙酰基-2-甲基环戊烯;顺,反-1,3,5-十一碳三烯;月桂烯;苯丙烯;γ-萜品烯)、酮( 6,10 二甲基-5,9-十一碳二烯-2-酮)柑橘、焦糖、甜味、苹果、糖果、玫瑰、发酵茶[47]
葡萄汁有孢汉逊酵母Hanseniaspora uvarum醇(乙醇)、酯(异戊酸香叶酯;乙酸苯乙酯;邻苯二甲酸二丁酯)、酮(2, 3-二氢-3, 5二羟基-6-甲基-4 (H)-吡喃-4-酮)花香、果香[48]
库德里阿兹威毕赤酵母
Pichia kudriavzevii
醇(乙醇)、酮(异戊醇呋喃)、吡嗪类、内酯类蜂蜜、麦芽和浆果味[36,49]
季里蒙毕赤酵母
Pichia guilliermondii
醛(乙醛;糠醛;癸醛) 、高级醇(糠醇)、萜烯(芳樟醇)陈皮、柑橘、花香、杏仁、甜味、焦糖、柠檬[50]
), ArticleFig(id=1321170168575807556, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1321170164104679426, language=CN, label=表2, caption=

不同酵母菌接种发酵咖啡中的挥发性香气物质

, figureFileSmall=null, figureFileBig=null, tableContent=
酵母菌挥发性物质特征香味文献
酿酒酵母
Saccharomyces cerevisiae
醛(乙醛;己醛)、乙酯(琥珀酸二乙酯;乙酸乙酯) 、高级醇(1-辛醇;2-甲基-1丁醇) ;酮(2-己酮;3,4 -二甲基- 2 -戊酮);萜烯(2 -甲基 - 1,3 -丁二烯)柑橘、焦糖、香蕉、腰果、红糖、苹果酸[41,43]
德氏圆酵母
Torulaspora delbrueckii
醛(2-苯基-2-丁烯醛; 2 -甲基- 2 -戊烯醛)、醇(乙醇;硫醇)、酯(果味酯)、萜烯、酮(3,4-二甲基-2-戊酮;1-羟基-2-丙烷酮)可可香、奶香、甜味[44]
发酵毕赤酵母
Pichia fermentans
醛(乙醛;己醛)、酯(乙酸己酯;乙酸异丁酯;乙酸异戊酯)、高级醇(2-甲基-1-丁醇;3-甲基-1-丁醇;1-戊醇;2-己醇;1-辛醇;1-癸醇)、二酮(2,3-丁二酮)果香、黄油、香草、花香[45]
平滑念珠菌
Candida parapsilosis
醛(乙醛;糠醛;癸醛)、 酮(乙偶姻)、高级醇(2-甲基丁醇;1-戊醇;2-庚醇;糠醇;2-苯乙醇) 、萜烯(芳樟醇;香茅醇;薄荷醇)焦糖、柠檬酸、苹果、樱桃、青草味[46]
解脂耶氏酵母
Yarrowia lipolytica
醛(辛醛)、酯(2-甲基丁酸己酯)、吡啶( 2-甲氧基吡啶)、萜烯(1-乙酰基-2-甲基环戊烯;顺,反-1,3,5-十一碳三烯;月桂烯;苯丙烯;γ-萜品烯)、酮( 6,10 二甲基-5,9-十一碳二烯-2-酮)柑橘、焦糖、甜味、苹果、糖果、玫瑰、发酵茶[47]
葡萄汁有孢汉逊酵母Hanseniaspora uvarum醇(乙醇)、酯(异戊酸香叶酯;乙酸苯乙酯;邻苯二甲酸二丁酯)、酮(2, 3-二氢-3, 5二羟基-6-甲基-4 (H)-吡喃-4-酮)花香、果香[48]
库德里阿兹威毕赤酵母
Pichia kudriavzevii
醇(乙醇)、酮(异戊醇呋喃)、吡嗪类、内酯类蜂蜜、麦芽和浆果味[36,49]
季里蒙毕赤酵母
Pichia guilliermondii
醛(乙醛;糠醛;癸醛) 、高级醇(糠醇)、萜烯(芳樟醇)陈皮、柑橘、花香、杏仁、甜味、焦糖、柠檬[50]
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咖啡发酵过程中的微生物及其与风味形成关系的研究进展
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倪婕 1, 2 , 汤回花 1 , 唐浩源 2 , 唐艳平 2 , 李宏 1, * , 胡永金 1, *
食品工业科技 | 特邀主编专栏—咖啡、可可、茶等特色饮料作物加工(客座主编:董文江、许勇泉、付才力) 2026,47(3): 12-21
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食品工业科技 |特邀主编专栏—咖啡、可可、茶等特色饮料作物加工(客座主编:董文江、许勇泉、付才力) 2026 , 47 (3) : 12 -21
咖啡发酵过程中的微生物及其与风味形成关系的研究进展
