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Article(id=1320175819566207982, tenantId=1146029695717560320, journalId=1260987677001138203, issueId=1320175721343987874, articleNumber=null, orderNo=null, doi=10.13386/j.issn1002-0306.2025050095, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1747152000000, receivedDateStr=2025-05-14, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1792684043064, onlineDateStr=2026-10-22, pubDate=1781452800000, pubDateStr=2026-06-15, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1792684043064, onlineIssueDateStr=2026-10-22, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1792684043064, creator=13701087609, updateTime=1792684043064, updator=13701087609, issue=Issue{id=1320175721343987874, tenantId=1146029695717560320, journalId=1260987677001138203, year='2026', volume='47', issue='12', pageStart='1', pageEnd='517', issueExtLink='null', onlineDate='null', pubDate='1781452800000', pubDateStr='2026-06-15', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1792684019647, creator='13701087609', updateTime=1792684019647, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext=null, issueFiles=null, downloadFileDto=null}, startPage=156, endPage=166, ext={EN=ArticleExt(id=1320175821088740335, articleId=1320175819566207982, tenantId=1146029695717560320, journalId=1260987677001138203, language=EN, title=Analysis of Bacterial Flora Structure and Flavor of Traditional Fermented Milk from Different Regions of Tibet, columnId=1261336275769016441, journalTitle=Science and Technology of Food Industry, columnName=Bioengineering, runingTitle=null, highlight=null, articleAbstract=

To investigate the microbial community structure and flavor quality of traditional Tibetan fermented milk, high-throughput sequencing and headspace solid-phase microextraction-gas chromatography-mass spectrometry (HS-SPME–GC–MS) were used to analyze the microbial diversity and volatile flavor components in traditional Tibetan fermented milk. The types and contents of organic acids were determined by liquid chromatography. The results showed that the main dominant bacterial flora in traditional Tibetan fermented milk products were Lactobacillus, Streptococcus, and Lactococcus, etc., while the main dominant fungal flora were Saccharomyces, Kluyveromyces, and Geotrichum, etc. Six organic acids were identified, including acetic acid, citric acid, lactic acid and so on, among which, lactic acid was the most abundant (6989.02 mg/L), followed by acetic acid (1047.25 mg/L). A total of 35 volatile flavor compounds were detected in the fermented milk, with esters accounting for the highest proportion (43.98%), followed by alcohols (28.98%) and acids (16.00%). The composition and concentration of volatile flavor compounds in fermented milk exhibited notable regional variations. Fermented milk from Shigatse was distinguished by pronounced fruity aromas, particularly pineapple and banana, complemented by creamy notes, which together contributed to a rich and complex flavor profile. By contrast, fermented milk from Nagqu was characterized by strong creamy aromas, nutty undertones, and a balanced sweet–sour taste, whereas fermented milk from Ali demonstrated a predominance of more intense fruity aromas. Spearman’s correlation analysis based on key odor–active compounds (odor activity value, OAV>1) revealed distinct associations between microbial taxa and flavor compounds. The bacterial genera most strongly correlated with key flavor compounds were primarily Lactobacillus, Acetobacter and Leuconostoc, while the fungal genera most closely linked to flavor development included Apiotrichum, Dipodascaceae and Pichia. This study elucidated the correlation between microbial community structure and the formation of aroma compounds in traditional Tibetan fermented milk, providing a theoretical foundation for the standardized production, targeted flavor regulation, and development of functional strains for traditional dairy products.

, authors=Min TIAN1, 2, 3, Jing ZHANG2, 3, Jiajun YU2, 3, 4, Yuhong ZHANG4, Huaying WEN4, Jumei LI2, 3, Jie XUE2, 3, *, Yanlin LIU1, *, authorsList=Min TIAN, Jing ZHANG, Jiajun YU, Yuhong ZHANG, Huaying WEN, Jumei LI, Jie XUE, Yanlin LIU, authorCompany=null, correspAuthors=Jie XUE, Yanlin LIU, 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=1320175822426723322, articleId=1320175819566207982, tenantId=1146029695717560320, journalId=1260987677001138203, language=CN, title=西藏不同地区传统发酵乳的菌群结构及其风味分析, columnId=1261336277547401341, journalTitle=食品工业科技, columnName=生物工程, runingTitle=null, highlight=null, articleAbstract=

为了探究西藏传统发酵乳中的微生物体系及其风味品质，利用高通量测序技术以及顶空固相微萃取-气相色谱-质谱联用技术，对西藏传统发酵乳中的微生物多样性及挥发性风味成分进行分析，并通过液相色谱法测定其有机酸的种类及含量。结果表明：西藏传统发酵乳制品中的主要优势细菌菌群为乳杆菌属（Lactobacillus）、链球菌属（Streptococcus）、乳球菌属（Lactococcus）等，主要优势真菌菌群为酵母菌属（Saccharomyces）、克鲁维酵母属（Kluyveromyces）、地霉属（Geotrichum）等。该乳制品中含乳酸、乙酸、柠檬酸等6种有机酸，其中乳酸含量最高，最高可达6989.02 mg/L；其次是乙酸，达1047.25 mg/L。该乳制品中共检测出35种挥发性风味物质，其中酯类物质相对含量占比最高（43.98%），其次是醇类（28.98%）和酸类物质（16.00%）。不同地区发酵乳中的挥发性风味物质种类和含量各有差异，其中日喀则市的发酵乳以菠萝和香蕉等果香和奶油香气为主，其风味较为丰富复杂；那曲地区的发酵乳以奶油香气、坚果香气和酸甜味为主；阿里地区的发酵乳水果香气较为明显。通过Spearman相关系数与OAV>1的特征风味物质进行相关性分析，与风味物质相关性较高的细菌菌属主要是Lactobacillus、Acetobacter、Leuconostoc；与风味物质相关性较高的真菌菌属主要是Apiotrichum、Dipodascaceae、Pichia。本研究阐明了西藏传统发酵乳中菌群结构与其风味物质形成的相关性，为传统乳制品的标准化生产、风味定向调控及功能性菌株开发提供理论依据。

, authors=田敏1, 2, 3, 张静2, 3, 于佳俊2, 3, 4, 张玉红4, 文华英4, 李菊梅2, 3, 薛洁2, 3, *, 刘延琳1, *, authorsList=田敏, 张静, 于佳俊, 张玉红, 文华英, 李菊梅, 薛洁, 刘延琳, authorCompany=null, correspAuthors=薛洁, 刘延琳, authorNote=

田敏（2001−），女，硕士研究生，研究方向：发酵工程，E-mail：

, correspAuthorsNote=
薛洁（1974−），女，博士，教授级高级工程师，研究方向：发酵工程，E-mail：。
刘延琳（1966−），女，博士，教授，研究方向：酿酒微生物，E-mail：。
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2China National Research Institute of Food and Fermentation Industries, Beijing 100125, China
3International Joint Research Center of Quality and Safety of Alcoholic Beverages, Beijing 100125, China, bio=null, bioImg=null, bioContent=null, aboutCorrespAuthor=null), CN=AuthorExt(id=1320175823240417292, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, authorId=1320175823060062215, language=CN, stringName=田敏, firstName=null, middleName=null, lastName=null, prefix=null, suffix=null, authorComment=null, nameInitials=null, affiliation=null, department=null, xref=1, 2, 3, address=1西北农林科技大学葡萄酒学院，陕西杨凌 712100
2中国食品发酵工业研究院有限公司，北京 100125
3国家酒类品质与安全国际联合研究中心，北京 100125, bio={"content":"

田敏（2001−），女，硕士研究生，研究方向：发酵工程，E-mail：

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田敏（2001−），女，硕士研究生，研究方向：发酵工程，E-mail：

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Food Science, 2018, 39(19): 329−335.], articleTitle=null, refAbstract=null), Reference(id=1320175836494418055, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=36, rfOrder=49, authorNames=null, journalName=null, refType=null, unstructuredReference=CHEN Zhifei, CAI Lili, HAO Hui, et al. Progress in the application of odor activity values in the characterization of key aroma components in foods[J]. 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Journal of Agricultural and Food Chemistry, 2016, 64(24): 4990−4999., articleTitle=null, refAbstract=null), Reference(id=1320175836657995913, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=38, rfOrder=51, authorNames=null, journalName=null, refType=null, unstructuredReference=卢灏泽, 吕嘉伟, 杨帆, 等. 西藏牦牛奶酪的微生物群落结构与风味物质分析[J]. 食品与发酵工业, 2023, 49(6): 179−186. [LU Haoze, LYU Jiawei, YANG Fan, et al. Microbial community structure and flavor analysis of Tibetan yak cheese[J]. Food and Fermentation Industries, 2023, 49(6): 179−186.], articleTitle=null, refAbstract=null), Reference(id=1320175836750270602, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=38, rfOrder=52, authorNames=null, journalName=null, refType=null, unstructuredReference=LU Haoze, LYU Jiawei, YANG Fan, et al. Microbial community structure and flavor analysis of Tibetan yak cheese[J]. Food and Fermentation Industries, 2023, 49（6）: 179−186., articleTitle=null, refAbstract=null)], funds=null, companyList=[AuthorCompany(id=1320175822669992955, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, xref=1, ext=[AuthorCompanyExt(id=1320175822678381564, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822669992955, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1College of Enology, Northwest A&F University, Yangling 712100, China), AuthorCompanyExt(id=1320175822686770173, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822669992955, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1西北农林科技大学葡萄酒学院，陕西杨凌 712100)]), AuthorCompany(id=1320175822741296126, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, xref=2, ext=[AuthorCompanyExt(id=1320175822749684735, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822741296126, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2China National Research Institute of Food and Fermentation Industries, Beijing 100125, China), AuthorCompanyExt(id=1320175822758073344, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822741296126, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2中国食品发酵工业研究院有限公司，北京 100125)]), AuthorCompany(id=1320175822846152704, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, xref=3, ext=[AuthorCompanyExt(id=1320175822854541313, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822846152704, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3International Joint Research Center of Quality and Safety of Alcoholic Beverages, Beijing 100125, China), AuthorCompanyExt(id=1320175822867124226, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822846152704, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3国家酒类品质与安全国际联合研究中心，北京 100125)]), AuthorCompany(id=1320175822951010307, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, xref=4, ext=[AuthorCompanyExt(id=1320175822959398916, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822951010307, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=4Institute of Agricultural Product Development and Food Science, Tibet Academy of Agricultural and Animal Husbandry Sciences, Lhasa 850030, China), AuthorCompanyExt(id=1320175822971981829, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, companyId=1320175822951010307, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=4西藏自治区农牧科学院农产品开发与食品科学研究所，西藏拉萨 850030)])], figs=[ArticleFig(id=1320175827438915647, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Fig.1, caption=Relative composition percentages of volatile flavor compound categories in different samples, figureFileSmall=jLvfzX07MogyApzIORXsfA==, figureFileBig=GjYJ8dQsgyyS7y6IcdX+HQ==, tableContent=null), ArticleFig(id=1320175827526996032, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=图1, caption=不同样品中挥发性风味物质类别的相对组成比例

注：图中百分比表示各类挥发性风味物质峰面积占该样品总峰面积的比例，用于反映不同样品挥发性风味物质类别组成特征，不代表样品间各类别化合物的相对含量。

, figureFileSmall=jLvfzX07MogyApzIORXsfA==, figureFileBig=GjYJ8dQsgyyS7y6IcdX+HQ==, tableContent=null), ArticleFig(id=1320175827728322625, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Fig.2, caption=Protein content of yogurt samples, figureFileSmall=UJ5zQAwtb7cvk1BehATUvQ==, figureFileBig=29ODu++rGqHAtRjRhLZiPw==, tableContent=null), ArticleFig(id=1320175827787042882, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=图2, caption=酸奶样品的蛋白质含量

注：不同小写字母表示数据差异显著，P<0.05。

, figureFileSmall=UJ5zQAwtb7cvk1BehATUvQ==, figureFileBig=29ODu++rGqHAtRjRhLZiPw==, tableContent=null), ArticleFig(id=1320175827858346052, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Fig.3, caption=Relative abundance at genus level of yogurt samples, figureFileSmall=ELPZ0nK6qW9K4AtBOPRewA==, figureFileBig=ZkzfWk8dhf6Ms+EM2DsyyA==, tableContent=null), ArticleFig(id=1320175827938037828, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=图3, caption=酸奶样品属水平相对丰度

注：a细菌属水平群落结构；b真菌属水平群落结构。

, figureFileSmall=ELPZ0nK6qW9K4AtBOPRewA==, figureFileBig=ZkzfWk8dhf6Ms+EM2DsyyA==, tableContent=null), ArticleFig(id=1320175828009340997, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Fig.4, caption=PCoA analysis of yogurt samples, figureFileSmall=foXqE+51bXPyju2FckIfRQ==, figureFileBig=Vev3qXQAsqZavnXcF+rjbQ==, tableContent=null), ArticleFig(id=1320175828089032774, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=图4, caption=酸奶样品的PCoA分析

注：a细菌；b真菌。

, figureFileSmall=foXqE+51bXPyju2FckIfRQ==, figureFileBig=Vev3qXQAsqZavnXcF+rjbQ==, tableContent=null), ArticleFig(id=1320175829745782855, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Fig.5, caption=Heat map of correlation between microbial species and volatile flavor substances in yogurt samples, figureFileSmall=YOY802uq6Ce+jVsvmFnffw==, figureFileBig=AM1pY3qA/4EX2S2v0lJzNw==, tableContent=null), ArticleFig(id=1320175829812891720, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=图5, caption=酸奶样品中的微生物种类与挥发性风味物质相关性

注：a.细菌；b.真菌；*表示P<0.05，**表示P<0.01，***表示P<0.001。

, figureFileSmall=YOY802uq6Ce+jVsvmFnffw==, figureFileBig=AM1pY3qA/4EX2S2v0lJzNw==, tableContent=null), ArticleFig(id=1320175829913555017, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Table 1, caption=