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倪婕1, 2 , 汤回花1, 唐浩源2, 唐艳平2, 李宏1, * , 胡永金1, *
作者信息
  • 1云南农业大学食品科学技术学院,云南昆明 650201
  • 2桂林师范学院化学与生命健康学院,广西桂林 541199
通讯作者:
李宏(1974−),男,博士,研究员,研究方向:农产品加工,E-mail:。
胡永金(1972−)(ORCID:0000−0001−5149−0079),男,博士,教授,研究方向:食品微生物资源与利用,E-mail:。
作者简介:

倪婕(1996−)(ORCID:0009−0007−1568−5283),女,博士,讲师,研究方向:食品微生物资源利用,E-mail:

Research Progress on Relationship between Microbial and Flavor Development in Coffee Fermentation
Jie NI1, 2 , Huihua TANG1, Haoyuan TANG2, Yanping TANG2, Hong LI1, * , Yongjin HU1, *
Affiliations
  • 1College of Food Science and Technology, Yunnan Agricultural University, Kunming 650201, China
  • 2College of Chemistry and Life Health Science, Guilin Normal University, Guilin 541199, China
出版时间: 2026-02-01 doi: 10.13386/j.issn1002-0306.2025010133
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发酵是咖啡果实加工中形成风味的重要工序,发酵过程中微生物产生的代谢物对烘焙咖啡豆的香气和风味有显著贡献。微生物降解咖啡果粘液的同时产生酯类、酮类、醇类、酸类和醛类代谢产物,这些化合物之间的相互作用形成了咖啡最终风味特征的复杂性。本文综述了咖啡湿法、干法和半干法发酵过程中的主要微生物群落组成和变化;论述了乳酸菌、酵母菌、其他细菌和丝状真菌对咖啡风味形成的影响及相关的代谢机制,旨在为咖啡风味定向调控提供理论基础,并进一步促进咖啡产业高效发展。

咖啡  /  发酵  /  微生物菌群  /  风味成分  /  风味机理

Fermentation serves as a critical process in coffee fruit processing for flavor development. Microbial metabolites produced during this biochemical transformation substantially contribute to the aromatic profile and flavor characteristics of roasted coffee beans. Microorganisms degrade coffee fruit mucilage and produce metabolites such as esters, ketones, alcohols, acids, and aldehydes. The interactions between these compounds contribute to the complexity of the final flavor profile of coffee. This review systematically summarizes the predominant microbial community structures and succession of coffee fermentation processes (wet, dry, and semi-dry methods), elucidates the impacts of lactic acid bacteria, yeasts, other bacterial, and filamentous fungi on flavor development, and the underlying metabolic pathways involved in these biochemical interactions. This review aims to establish a theoretical framework for targeted flavor modulation in coffee and to advance the sustainable efficiency of coffee industry practices.

coffee  /  fermentation  /  microbial community  /  flavor components  /  flavor mechanism
倪婕, 汤回花, 唐浩源, 唐艳平, 李宏, 胡永金. 咖啡发酵过程中的微生物及其与风味形成关系的研究进展. 食品工业科技, 2026 , 47 (3) : 12 -21 . DOI: 10.13386/j.issn1002-0306.2025010133
Jie NI, Huihua TANG, Haoyuan TANG, Yanping TANG, Hong LI, Yongjin HU. Research Progress on Relationship between Microbial and Flavor Development in Coffee Fermentation[J]. Science and Technology of Food Industry, 2026 , 47 (3) : 12 -21 . DOI: 10.13386/j.issn1002-0306.2025010133
成熟的咖啡果在采摘后经初加工得到咖啡豆。我国对咖啡果常见的初加工方法有湿法、干法和半干法[1]。传统的湿法加工是将成熟的咖啡果在采摘后立即进行浮选清洗,机械去除果皮和果肉,并放入水槽中发酵24~96 h以去除果胶层,再干燥至含水量达到10%~12%[2]。干法加工是将完整的咖啡果置于露天晒台中,同时进行发酵和干燥过程,再将外壳去除得到咖啡豆[3]。半干法则是介于干法和湿法之间的一种方法,将机械脱皮和脱胶后的咖啡豆不进行清洗而直接晾晒干燥,在干燥过程中进行发酵[4]。初加工过程中,微生物与咖啡果实的内源物质之间相互作用产生多种挥发性风味物质,这些物质大部分在烘焙后仍保留在咖啡豆中,赋予了咖啡独特的风味[5−6]。酵母菌、细菌和丝状真菌等多种微生物共同作用,降解了咖啡豆的粘液层,抑制不良微生物的生长,还能产生酯类、酮类、醇类、酸类和醛类等代谢物[7−8]。这些代谢化合物之间的相互作用对焙炒咖啡豆的风味有显著贡献,导致了咖啡饮料感官特征的复杂性。目前相关研究多集中于咖啡加工工艺对风味形成的影响,对咖啡发酵过程中涉及的微生物与风味形成关系及相关代谢机制缺乏系统总结。因此,本文聚焦于咖啡加工中的发酵过程,综述了咖啡湿法、干法和半干法发酵过程中的主要微生物,阐述了乳酸菌、酵母菌、其他细菌和丝状真菌对咖啡风味形成的影响及咖啡发酵代谢风味的形成机制。
咖啡加工过程中,酵母菌、乳酸菌、其他细菌和丝状真菌大量存在,不同加工方式产生的主要微生物有所不同,如表1所示。发酵开始时,细菌数量大于酵母菌数量,最常见的细菌属有乳杆菌属(Lactobacillus)、明串珠菌属(Leuconostoc)、芽孢杆菌属(Bacillus)、不动杆菌属(Acinetobacter)和魏斯氏菌属(Weissella)[9]。随着发酵的进行,pH下降,酵母菌因耐酸能力强而成为主要微生物群,常见的酵母菌属有酿酒酵母属(Saccharomyces)、毕赤酵母属(Pichia)、假丝酵母属(Candida)、红酵母属(Rhodotorula)和汉逊酵母属(Hanseniaspora)[10]。发酵过程中产生的丝状真菌相对较少,主要包括曲霉属(Aspergillus)和青霉属(Penicillium)[11]。