Yoghurt sample information sheet

, figureFileSmall=null, figureFileBig=null, tableContent=
样品名称采样地点经纬度海拔
（m）		样品来源
L3那曲市班戈县90.01°E，31.40°N4681.5牦牛奶
L5那曲市班戈县门当乡4号桥89.59°E，31.40°N4564.2牦牛奶
L7那曲市申扎县布嘎村88.52°E，31.76°N4565绵羊奶
L10那曲市尼玛县阿索乡81.06°E，31.78°N4753绵羊奶
L14阿里地区革吉县雄巴乡82.65°E，32.23°N4412山羊奶
L18阿里地区革吉县
革吉镇森布村2组		81.31°E，32.18°N4620山羊奶
L20日喀则萨嘎县达琼村85.57°E，29.04°N4860牦牛奶
L23日喀则吉隆县折巴乡恰门巴村85.43°E，28.95°N4678牦牛奶
L25日喀则吉隆县吉隆镇如嘎自然村85.24°E，28.45°N3012牦牛奶
L28日喀则聂拉木县门布乡孔措村85.84°E，28.67°N4563牦牛奶
L30日喀则定日县协格尔镇罗罗辖镇87.21°E，28.72°N4425牦牛奶
L31日喀则拉孜县查务乡吉角村87.46°E，28.98°N4850牦牛奶
L32日喀则萨迦县冲达村88.47°E，29.17°N4346牦牛奶
), ArticleFig(id=1320175829984858186, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=表1, caption=

酸奶样品信息

, figureFileSmall=null, figureFileBig=null, tableContent=
样品名称采样地点经纬度海拔
（m）		样品来源
L3那曲市班戈县90.01°E，31.40°N4681.5牦牛奶
L5那曲市班戈县门当乡4号桥89.59°E，31.40°N4564.2牦牛奶
L7那曲市申扎县布嘎村88.52°E，31.76°N4565绵羊奶
L10那曲市尼玛县阿索乡81.06°E，31.78°N4753绵羊奶
L14阿里地区革吉县雄巴乡82.65°E，32.23°N4412山羊奶
L18阿里地区革吉县
革吉镇森布村2组		81.31°E，32.18°N4620山羊奶
L20日喀则萨嘎县达琼村85.57°E，29.04°N4860牦牛奶
L23日喀则吉隆县折巴乡恰门巴村85.43°E，28.95°N4678牦牛奶
L25日喀则吉隆县吉隆镇如嘎自然村85.24°E，28.45°N3012牦牛奶
L28日喀则聂拉木县门布乡孔措村85.84°E，28.67°N4563牦牛奶
L30日喀则定日县协格尔镇罗罗辖镇87.21°E，28.72°N4425牦牛奶
L31日喀则拉孜县查务乡吉角村87.46°E，28.98°N4850牦牛奶
L32日喀则萨迦县冲达村88.47°E，29.17°N4346牦牛奶
), ArticleFig(id=1320175830064549963, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Table 2, caption=

Volatile flavour components of yoghurt (μg/kg)

, figureFileSmall=null, figureFileBig=null, tableContent=
类别风味物质保留时间（min）L3L5L7L10L14L18L20L23L25L28L30L31L32香气特征[15–16]
酯类丁酸乙酯7.36783.30±5.63NDNDNDNDNDND93.76±3.97344.72±17.65NDNDND121.17±6.28菠萝、香蕉和苹果香
庚酸乙酯17.2596.42±0.35NDNDNDNDNDND27.60±1.05147.54±10.88NDNDNDND菠萝香
辛酸乙酯26.76990.90±52.55342.16±26.833520.62±189.00765.49±26.16868.43±53.76783.91±58.512066.32±145.90831.11±63.2810044.03±467.67718.21±35.18147.43±10.091802.86±60.36ND脂香味
月桂酸乙酯40.4161.44±0.053.31±0.18NDNDNDNDNDND139.61±6.1712.44±0.43NDNDND果香和花香
癸酸乙酯34.774111.65±7.3160.14±4.631001.59±59.9681.70±3.15318.60±11.0647.93±2.66191.19±5.76194.13±10.682633.64±127.28191.36±14.5816.64±1.1035.58±1.78ND椰子香
己酸乙酯18.254507.24±31.16ND676.25±39.28ND66.53±5.26ND41.62±1.74666.28±27.926657.53±527.00128.44±5.9640.23±1.28894.75±65.46ND菠萝香
乙酸乙酯3.8441446.18±49.70NDND832.48±26.62968.14±54.211004.14±61.788807.02±455.931155.72±81.973730.71±248.231394.10±44.012984.88±195.077906.71±504.67ND橙子味
乳酸异戊酯29.9585.93±0.44NDNDNDNDND116.85±8.2916.26±0.74130.95±5.68ND53.21±3.04156.76±11.93ND刺激味
棕榈酸乙酯45.553NDND85.43±6.3911.90±0.6411.31±0.376.45±0.508.43±0.40ND37.10±1.86NDNDNDND蜡香、奶油香
9-癸烯酸乙酯33.2918.43±1.233.74±0.15NDNDNDNDND24.66±1.91285.49±21.27NDNDNDND果香、蜡香、脂香
L-乳酸乙酯19.73195.83±3.7971.59±2.831305.07±66.83NDNDND2364.86±152.12440.17±28.631658.33±123.40ND457.20±23.85898.38±54.03929.50±62.60果香、奶油香
甲酸乙烯酯2.645NDNDNDNDNDNDNDNDND374.28±22.36ND2359.08±137.88ND刺激味
丁位癸内酯44.4421.85±0.061.13±0.05NDNDNDNDND1.15±0.05NDNDNDNDND椰子香、奶油香
乙酸戊酯10.012241.08±10.6698.07±5.62262.62±17.66168.36±8.21253.34±8.92226.19±12.55536.23±26.64166.52±8.49576.60±37.33ND76.91±5.07848.27±37.92ND香蕉味
酸类丁酸30.86ND21.20±1.46NDNDNDNDND21.21±0.751200.03±62.32129.37±5.1668.85±3.06193.39±15.20511.37±25.79刺激味
辛酸42.108NDND581.78±32.08165.32±5.53103.43±3.89281.90±19.29115.11±6.53ND3130.22±249.72160.90±9.49NDNDND焦糖香味
己酸37.66917.18±0.9736.37±1.88669.38±48.42155.30±7.46ND401.11±30.56327.78±19.2858.42±4.138127.88±452.85646.78±20.91832.41±44.04832.85±30.89727.09±36.85甜香味
正癸酸44.786NDND34.82±2.3612.09±0.80ND18.11±0.69NDND320.92±11.2547.24±1.5718.63±1.10NDND奶香味
2-甲基己酸32.454ND35.61±1.49NDNDNDNDND46.63±1.50NDNDND126.96±8.14152.47±6.48水果香
棕榈酸54.4683.73±0.166.05±0.19NDNDNDNDND9.75±0.66NDNDNDNDND微弱油脂香、蜡香
醋酸23.688ND212.25±6.52541.37±37.1962.48±2.63122.62±3.8289.35±3.53219.17±11.36147.52±7.66412.88±17.29575.55±44.161423.08±48.031200.01±88.853489.31±276.58刺激性气味
苯甲酸46.589NDND650.95±21.3668.39±5.44ND226.84±7.2776.03±4.7112.46±0.54ND535.77±21.49NDNDND苦杏仁味
丙酸27.392NDNDNDNDNDNDNDNDNDNDNDND56.28±1.78刺激味
环己甲酸42.369NDNDNDNDNDNDNDNDNDND18.31±0.6377.88±4.26ND山楂香
醇类异戊醇14.0661416.71±113.221060.64±48.743466.50±163.73846.72±67.032166.35±106.452146.33±78.375179.61±330.371531.43±56.603928.38±167.582505.03±189.911539.04±74.416484.65±239.814059.69±257.59苦杏仁味
丁二醇27.81ND590.12±33.26NDNDNDNDND143.83±6.7030.24±2.09ND536.96±26.20546.31±24.232136.51±128.50奶油香、果香
3-甲硫基丙醇33.99315.34±0.676.38±0.31NDNDND51.55±2.80ND9.55±0.67ND109.90±7.04ND72.60±5.40ND硫醇香气
β-苯乙醇39.264301.43±20.64167.99±8.70847.89±34.57440.51±24.20458.69±16.45964.28±33.341325.65±48.45540.61±38.681039.81±58.562516.07±91.74684.58±39.095045.64±391.75450.27±21.06蜜香玫瑰香
酮类2-庚酮12.64320.61±1.23NDND16.90±0.52ND94.84±2.90NDNDNDND81.92±4.08NDND梨子香
3-羟基-2-丁酮16.748ND18.35±0.72NDNDND13.44±0.549127.21±576.7225.59±1.93NDND146.49±9.941846.11±91.44ND奶油香
2-壬酮21.38217.80±1.40NDNDNDNDNDND14.39±0.83333.72±12.16NDND0.00±0.00ND果香味
其他苯乙烯16.09767.28±4.2954.94±4.032215.63±91.251078.52±46.621312.33±49.321173.00±87.671065.09±34.79190.57±10.28NDND52.85±2.96746.62±39.64125.59±7.91树脂香、花香
十二烷11.70934.54±1.1457.28±3.84NDNDNDNDND40.94±2.19426.15±27.04NDNDNDND微弱烃类气味
异丁烷9.44130.43±9.04NDNDNDNDND262.54±19.91ND277.99±11.77NDNDNDND微弱石油样气味
2,3,5-三甲基吡嗪19.98713.43±0.699.36±0.38NDNDNDNDND3.69±0.25NDNDNDNDND奶油香
), ArticleFig(id=1320175830161018956, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=表2, caption=