在湿法加工中,咖啡豆上的粘液层是发酵的主要基质,水浸泡处理为微生物的生长提供了高湿度和低氧条件,使细菌和酵母菌成为其中主要的种群,丝状真菌则受到抑制[12]。Elhalis等[13]研究了澳洲咖啡湿法加工过程中的微生物变化,结果表明乳酸菌以肠膜明串珠菌(Leuconostoc mesenteroides)和乳酸乳球菌(Lactococcus lactis)为主,分别在发酵前24 h增长到5.5和5.0 log CFU/g后下降;酵母菌数量呈上升趋势,发酵结束时(36 h)达到最大值5.5 log CFU/g,主要由汉逊酵母属和毕赤酵母属组成。Haile等[14]采用湿法发酵咖啡鲜果(96 h),观察到酵母数量在发酵48 h达到最大值,其中酿酒酵母数量(8.23 log CFU/g)在发酵48 h时显著高于其他酵母。da Silva等[15]研究了巴西咖啡果在湿法发酵过程中的细菌群落变化,发酵初期以明串珠菌属(52.47%)为主,其次为葡糖杆菌属(Gluconobacter)(27.20%);12 h后,葡糖杆菌属显著增加,达到57.40%;发酵结束时(24 h),明串珠菌属增加到69.92%,葡糖杆菌属稳定在20.16%。Shen等[16]研究了云南阿拉比卡咖啡湿法发酵过程中的微生物群落,结果表明发酵初期细菌中无色杆菌属(Achromobacter)的丰度最高(12.95%),在24 h达到24.69%后呈下降趋势;魏斯氏菌属和明串珠菌属分别从0.15%增加到33.74%和0.00%增加到6.39%(24 h)。就真菌而言,曲霉属在12 h达到10.76%后开始减少,36 h时为最低值(0.90%);酵母菌以汉逊酵母属的丰度最高,由3.45%(0 h)增加至26.80%(24 h)。综上,在湿法发酵过程中,以乳酸菌为代表的细菌通常在前期较为活跃,酵母菌因耐酸在发酵中后期占据主导地位,抑制产毒素真菌的生长。但不同地区不同品种咖啡的微生物多样性有很大差异,发酵过程中优势微生物的组成也有所不同。
干法加工通常需经过10~25 d的日晒过程,由于自然发酵时间长,涉及的微生物群更具多样性和复杂性。de Brnyn等[17]研究了干法加工咖啡豆的微生物群落变化,结果表明咖啡鲜果在堆积发酵过程中,细菌群落中明串珠菌、醋酸杆菌的丰度较高,其中明串珠菌在发酵的第10 d达到最大值(约62%)后下降;真菌群落则以毕赤酵母、假丝酵母属占主导,毕赤酵母属丰度在发酵第20 d达到最大值(约94%)。Silva等[18]研究表明阿拉比卡咖啡在干法发酵初期(0 d)由细菌主导(96.3%),发酵第8 d细菌数量减少至50%,并出现丝状真菌。第14 d细菌数量下降至<10%,与此同时,酵母菌的丰度不断升高,最常见的属有德巴利酵母(Debaryomyces)(27%)、毕赤酵母(18.9%)和假丝酵母(8.0%)。在干法发酵中,微生物分布主要以细菌为主,初期以革兰氏阴性菌占主导地位,随着果肉中水分和糖含量降低,其数量和种类逐渐减少,酵母菌和丝状真菌的丰度则不断升高。
半干法是介于咖啡干法和湿法加工的一种中间工艺。Vilela等[8]从半干法发酵咖啡豆中共分离出728株菌,酵母菌占52.47%,细菌占39.28%,丝状真菌占8.24%。在发酵的前24 h细菌数量最多(7.0 log CFU/g),酵母菌数量次之(3.7 log CFU/g),丝状真菌占少数(2.0 log CFU/g);发酵后期酵母菌占主导地位,120 h后达到6.9 log CFU/g,细菌降至2.0 log CFU/g。Ribeiro等[19]重点监测了酿酒酵母属在半干法发酵过程中的变化,结果表明咖啡果实中存在酿酒酵母(3.8 log CFU/g),发酵24 h后其种群数量增长约33%,在36 h后呈下降趋势。这种微生物优势群落的更替,与发酵条件(如pH、底物利用)密切相关,直接影响咖啡豆的代谢产物积累与最终风味形成。
咖啡的自然发酵是以促进果胶降解为主要目的,而科学研究表明,咖啡豆的品质特性与发酵过程中的微生物及其转化的风味物质密切相关[20]。咖啡的发酵是一个复杂的过程,微生物群落间通过代谢产物交换、群体感应调控及生存竞争等作用,决定了咖啡的最终品质[21]。微生物的生长会产生一系列代谢物,这些物质扩散到咖啡豆中并释放香气,包括高级醇类(2,3-丁二醇、正丁醇、异丁醇、2-甲基-1-丁醇、1-丙醇、2-苯乙醇和异戊醇)、酮类(2,3-丁二酮)、醛类(乙醛和2-甲基-丁烯醛)、酯类(己酸乙酯、乙酸异戊酯、乙酸正丁酯、乙酸丙酯和乙酸乙酯)、萜类(α-萜醇、芳樟醇、香叶醇、β-香茅醇)[22]。与之密切相关的优势微生物主要是乳酸菌和酵母菌,部分除乳酸菌外的其他细菌和丝状真菌也会对咖啡风味产生一定影响。
乳酸菌是咖啡发酵过程中的关键菌群,能通过柠檬酸代谢和氨基酸代谢分解果胶和其他化合物[23−24]。在特定条件下,乳酸菌经代偿途径利用丙酮酸形成挥发性芳香化合物,如乙酸乙酯、异丁酸乙酯和乙醛等,有助于花香、果香和黄油香气形成[8]。Zhao等[25]从发酵咖啡豆中分离得到蒙氏肠球菌(Enterococcus mundtii),通过共接种发酵试验,发现有机酸(乙酸、柠檬酸、苹果酸、乳酸和奎宁酸)、酯类、醇类和酮类物质显著变化(P<0.05),有助于提高咖啡豆的香气和酸感。Wang等[26]采用鼠李糖乳酪杆菌(Lacticaseibacillus rhamnosus)作为发酵剂并补充1%(w/w)葡萄糖进行咖啡发酵,所得到的烘焙咖啡乳酸和糠醛含量更高,焦糖味更明显。de Jesus等[27]在接种肠膜明串珠菌发酵的咖啡中检测到十八烷酸、苯乙酸、苯甲酸、2-羟基乙酯,使焙烤咖啡具有焦糖、水果和香料等特征香气,并带来高酸度的差异化感官特征。近年来,植物乳植杆菌(Lactiplantibacillus plantarum)是咖啡发酵工艺中研究最多的产香乳酸菌。de Melo等[28]首次将植物乳植杆菌作为发酵剂用于农场湿法加工,缩短了咖啡发酵周期,提高了乙酸乙酯、乙醛和异丁酸乙酯等挥发性物质的含量,带来香蕉、橙子和菠萝特征风味。de Carvalho等[29]采用搅拌槽生物反应器发酵,通过接种植物乳植杆菌显著提高了烘焙咖啡的挥发性芳香化合物含量,如乙酸丁酯、乙酸乙酯、1-辛醇、乙酸异戊酯、己酸乙酯、琥珀酸乙酯和辛酸乙酯;与自然发酵相比,其香气、风味、酸度、体度和平衡性得分更高。叶宇娴[30]在接种发酵咖啡豆中微生物和风味的相关性研究中发现,植物乳植杆菌提高了特征性芳香醛类和酯类物质的含量,表现出柔和的水果和奶油香气;肠膜明串珠菌促进了酸类、醛类、酯类和吡咯类物质产生,使咖啡豆兼具典型的焙烤香气和丰富的花香、果香。由于乳酸菌可以在低氧环境下生长,产生有机酸和挥发性物质,为咖啡饮料带来酸味感官特征和果香类风味。因此,在咖啡初加工中接种乳酸菌发酵是咖啡风味调节的新策略,可以为咖啡饮料创造新的香气特征。
酵母菌是咖啡豆中最常分离的微生物之一,在发酵过程中对恶劣环境和应激条件有较强的适应能力[31]。酵母菌群对于优化咖啡的发酵性能至关重要,不仅能分解果胶,还能利用果肉中的糖类进行酒精发酵,产生乙醇、二氧化碳和大量次级代谢产物[32−33]。咖啡发酵过程中常见的酵母菌及其特征呈香物质及香味描述如表2所示。苯乙醇和乙酸异戊酯是酵母菌在代谢过程中产生的典型化合物,赋予咖啡花香和果香特征;乙醛作为酵母代谢乙醇的中间产物,能为咖啡带来杏仁味、果香和甜味;而高级醇类(如2-苯乙醇、异戊醇和糠醇)则显著影响咖啡香气的复杂度和层次感[34]。
葡萄汁有孢汉逊酵母(Hanseniaspora uvarum)因具有高产乙酸酯(果味)的能力,常被用作发酵剂以提升咖啡饮料的感官品质[35]。Elhalis等[36]研究证实了葡萄汁有孢汉逊酵母和库德里阿兹威毕赤酵母(Pichia kudriavzevii)复合发酵有利于产生乙酸乙酯和乙酸异戊酯,赋予咖啡香蕉、梨和苹果等果香风味品质。此外,酿酒酵母也是咖啡加工中广泛应用的发酵菌株,在厌氧发酵条件下主要产生乙醇,但在半厌氧和好氧条件下,能够产生琥珀酸、乳酸、葡萄糖酸盐、1,2-丙二醇、异丁醇、2,3-丁二醇[37]。黄梅等[38]研究表明,接种酿酒酵母发酵的烘焙咖啡豆中3-乙基-2,5-二甲基吡嗪、2,6-二甲基吡嗪、2-乙酰基吡咯、2,5-二甲基吡嗪香气活度值最高,对咖啡香气体系的贡献作用较大,且不同酿酒酵母所接种的烘焙豆之间挥发性物质存在差异。Vaz等[39]通过接种酿酒酵母发酵提高了阿拉比卡咖啡的理化特性和感官品质,在发酵过程中果糖、蔗糖、葡萄糖含量减少,乙酸、丙酸、琥珀酸和乳酸等有机酸显著变化。Martins等[40]研究表明酿酒酵母、德尔布有孢圆酵母(Torulaspora delbrueckii)和毕赤酵母菌混合发酵有利于产生乙酸乙酯、乙酸异戊酯、乙醇和乙醛等挥发性风味物质,增加了咖啡的杯测得分。Rocha等[41]添加酿酒酵母进行咖啡自诱导厌氧发酵(SIAF),提升了琥珀酸二乙酯、乙酸乙酯、1-辛醇、2-甲基-1-丁醇等挥发性风味物质,使咖啡具有焦糖、坚果特征风味,将杯测分数增加到84.50。赵林芬等[42]利用酿酒酵母在低温厌氧条件下发酵咖啡鲜果,将醛、醇、酯类挥发性成分提升至60%以上,赋予云南咖啡的花果甜香特质。综上,将单一或复配的特定酵母发酵剂用于咖啡初加工具有一定的商业潜力,不仅可以缩短发酵周期,提高产品的稳定性,还可获得更高的杯测分数和更明显的风味特征。