酸奶样品中的挥发性风味成分（μg/kg）

, figureFileSmall=null, figureFileBig=null, tableContent=
类别风味物质保留时间（min）L3L5L7L10L14L18L20L23L25L28L30L31L32香气特征[15–16]
酯类丁酸乙酯7.36783.30±5.63NDNDNDNDNDND93.76±3.97344.72±17.65NDNDND121.17±6.28菠萝、香蕉和苹果香
庚酸乙酯17.2596.42±0.35NDNDNDNDNDND27.60±1.05147.54±10.88NDNDNDND菠萝香
辛酸乙酯26.76990.90±52.55342.16±26.833520.62±189.00765.49±26.16868.43±53.76783.91±58.512066.32±145.90831.11±63.2810044.03±467.67718.21±35.18147.43±10.091802.86±60.36ND脂香味
月桂酸乙酯40.4161.44±0.053.31±0.18NDNDNDNDNDND139.61±6.1712.44±0.43NDNDND果香和花香
癸酸乙酯34.774111.65±7.3160.14±4.631001.59±59.9681.70±3.15318.60±11.0647.93±2.66191.19±5.76194.13±10.682633.64±127.28191.36±14.5816.64±1.1035.58±1.78ND椰子香
己酸乙酯18.254507.24±31.16ND676.25±39.28ND66.53±5.26ND41.62±1.74666.28±27.926657.53±527.00128.44±5.9640.23±1.28894.75±65.46ND菠萝香
乙酸乙酯3.8441446.18±49.70NDND832.48±26.62968.14±54.211004.14±61.788807.02±455.931155.72±81.973730.71±248.231394.10±44.012984.88±195.077906.71±504.67ND橙子味
乳酸异戊酯29.9585.93±0.44NDNDNDNDND116.85±8.2916.26±0.74130.95±5.68ND53.21±3.04156.76±11.93ND刺激味
棕榈酸乙酯45.553NDND85.43±6.3911.90±0.6411.31±0.376.45±0.508.43±0.40ND37.10±1.86NDNDNDND蜡香、奶油香
9-癸烯酸乙酯33.2918.43±1.233.74±0.15NDNDNDNDND24.66±1.91285.49±21.27NDNDNDND果香、蜡香、脂香
L-乳酸乙酯19.73195.83±3.7971.59±2.831305.07±66.83NDNDND2364.86±152.12440.17±28.631658.33±123.40ND457.20±23.85898.38±54.03929.50±62.60果香、奶油香
甲酸乙烯酯2.645NDNDNDNDNDNDNDNDND374.28±22.36ND2359.08±137.88ND刺激味
丁位癸内酯44.4421.85±0.061.13±0.05NDNDNDNDND1.15±0.05NDNDNDNDND椰子香、奶油香
乙酸戊酯10.012241.08±10.6698.07±5.62262.62±17.66168.36±8.21253.34±8.92226.19±12.55536.23±26.64166.52±8.49576.60±37.33ND76.91±5.07848.27±37.92ND香蕉味
酸类丁酸30.86ND21.20±1.46NDNDNDNDND21.21±0.751200.03±62.32129.37±5.1668.85±3.06193.39±15.20511.37±25.79刺激味
辛酸42.108NDND581.78±32.08165.32±5.53103.43±3.89281.90±19.29115.11±6.53ND3130.22±249.72160.90±9.49NDNDND焦糖香味
己酸37.66917.18±0.9736.37±1.88669.38±48.42155.30±7.46ND401.11±30.56327.78±19.2858.42±4.138127.88±452.85646.78±20.91832.41±44.04832.85±30.89727.09±36.85甜香味
正癸酸44.786NDND34.82±2.3612.09±0.80ND18.11±0.69NDND320.92±11.2547.24±1.5718.63±1.10NDND奶香味
2-甲基己酸32.454ND35.61±1.49NDNDNDNDND46.63±1.50NDNDND126.96±8.14152.47±6.48水果香
棕榈酸54.4683.73±0.166.05±0.19NDNDNDNDND9.75±0.66NDNDNDNDND微弱油脂香、蜡香
醋酸23.688ND212.25±6.52541.37±37.1962.48±2.63122.62±3.8289.35±3.53219.17±11.36147.52±7.66412.88±17.29575.55±44.161423.08±48.031200.01±88.853489.31±276.58刺激性气味
苯甲酸46.589NDND650.95±21.3668.39±5.44ND226.84±7.2776.03±4.7112.46±0.54ND535.77±21.49NDNDND苦杏仁味
丙酸27.392NDNDNDNDNDNDNDNDNDNDNDND56.28±1.78刺激味
环己甲酸42.369NDNDNDNDNDNDNDNDNDND18.31±0.6377.88±4.26ND山楂香
醇类异戊醇14.0661416.71±113.221060.64±48.743466.50±163.73846.72±67.032166.35±106.452146.33±78.375179.61±330.371531.43±56.603928.38±167.582505.03±189.911539.04±74.416484.65±239.814059.69±257.59苦杏仁味
丁二醇27.81ND590.12±33.26NDNDNDNDND143.83±6.7030.24±2.09ND536.96±26.20546.31±24.232136.51±128.50奶油香、果香
3-甲硫基丙醇33.99315.34±0.676.38±0.31NDNDND51.55±2.80ND9.55±0.67ND109.90±7.04ND72.60±5.40ND硫醇香气
β-苯乙醇39.264301.43±20.64167.99±8.70847.89±34.57440.51±24.20458.69±16.45964.28±33.341325.65±48.45540.61±38.681039.81±58.562516.07±91.74684.58±39.095045.64±391.75450.27±21.06蜜香玫瑰香
酮类2-庚酮12.64320.61±1.23NDND16.90±0.52ND94.84±2.90NDNDNDND81.92±4.08NDND梨子香
3-羟基-2-丁酮16.748ND18.35±0.72NDNDND13.44±0.549127.21±576.7225.59±1.93NDND146.49±9.941846.11±91.44ND奶油香
2-壬酮21.38217.80±1.40NDNDNDNDNDND14.39±0.83333.72±12.16NDND0.00±0.00ND果香味
其他苯乙烯16.09767.28±4.2954.94±4.032215.63±91.251078.52±46.621312.33±49.321173.00±87.671065.09±34.79190.57±10.28NDND52.85±2.96746.62±39.64125.59±7.91树脂香、花香
十二烷11.70934.54±1.1457.28±3.84NDNDNDNDND40.94±2.19426.15±27.04NDNDNDND微弱烃类气味
异丁烷9.44130.43±9.04NDNDNDNDND262.54±19.91ND277.99±11.77NDNDNDND微弱石油样气味
2,3,5-三甲基吡嗪19.98713.43±0.699.36±0.38NDNDNDNDND3.69±0.25NDNDNDNDND奶油香
), ArticleFig(id=1320175830257487949, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Table 3, caption=