除乳酸菌外,从不同发酵方式处理的咖啡中分离出的部分其他细菌也会对风味产生重要影响。这些细菌在发酵前期高速生长,产生大量的有机酸(苹果酸、柠檬酸、乳酸、乙酸、琥珀酸和草酸等),从而降低pH[51]。有机酸有助于形成咖啡的酸味感官特征,但若过度发酵则会影响咖啡饮料的质量,尤其是过量丁酸和丙酸带来的不良风味[52]。最常见的是肠杆菌科细菌和醋酸菌的产酸发酵,这是加工过程中出现咖啡豆异味的主要原因[53]。但通过筛选和培养高产酯类或有机酸的菌株,也可以实现优化风味特性提高咖啡感官质量的目的。Janne等[54]报道,接种解淀粉芽孢杆菌(Bacillus amyloliquefaciens)发酵咖啡的乳酸、乙酸、乙醇含量较高,呈现出明显的酸味和坚果、可可风味。魏敏等[55]研究了接种桑肠杆菌(Enterobacter mori)对咖啡挥发性成分的影响,结果表明发酵咖啡中的坚果、奶香等香气显著提升,新生成异戊醛、3-甲基丁酸、2-甲基丁酸、苯甲醇、香兰素等17种挥发性成分,同时刺激性成分如苯酚类、邻苯二甲酸二丁酯等含量降低。Martinez等[56]使用枯草芽孢杆菌(Bacillus subtilis)和分散泛菌(Pantoea dispersa)进行湿法发酵,促进了苹果酸、乳酸、琥珀酸和乙酸等有机酸的产生。有研究表明,在湿法发酵中接种地衣芽孢杆菌(Bacillus licheniformis)对提高咖啡品质有显著影响,可产生5-甲基糠醛、2-氧乙酸丁酯、1-糠基吡咯和麦芽醇酚等挥发性化合物,带来杏仁、焦糖和巧克力风味,并提高咖啡杯测评分[57]。因此,通过定向接种功能细菌菌株,可有效优化有机酸与挥发性化合物的组成,从而提升咖啡的感官品质。
目前,对丝状真菌的研究主要集中在对咖啡质量带来的不利影响,比如部分曲霉、青霉和镰刀菌次生代谢产生的毒素与异味[58]。尽管如此,丝状真菌因具有产多种胞外酶、对水分要求低等优点,能有效转化咖啡豆的前体物质,而在咖啡固态发酵中受到关注。Lee等[59]研究表明少孢根霉(Rhizopus oligosporus)可以改变咖啡生豆香气前体物质的组成,使得脯氨酸和天冬氨酸的浓度增加1.5倍,阿魏酸和咖啡酸的降解使得挥发性酚衍生物的总浓度增加2倍,在烘焙过程中产生咖啡特有的挥发性物质(吡嗪类、硫醇类、呋喃酮类和愈创木酚类)。Tang等[60]接种曲霉属和毛霉属(Mucor)对咖啡豆进行固态发酵提高了游离糖、游离氨基酸和多酚类化合物(如咖啡酰奎宁酸)的含量。
综上所述,微生物在塑造咖啡风味方面具有突出的优势,不同微生物发酵可以赋予咖啡豆不同的风味特征。对咖啡香气的定向调控,应更多关注不同菌种与风味物质的形成关系,利用多菌种混合发酵赋予咖啡更加均衡、独特的口感,以此来提高精品咖啡的特征风味。
在咖啡发酵过程中,微生物代谢可以增加咖啡香气和风味化合物的多样性[61]。酵母菌、细菌、丝状真菌利用咖啡果多糖通过糖酵解途径(EMP)、三羧酸循环(TCA)等产生酒精、甘露醇、甘油和酸;通过蛋白质水解、氨基酸代谢形成杂醇类、醛类、含硫化合物;通过脂肪酸的生物合成/分解,进一步形成次生代谢产物,如酯、甲基酮、醇和内酯[62]。这些化合物迁移并保留在咖啡豆中,影响最终风味[63]。据报道,在咖啡生豆中共检测到170多种挥发性化合物,主要包括26%的醇类,23%的酯类,14%的醛类,7%的萜烯/萜类,7%的呋喃/呋喃酮,这些挥发性化合物中有70%是在发酵过程中发现的[64]。它们赋予了咖啡饮料丰富的感官特性:呋喃(甜味)、吡嗪(榛子味)、醛类(水果味)、酮类(焦糖和黄油味)、酯类(水果味)、醇类(花香和葡萄酒味)、吡啶类(巧克力味)[65]。因此,微生物的代谢反应决定了糖、蛋白质、脂肪等大分子物质水解产生相应的挥发性风味物质、有机酸以及氨基酸等一系列风味物质的浓度,使咖啡豆在后续的烘焙过程中出现差异化以形成丰富各异的风味。在咖啡发酵过程中,微生物参与风味物质形成的相关代谢途径包括糖代谢、蛋白质与氨基酸代谢及脂肪代谢。
咖啡果中存在丰富的碳水化合物,包括戊糖(木糖、核糖和阿拉伯糖)、己糖(葡萄糖、果糖、半乳糖和甘露糖)和多糖(果胶)等,是微生物利用的主要碳源[30]。在咖啡发酵过程中,约60%的糖被细菌和酵母利用,从而产生大量的乙醇、乙酸和乳酸[62]。微生物将糖类物质转化为风味物质的代谢路径如图1所示。芽孢杆菌属、毕赤酵母属、乳杆菌属等能够产生不同的水解酶,如果胶甲酯酶、果胶裂解酶和多聚半乳糖醛酸酶,与咖啡内源酶共同作用促进复杂碳水化合物的分解,释放单糖(即葡萄糖、果糖、鼠李糖、L-阿拉伯糖)作为进一步代谢和香气物质形成的额外碳源[66−67]。de Jesus等[68]进行SIAF发酵结果表明接种肠膜明串珠菌、植物乳植杆菌和酿酒酵母导致了咖啡鲜果中葡萄糖(98.6%)、果糖(97.6%)和蔗糖(100%)的大量消耗和乳酸、乙酸的积累,由乳酸菌代谢的2,3-丁二醇对咖啡饮料的芳香特性产生显著影响。
咖啡发酵前期,以明串珠菌属、葡糖杆菌属为代表的细菌通过EMP途径将葡萄糖转化为1,6-二磷酸果糖,在醛缩酶的作用下裂解成两个三碳化合物,进而转化为两个丙酮酸分子;或者通过磷酸戊糖途径(HMP)生成6-磷酸葡萄糖酸后,被脱氢酶继续降解为磷酸戊糖,最终也转化为丙酮酸[69−70]。一方面,丙酮酸分子在相关酶系的作用下,生成乙醛、乙醇、乙酸等典型的代谢风味物质,或进入三羧酸循环(TCA)产生琥珀酸、柠檬酸、苹果酸等有机酸[71]。这些有机酸浓度高且不易挥发,在干燥过程中得以保存并影响咖啡饮料的酸感。比如,琥珀酸是一种由酵母菌、异型发酵乳酸菌和芽孢杆菌产生的有机酸,通过乙醛酸盐和TCA代谢产生,广泛存在于咖啡饮料中[57]。另一方面,在酿酒酵母细胞内,丙酮酸可通过α-乙酰乳酸的化学驱动脱羧反应产生二酮类化合物,其中2,3-丁二酮的浓度与奶油香气的强度呈显著正相关[72]。对于乳酸菌而言,同型发酵乳酸菌(如植物乳植杆菌)可以通过HMP将咖啡果肉中的己糖转化为乳酸(转化率>85%),并产生乙酸盐、乙醇、二乙酰和乙醛等其他风味代谢物[73]。异型发酵乳酸菌(如肠膜明串珠菌)则降解丙酮酸产生乳酸、乙醇以及多种短链脂肪酸[74]。此外,一些在咖啡发酵过程中分离的酵母菌种,如酿酒酵母、有孢圆酵母和汉逊酵母,能够利用3-磷酸甘油醛通过甲羟戊酸途径(MEP)产生萜类衍生物[75]。萜类化合物包括β-香叶醇、芳樟醇、α-萜品醇和γ-萜品烯,主要源自于糖苷前体,通过酵母菌产生的β-葡萄糖苷酶的作用释放[76]。Silva等[77]研究表明在酿酒酵母和毕赤酵母发酵咖啡中检测到芳樟醇,赋予了咖啡饮料柑橘和木质香气。
微生物的活性和发酵程度决定了游离氨基酸的浓度,低分子肽和游离氨基酸本身具有一定的滋味,同时也是重要的风味前体物质,构成了最终产品的基本味道[62]。咖啡发酵过程中微生物将蛋白质和氨基酸类转化为多种风味物质的代谢路径如图2所示。以风味著称的麝香猫咖啡通过利用胃肠道微生物水解咖啡果实中的蛋白质,造成氨基酸组成的变化进而对咖啡香气产生显著影响,同时,使发酵后苦味和酸度偏低[78]。咖啡果肉富含氨基酸(亮氨酸、缬氨酸、苯丙氨酸、苏氨酸和异亮氨酸),这些氨基酸可被艾利希途径(Ehrlich)同化[52]。游离氨基酸在微生物酶(转氨酶、脱羧酶、脱氢酶和裂解酶)的协同作用下进一步产生醛类、羧酸类、醇类和羟基短链脂肪酸类物质[79]。酵母菌在发酵过程中通过Ehrlich途径分解氨基酸,α-酮戊二酸在其中作为关键的氨基受体,可将支链氨基酸和芳香族氨基酸转化为相应的高级醇,例如正丁醇、异丁醇、1-丙醇、2-苯乙醇、异戊醇、2,3-丁二醇和2-甲基-1-丁醇[80−81]。高级醇在酿酒酵母细胞中通过酶促反应与乙酰-CoA酯化,生成具有高香气活性的乙酸异戊酯和乙酸苯乙酯等酯类,进而提供果香和葡萄酒香特征,因此咖啡豆中芳香酯的生成与氨基酸的减少呈正相关[82]。Lee等[59]研究了少孢根霉发酵对咖啡豆氨基酸的影响,结果表明发酵后的咖啡豆总游离氨基酸浓度增加了3~10倍,苯丙氨酸通过Ehrlich途径生物转化为2-苯乙醇,使得发酵后的咖啡豆中2-苯乙醇的含量相对较高;缬氨酸生物合成途径的中间体α-乙酰乳酸通过氧化脱羧反应生成双乙酰,进而还原为2,3-丁二醇,使得发酵后的咖啡豆中2,3-丁二醇浓度显著提高。
与真菌不同,由于乳酸菌缺乏谷氨酸脱氢酶,无法生成α-酮戊二酸,因此,在咖啡豆发酵过程中,乳酸菌的转氨反应及后续香气物质的生成主要依赖于培养基质中的α-酮戊二酸[81]。植物乳杆菌能够在苯丙氨酸分解代谢过程中生成苯乙醛、苯乙酸和苯乙醇,而乳酸乳球菌则能够将亮氨酸代谢为3-甲基丁醛和3-甲基丁酸,这些代谢产物赋予咖啡独特的香气特征,如麦芽香、花香、奶酪或水果味[73]。鼠李糖乳酪杆菌(Lacticaseibacillus rhamnosus)促进了芳香族氨基酸的分解代谢,使得对羟基苯基乳酸和3-苯基乳酸浓度升高,为咖啡带来花香、蜂蜜和焦糖风味[83]。由于两种氨基酮的缩合可以产生吡嗪衍生物,因此接种发酵咖啡豆中吡嗪的含量和类型受到氮源类型(游离氨或与氮相连的氨基酸)的影响[52]。研究证明,以2,3-二甲基吡嗪为代表的吡嗪类物质均为自然发酵和微生物强化发酵咖啡的关键挥发性成分的标志物,接种植物乳植杆菌发酵咖啡中的吡嗪类含量是对照组的3.14倍,赋予咖啡坚果香、烘焙香以及甜香[84−85]。