Organic acid content in yogurt samples (mg/L)

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号酒石酸苹果酸乳酸乙酸柠檬酸琥珀酸
L3147.13±0.7183.37±4.245741.28±4.24226.64±3.5412.5±6.36341.96±7.07
L5162.13±1.4154.75±4.246015.58±2.12502.73±5.667.55±2.12218.1±4.95
L7151.17±4.9555.68±4.954986.71±4.951047.25±03.85±4.95295.56±2.83
L10199.72±7.0752.8±4.246552.89±5.66509.79±2.837.89±5.66242.86±2.12
L14158.61±7.0762.00±7.075638.63±1.41368.43±2.1212.77±4.95249.33±2.12
L18169.6±5.666.83±4.246055.71±4.24264.42±021.07±3.54146.11±0.71
L20229.47±0.7188.71±2.125151.28±0.71219.25±0.7121.32±0.71718.96±6.36
L23169.84±6.3627.03±1.415391.73±6.36200.02±7.0727.35±4.95597.76±1.41
L25303.1±1.4126.72±7.076056.34±3.5485.6±2.1213.87±6.36564.1±6.36
L28152.97±6.36177.91±5.666984.52±6.36102.89±5.6642.99±3.54424.75±4.24
L30376.01±4.24130.32±0.715846.9±6.36395.57±6.3634.34±1.41174.23±1.41
L31141.71±5.6645.72±4.245962.26±5.66174.99±4.9527.86±6.36287.45±1.41
L32120.9±6.36152.12±2.835823.49±0.71231.73±5.6640.69±0.71280.71±6.36
), ArticleFig(id=1320175830345568334, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=表3, caption=

酸奶样品中的有机酸含量（mg/L）

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号酒石酸苹果酸乳酸乙酸柠檬酸琥珀酸
L3147.13±0.7183.37±4.245741.28±4.24226.64±3.5412.5±6.36341.96±7.07
L5162.13±1.4154.75±4.246015.58±2.12502.73±5.667.55±2.12218.1±4.95
L7151.17±4.9555.68±4.954986.71±4.951047.25±03.85±4.95295.56±2.83
L10199.72±7.0752.8±4.246552.89±5.66509.79±2.837.89±5.66242.86±2.12
L14158.61±7.0762.00±7.075638.63±1.41368.43±2.1212.77±4.95249.33±2.12
L18169.6±5.666.83±4.246055.71±4.24264.42±021.07±3.54146.11±0.71
L20229.47±0.7188.71±2.125151.28±0.71219.25±0.7121.32±0.71718.96±6.36
L23169.84±6.3627.03±1.415391.73±6.36200.02±7.0727.35±4.95597.76±1.41
L25303.1±1.4126.72±7.076056.34±3.5485.6±2.1213.87±6.36564.1±6.36
L28152.97±6.36177.91±5.666984.52±6.36102.89±5.6642.99±3.54424.75±4.24
L30376.01±4.24130.32±0.715846.9±6.36395.57±6.3634.34±1.41174.23±1.41
L31141.71±5.6645.72±4.245962.26±5.66174.99±4.9527.86±6.36287.45±1.41
L32120.9±6.36152.12±2.835823.49±0.71231.73±5.6640.69±0.71280.71±6.36
), ArticleFig(id=1320175830425260111, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Table 4, caption=

Sequencing results and alpha diversity indices for bacteria

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号序列数OTU数ACEChaoShannonSimpsonCoverage
L10533711415.5214.200.730.500.999963
L14533711213.2012.250.730.500.999963
L185337155.755.000.040.990.999981
L20533714451.9847.060.580.640.999794
L23533711729.0132.000.350.850.999888
L25533711313.4813.000.160.950.999981
L285337166.006.000.660.611.000000
L3533711115.6712.500.390.790.999925
L30533713537.7235.861.020.500.999925
L31533713838.8938.100.740.600.999963
L32533714649.8148.150.690.720.999850
L5533711617.1516.500.180.940.999963
L7533713648.3845.430.510.730.999775
), ArticleFig(id=1320175830504951888, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=表4, caption=

细菌测序结果统计与alpha多样性指数

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号序列数OTU数ACEChaoShannonSimpsonCoverage
L10533711415.5214.200.730.500.999963
L14533711213.2012.250.730.500.999963
L185337155.755.000.040.990.999981
L20533714451.9847.060.580.640.999794
L23533711729.0132.000.350.850.999888
L25533711313.4813.000.160.950.999981
L285337166.006.000.660.611.000000
L3533711115.6712.500.390.790.999925
L30533713537.7235.861.020.500.999925
L31533713838.8938.100.740.600.999963
L32533714649.8148.150.690.720.999850
L5533711617.1516.500.180.940.999963
L7533713648.3845.430.510.730.999775
), ArticleFig(id=1320175830597226577, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=EN, label=Table 5, caption=

Sequencing results and alpha diversity indices for fungi

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号序列数OTU数ACEChaoShannonSimpsonCoverage
L10399473237.5035.500.890.560.999800
L14399473940.7040.500.710.610.999900
L18399474954.3953.000.470.830.999775
L2039947143150.62147.690.700.720.999574
L233994776111.0399.631.190.410.999299
L25399473033.7833.750.980.460.999850
L28399475357.9558.601.090.470.999800
L3399472631.1729.500.650.580.999825
L30399472831.8331.000.720.540.999825
L31399474548.0146.431.650.280.999875
L32399471718.0417.250.660.680.999950
L5399472228.6426.200.820.540.999825
L7399474445.3744.601.010.530.999925
), ArticleFig(id=1320175830664335442, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320175819566207982, language=CN, label=表5, caption=

真菌测序结果统计与alpha多样性指数

, figureFileSmall=null, figureFileBig=null, tableContent=
样品编号序列数OTU数ACEChaoShannonSimpsonCoverage
L10399473237.5035.500.890.560.999800
L14399473940.7040.500.710.610.999900
L18399474954.3953.000.470.830.999775
L2039947143150.62147.690.700.720.999574
L233994776111.0399.631.190.410.999299
L25399473033.7833.750.980.460.999850
L28399475357.9558.601.090.470.999800
L3399472631.1729.500.650.580.999825
L30399472831.8331.000.720.540.999825
L31399474548.0146.431.650.280.999875
L32399471718.0417.250.660.680.999950
L5399472228.6426.200.820.540.999825
L7399474445.3744.601.010.530.999925
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西藏不同地区传统发酵乳的菌群结构及其风味分析
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田敏 1, 2, 3 , 张静 2, 3 , 于佳俊 2, 3, 4 , 张玉红 4 , 文华英 4 , 李菊梅 2, 3 , 薛洁 2, 3, * , 刘延琳 1, *
食品工业科技 | 生物工程 2026,47(12): 156-166
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食品工业科技 |生物工程 2026 , 47 (12) : 156 -166
西藏不同地区传统发酵乳的菌群结构及其风味分析
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田敏1, 2, 3 , 张静2, 3, 于佳俊2, 3, 4, 张玉红4, 文华英4, 李菊梅2, 3, 薛洁2, 3, * , 刘延琳1, *
作者信息
  • 1西北农林科技大学葡萄酒学院，陕西杨凌 712100
  • 2中国食品发酵工业研究院有限公司，北京 100125
  • 3国家酒类品质与安全国际联合研究中心，北京 100125
  • 4西藏自治区农牧科学院农产品开发与食品科学研究所，西藏拉萨 850030
通讯作者:
薛洁（1974−），女，博士，教授级高级工程师，研究方向：发酵工程，E-mail：。
刘延琳（1966−），女，博士，教授，研究方向：酿酒微生物，E-mail：。
作者简介:

田敏（2001−），女，硕士研究生，研究方向：发酵工程，E-mail：

Analysis of Bacterial Flora Structure and Flavor of Traditional Fermented Milk from Different Regions of Tibet
Min TIAN1, 2, 3 , Jing ZHANG2, 3, Jiajun YU2, 3, 4, Yuhong ZHANG4, Huaying WEN4, Jumei LI2, 3, Jie XUE2, 3, * , Yanlin LIU1, *
Affiliations
  • 1College of Enology, Northwest A&F University, Yangling 712100, China
  • 2China National Research Institute of Food and Fermentation Industries, Beijing 100125, China
  • 3International Joint Research Center of Quality and Safety of Alcoholic Beverages, Beijing 100125, China
  • 4Institute of Agricultural Product Development and Food Science, Tibet Academy of Agricultural and Animal Husbandry Sciences, Lhasa 850030, China
出版时间: 2026-06-15 doi: 10.13386/j.issn1002-0306.2025050095
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为了探究西藏传统发酵乳中的微生物体系及其风味品质，利用高通量测序技术以及顶空固相微萃取-气相色谱-质谱联用技术，对西藏传统发酵乳中的微生物多样性及挥发性风味成分进行分析，并通过液相色谱法测定其有机酸的种类及含量。结果表明：西藏传统发酵乳制品中的主要优势细菌菌群为乳杆菌属（Lactobacillus）、链球菌属（Streptococcus）、乳球菌属（Lactococcus）等，主要优势真菌菌群为酵母菌属（Saccharomyces）、克鲁维酵母属（Kluyveromyces）、地霉属（Geotrichum）等。该乳制品中含乳酸、乙酸、柠檬酸等6种有机酸，其中乳酸含量最高，最高可达6989.02 mg/L；其次是乙酸，达1047.25 mg/L。该乳制品中共检测出35种挥发性风味物质，其中酯类物质相对含量占比最高（43.98%），其次是醇类（28.98%）和酸类物质（16.00%）。不同地区发酵乳中的挥发性风味物质种类和含量各有差异，其中日喀则市的发酵乳以菠萝和香蕉等果香和奶油香气为主，其风味较为丰富复杂；那曲地区的发酵乳以奶油香气、坚果香气和酸甜味为主；阿里地区的发酵乳水果香气较为明显。通过Spearman相关系数与OAV>1的特征风味物质进行相关性分析，与风味物质相关性较高的细菌菌属主要是Lactobacillus、Acetobacter、Leuconostoc；与风味物质相关性较高的真菌菌属主要是Apiotrichum、Dipodascaceae、Pichia。本研究阐明了西藏传统发酵乳中菌群结构与其风味物质形成的相关性，为传统乳制品的标准化生产、风味定向调控及功能性菌株开发提供理论依据。

发酵乳  /  高通量测序  /  菌群结构  /  挥发性风味  /  相关性分析

To investigate the microbial community structure and flavor quality of traditional Tibetan fermented milk, high-throughput sequencing and headspace solid-phase microextraction-gas chromatography-mass spectrometry (HS-SPME–GC–MS) were used to analyze the microbial diversity and volatile flavor components in traditional Tibetan fermented milk. The types and contents of organic acids were determined by liquid chromatography. The results showed that the main dominant bacterial flora in traditional Tibetan fermented milk products were Lactobacillus, Streptococcus, and Lactococcus, etc., while the main dominant fungal flora were Saccharomyces, Kluyveromyces, and Geotrichum, etc. Six organic acids were identified, including acetic acid, citric acid, lactic acid and so on, among which, lactic acid was the most abundant (6989.02 mg/L), followed by acetic acid (1047.25 mg/L). A total of 35 volatile flavor compounds were detected in the fermented milk, with esters accounting for the highest proportion (43.98%), followed by alcohols (28.98%) and acids (16.00%). The composition and concentration of volatile flavor compounds in fermented milk exhibited notable regional variations. Fermented milk from Shigatse was distinguished by pronounced fruity aromas, particularly pineapple and banana, complemented by creamy notes, which together contributed to a rich and complex flavor profile. By contrast, fermented milk from Nagqu was characterized by strong creamy aromas, nutty undertones, and a balanced sweet–sour taste, whereas fermented milk from Ali demonstrated a predominance of more intense fruity aromas. Spearman’s correlation analysis based on key odor–active compounds (odor activity value, OAV>1) revealed distinct associations between microbial taxa and flavor compounds. The bacterial genera most strongly correlated with key flavor compounds were primarily Lactobacillus, Acetobacter and Leuconostoc, while the fungal genera most closely linked to flavor development included Apiotrichum, Dipodascaceae and Pichia. This study elucidated the correlation between microbial community structure and the formation of aroma compounds in traditional Tibetan fermented milk, providing a theoretical foundation for the standardized production, targeted flavor regulation, and development of functional strains for traditional dairy products.