咖啡豆的脂质部分主要由甘油三酯、甾醇、生育酚和二萜组成,脂肪酸主要以结合态存在,其中大部分与甘油结合形成甘油三酯;约20%与二萜类物质结合,小部分存在于甾醇中[52]。咖啡发酵过程中微生物将脂肪转化为多种风味物质的代谢路径如图3所示。乳球菌和乳杆菌产生的脂酶含量相对较高,酯酶将脂肪水解为游离脂肪酸,再通过氧化反应产生醛、醇、酮和酯类挥发性化合物[86]。据报道,少孢根霉在固态发酵条件下也有产脂肪酶的能力,且能够通过直接酯化反应催化酯类合成[59]。在酵母菌、乳酸菌细胞内,饱和脂肪酸通过β-氧化使β-酮酸脱羧产生甲基酮,甲基酮可被还原酶还原为相应的次级醇;不饱和脂肪酸在自由基存在下通过β-氧化形成氢过氧化物、醛类和酯类[87]。此外,脂肪酸可直接形成4-/5-羟基酸,这些酸环化为内酯类化合物,从而提供浓郁的果味[88]。研究显示在咖啡豆中接种异常威克汉姆酵母(Wickerhamomyces anomalus)和解脂耶氏酵母(Yarrowia lipolytica)进行发酵,分别可产生高于对照组(未接种)8倍和5倍的丁内酯,并带来显著的黄油/椰子风味[47]。
咖啡的发酵时间越长,产生酯类物质的种类越丰富。咖啡中常见的酯类化合物包括乙酸酯、脂肪酸甲酯和脂肪酸乙酯等,与发酵初期乳酸菌和酵母菌的生长密切相关,其合成途径主要包括酯化、醇解、酸解和酯交换等反应[89]。例如,由酿酒酵母产生的癸酸乙酯是一种由癸酸和乙醇形成的挥发性酵母代谢产物,可以促进花香和果味[90]。由少孢根霉产生的棕榈酸乙酯具有水果味特性,可通过含有棕榈酸的甘油三酯与乙醇的酯交换反应生成[59]。Chin等[91]研究表明库德里阿兹威毕赤酵母和乳酸乳球菌共发酵咖啡豆提高了异戊醇和苯乙醇含量,可作为芳香酯的前体;酿酒酵母、汉逊酵母、库德里阿兹威毕赤酵母共发酵可显著提高脂肪酸甲酯、乙酯以及乙酸酯的浓度,为咖啡饮料带来水果和花香,其中,库德里阿兹威毕赤酵母具有高效将异戊醇转化为乙酸异戊酯(香蕉味)的能力。
随着中国咖啡市场的扩大,人们对咖啡的种类和品质有了更高的要求,生产过程中仍出现精品率低、风味不稳定、安全性不高等问题。因此,本文系统阐述了咖啡发酵过程中的微生物组成及代谢机制,明确微生物与特征风味物质的关系,对咖啡鲜果加工后期风味的定向调控具有重要意义。基于研究现状,今后对咖啡发酵过程中的微生物和风味研究可以围绕以下几方面开展:a.以目标风味为导向,可通过数据分析挖掘咖啡发酵过程中的功能微生物资源,了解菌株间的互作机理,筛选纯种发酵剂或复配强化微生物群用于发酵接种以调控香气和滋味的形成;b.结合人工智能和多组学联用开发预测模型,将组合微生物与代谢特征、特定的感官特征相关联,从而精确调控发酵过程以优化感官质量;c.开展环境因子对微生物多样性以及咖啡风味形成的研究,以便在生产过程中精准化调控,提高咖啡的品质、风味稳定性和缩短生产周期,为促进精品咖啡的标准化生产和产业化发展提供基础。

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2026年第47卷第3期
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doi: 10.13386/j.issn1002-0306.2025010133
  • 接收时间:2025-01-14
  • 首发时间:2026-10-25
  • 出版时间:2026-02-01
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  • 收稿日期:2025-01-14
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    1云南农业大学食品科学技术学院,云南昆明 650201
    2桂林师范学院化学与生命健康学院,广西桂林 541199

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李宏(1974−),男,博士,研究员,研究方向:农产品加工,E-mail:。
胡永金(1972−)(ORCID:0000−0001−5149−0079),男,博士,教授,研究方向:食品微生物资源与利用,E-mail:。
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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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