fermented milk  /  high-throughput  /  community structure  /  volatile flavor components  /  correlation analysis
田敏, 张静, 于佳俊, 张玉红, 文华英, 李菊梅, 薛洁, 刘延琳. 西藏不同地区传统发酵乳的菌群结构及其风味分析. 食品工业科技, 2026 , 47 (12) : 156 -166 . DOI: 10.13386/j.issn1002-0306.2025050095
Min TIAN, Jing ZHANG, Jiajun YU, Yuhong ZHANG, Huaying WEN, Jumei LI, Jie XUE, Yanlin LIU. Analysis of Bacterial Flora Structure and Flavor of Traditional Fermented Milk from Different Regions of Tibet[J]. Science and Technology of Food Industry, 2026 , 47 (12) : 156 -166 . DOI: 10.13386/j.issn1002-0306.2025050095
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西藏地区以其独特的自然环境和悠久的畜牧业传统，孕育出了丰富多样的传统发酵乳制品[1]。这些乳制品不仅是当地居民饮食文化的重要组成部分，也因其独特的风味和营养价值而备受关注，其中酸奶是西藏最具有代表性的传统发酵乳制品，具有鲜明的地域特色和民族特色[2]。由于西藏地区具有低温、低压、高海拔、强紫外线和缺氧等特点，传统发酵乳制品中的微生物在自然选择和遗传进化的作用下，形成了复杂独特的菌群结构，具有很高的研究价值。然而，与现代工业化的乳制品相比，西藏传统乳制品的生产过程往往具有较强的地域性和传统性，其微生物群落结构和风味形成机制尚未得到充分研究[3]。研究表明，由于西藏不同地区的环境多样性以及传统发酵工艺的多样性，西藏不同地区的传统发酵乳的风味不同，微生物群落结构也有差异[4]。近来，随着消费者对食品风味、营养和安全性要求的提高，解析传统发酵乳制品中微生物菌群结构及其挥发性风味物质，成为提升产品品质、推动工业化生产的关键科学问题[5]。
高通量测序技术（High-Throughput Sequencing，HTS）能够在短时间内对大量DNA或RNA序列进行快速测序，具有高通量、高准确性、低成本、快速和应用广泛等优点[6]。目前高通量测序技术已经被广泛运用于各个领域，在食品领域主要用于分析微生物群落的多样性[7]。高通量测序技术的出现彻底改变了传统发酵乳中微生物多样性的研究方法，它可以揭示更复杂的微生物多样性，这为传统发酵食品的监控、操作和安全性提供了强大的技术支持，推动了传统发酵食品产业的升级和发展[8–9]。Wu等[10]基于高通量测序技术，对青藏高原传统发酵牦牛奶的细菌群落结构特征进行分析，在属水平上，共鉴定出了82个细菌菌属，其中优势菌属为乳杆菌属Lactobacillus（64.73%）和链球菌属Streptococcus（28.48%）。气相色谱-质谱联用技术（GC-MS）在食品领域中主要用于分析食品中的挥发性风味化合物，具有高灵敏度、多功能性和数据库支持等优点[11]。Fang等[12]利用GC-MS技术分析了青藏高原不同生态区生产的传统发酵牦牛奶挥发性风味成分特征，发现挥发性风味物质的相对百分比组成在不同生态区之间有所不同。
本研究以西藏传统发酵乳为原料，利用高通量测序技术分析西藏传统发酵乳的微生物群落组成及其多样性，明确其中的优势菌属，并对西藏传统发酵乳中的蛋白质、有机酸和挥发性风味物质等进行分析。研究成果有望为传统发酵乳制品的标准化生产、风味定向调控及功能性菌株开发提供理论依据，同时为高原特色食品的资源保护与产业化升级注入科技动力。
1　材料与方法
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1.1　材料与仪器
酸奶　2024年8月，从西藏不同地区共采集13个实验样品，样品均由西藏牧民采用传统方式自然发酵而成，主要是山羊奶、绵羊奶以及牦牛奶发酵而成的酸奶样品，具体信息如表1所示。在采集新鲜样品后，立即将其收集于50 mL的无菌离心管内，并先进行冷藏保存，随后转移至实验室−20 ℃冰箱中保存备用；乳酸、柠檬酸、苹果酸、酒石酸、琥珀酸、乙酸标准品　纯度≥98%，北京化学试剂公司；磷酸二氢钾、磷酸（分析纯）、甲醇（色谱纯）　西陇化工股份有限公司；2-辛醇（内标物，纯度≥98%） TCI；E.Z.N.A.®Soil DNA kit　美国OMEGA公司；琼脂糖　西班牙BIOWESTE公司；DNA凝胶回收试剂盒　南京诺唯赞生物科技股份有限公司；PCR所用试剂　大连TaKaRa公司；细菌基因组DNA提取试剂盒　天根生化科技（北京）有限公司。
E-WAX ETR色谱柱（30 m×0.25 mm，0.50 μm）　美国PerkinElmer公司；MiSeq PE300高通量测序仪　美国Illumina公司；2-16N型高速微量离心机　湖南恒诺仪器设备有限公司；T20型PCR仪　杭州朗基科学仪器有限公司；1300 SERIES A2型超净工作台　美国Thermo公司；5977B型气相色谱-质谱联用仪　美国Agilent公司。
1.2　实验方法
1.2.1　挥发性风味成分测定
参考Jiang等[4]研究方法略有修改，准确称取酸奶样本3.50 g、氯化钠2 g、超纯水3.5 mL置于20 mL顶空瓶中，以120 mg/L的2-辛醇作为内标，其添加量为10 μL，进行固相微萃取，在55 ℃条件下萃取45 min，在250 ℃条件下解析2 min。
GC条件：柱箱温度40 ℃保持2 min，以3 ℃/min的速率升至180 ℃，再以6 ℃/min的速率升至240 ℃保持5 min。进样口温度为250 ℃，载气（高纯氦气）流速1.5 mL/min。MS条件：EI源电离，离子源温度220 ℃，70 eV电子能量，质量扫描范围 m/z 35～500，扫描方式为全扫描。
定性：在NIST数据库中，通过对各色谱峰的保留时间进行匹配，选择匹配度大于85的化合物为有效风味物质。定量：根据各物质峰面积比等于物质浓度比，采用半定量方法，将各化合物的保留峰面积与2-辛醇的峰面积对比，从而计算出各化合物的相对含量。
挥发性风味物质相对含量=（风味物质峰面积×内标物质含量）/内标物质峰面积
1.2.2　酸奶样品中蛋白质含量的测定
参照GB 5009.5-2016《食品安全国家标准食品中蛋白质的测定》对酸奶中的蛋白质含量进行测定。
1.2.3　酸奶样品中有机酸含量的测定
参照GB 5009.157-2016《食品安全国家标准 食品中有机酸的测定》对酸奶中的有机酸含量进行测定。
1.2.4　高通量测序
利用试剂盒提取样本DNA，利用1%的琼脂糖凝胶电泳检测DNA的提取质量，并测定其浓度和纯度。PCR扩增体系：10×PCR buffer（Mg2+plus）2.5 μL，dNTP Mixture 2.0 μL，上游引物1.0 μL，下游引物1.0 μL，DNA模板1.0 μL，rTaq DNA聚合酶（5 U/μL）0.3 μL，ddH2O补足至25 μL。细菌16S rRNA基因V3~V4可变区：上游引物27F:5’-AGAGTTTGATCMTGGCTCAG-3’，下游引物1492R：5’-GGTTACCTTGTTACGACTT-3’。PCR扩增条件：95 ℃预变性3 min，35个循环（95 ℃变性30 s，55 ℃退火30 s，72 ℃延伸30 s），72 ℃稳定延伸10 min，最后4 ℃保存。真菌ITS区域的通用引物为ITS1F。上游引物：ITS1F（5’-CTTGGTCATTTAGAGGAAGTAA-3’），下游引物：ITS2R（5’-GCTGCGTTCTTCATCGATGC-3’）。PCR扩增条件：95 ℃预变性3 min，95 ℃变性30 s，55 ℃退火30 s，72 ℃延伸45 s，扩增30个循环；72 ℃再延伸10 min，4 ℃保存[13–14]。PCR产物在Illumina MiseqPE300平台进行高通量测序。
1.3　数据处理
利用Excel对数据进行整理和分析；利用SPSS 27.0方差分析（ANOVA）进行样本间的差异显著性分析；使用Chroma TOF软件（V 4.3x，LECO）和NIST库对质谱数据进行分析；基于美吉生物信息平台进行主坐标分析（principal co-ordinates analysis，PCoA）；利用软件Origin 2024对数据进行统计分析和绘图，其中组间差异显著性的检验采用ANOVA方法进行（P<0.05）。
2　结果与分析
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2.1　酸奶样品中挥发性风味成分分析
13种酸奶样本中挥发性风味成分检测结果见表2，由表2可知，样本中挥发性风味成分种类复杂。在13种样本中共检测出35种主要的挥发性风味成分，包括酯类（14种）、酸类（10种），醇类（4种），酮类（3种）和其他化合物（4种）。图1展示了不同样品中挥发性风味物质类别的相对组成，由图1可知，不同样品中挥发性风味物质的分布存在明显差异，大多数化合物在各样品中的相对含量并不均衡，而是在部分样品中呈现明显富集。13种样品挥发性成分占比从高到低依次为：酯类>醇类>酸类>酮类>其他化合物。整体来看，酯类、醇类和部分酸类化合物在样品间的变化幅度较大，是造成样品聚类差异的主要因素。L25、L20和L31样品在多个酯类、醇类及酮类化合物上表现出较高信号强度，形成较为集中的高丰度区域，说明这几个样品挥发性风味物质种类较丰富、含量较高，香气特征更为突出。
酯类化合物由脂肪酸和醇酯化产生，其风味阈值低，风味贡献明显，可以赋予酸奶水果香气和奶油香气[18]。从各样品酯类物质组成来看，乙酸乙酯、辛酸乙酯、己酸乙酯和 L-乳酸乙酯是造成不同样品酯类风味差异的主要化合物，但其在各样品中的分布并不一致。L25 样品酯类物质总相对含量最高，为 26386.26 μg/kg，主要由辛酸乙酯、己酸乙酯、乙酸乙酯和癸酸乙酯贡献，说明其果香和脂香特征较为突出；L31 和 L20 酯类物质总相对含量也较高，分别为 14902.39 μg/kg 和 14132.52 μg/kg，二者均以乙酸乙酯为主要酯类成分，可能赋予样品较明显的果香特征。L7 样品中辛酸乙酯、L-乳酸乙酯和癸酸乙酯含量较高，酯类香气组成相对丰富。相比之下，L5 样品酯类物质总相对含量最低，为 580.16 μg/kg；L32 样品酯类物质检出种类最少，仅检出丁酸乙酯和 L-乳酸乙酯 2 种。总体来看，不同样品中酯类物质的组成和含量差异明显，说明不同地区传统发酵乳在果香、奶油香和脂香等酯类风味特征上具有一定差异性。
乳糖代谢、甲基酮还原、氨基酸代谢、亚油酸和亚麻酸降解等会影响醇类化合物的生成，但其风味阈值高，对发酵乳的风味贡献较低[19]。样品中检出的醇类物质主要包括异戊醇、β-苯乙醇、丁二醇和 3-甲硫基丙醇。其中，异戊醇和 β-苯乙醇在 13 个样品中均有检出，是主要醇类风味物质；异戊醇在 L31 中含量最高，为 6484.65 μg/kg，β-苯乙醇同样在 L31 中含量最高，为 5045.64 μg/kg。丁二醇主要在 L5、L23、L25、L30、L31 和 L32 中检出，其中 L32 含量最高，为 2136.51 μg/kg；3-甲硫基丙醇仅在部分样品中检出，其中 L28 含量最高，为 109.90 μg/kg。从样品间差异来看，L31 醇类物质总相对含量最高，为 12149.20 μg/kg，主要由异戊醇和 β-苯乙醇贡献，说明其发酵香、花香和蜜香特征较为突出；L32 和 L20 的醇类物质总相对含量也较高，分别为 6646.47 μg/kg 和 6505.26 μg/kg；而 L10 醇类物质总相对含量最低，为 1287.23 μg/kg，说明其醇类香气贡献相对较弱。总体来看，不同样品中醇类物质的组成和含量差异明显，体现出不同地区传统发酵乳在发酵香和花香特征上的样品差异。
酸类化合物主要与脂肪分解和微生物发酵等代谢途径有关，它是酸奶的主要风味物质，也是决定酸奶滋味的关键[20]。由表2可知，不同样品中酸类物质的组成和含量差异明显。己酸和醋酸在多数样品中均有检出，是分布较广的酸类物质；而辛酸、丁酸、苯甲酸、正癸酸、2-甲基己酸、丙酸和环己甲酸等在不同样品中的检出情况差异较大。各样品中，L25 酸类物质总相对含量最高，为 13191.92 μg/kg，主要由己酸、辛酸和丁酸贡献，说明其甜香、焦糖香及酸刺激性风味较为突出；L32 酸类物质总相对含量也较高，主要由醋酸、己酸和丁酸贡献，可能赋予其较明显的酸味和刺激性气味；L31、L30、L28 和 L7 样品中酸类物质含量也相对较高，但主要贡献物质有所不同，其中 L31 和 L30 以醋酸、己酸为主，L28 以己酸、醋酸和苯甲酸为主，L7 则以己酸、苯甲酸、辛酸和醋酸为主。相比之下，L3 样品酸类物质总相对含量最低，为 20.91 μg/kg，仅检出己酸和棕榈酸。总体来看，不同样品酸类物质的组成和含量差异明显，说明不同地区传统发酵乳在酸味强度及甜香、焦糖香等风味特征上存在差异。
酮类化合物主要由不饱和脂肪酸经过一系列化学反应而形成的化合物[21]。由表2可知，不同样品中酮类物质的检出情况和含量差异明显。L20 样品中 3-羟基-2-丁酮含量最高，为 9127.21 μg/kg，具有奶油香，是该样品酮类风味的主要贡献物质；L31 样品中 3-羟基-2-丁酮含量也较高，为 1846.11 μg/kg，说明其奶油香特征相对明显。L25 样品中主要检出 2-壬酮，含量为 333.72 μg/kg，可能赋予其一定果香特征；L30 样品中检出 2-庚酮和 3-羟基-2-丁酮，总相对含量为 228.41 μg/kg；L18 样品中 2-庚酮含量较高，为 94.84 μg/kg，同时检出少量 3-羟基-2-丁酮。相比之下，L7、L14、L28 和 L32 样品中未检出酮类物质。总体来看，不同样品中酮类物质的组成差异明显，其中 L20 和 L31 主要表现为奶油香特征，L25 则具有一定果香特征。烷烃类化合物和含氮类化合物的风味阈值相对较高，所以其对酸奶样品风味的形成影响不明显，但一定条件下，这些物质可使酸奶的口感更加饱满，对酸奶的香气品质也有优化作用。
总体来看，日喀则市的酸奶样本主要以酯类化合物带来的菠萝和香蕉等果香和奶油香气为主，酸类物质含量也较高，其风味较为丰富复杂；那曲地区的酸奶样本中的风味物质并不突出，整体以酯类化合物和酸类化合物带来的奶油香气、坚果香气和酸甜味；阿里地区的酸奶样本中由酮类物质带来的水果香气较为明显。而来自日喀则吉隆县的海拔最低的酸奶样本L25中的酯类物质、酸类物质和酮类物质的含量和种类最为突出，与高海拔地区的酸奶样本相比差异很大。推测海拔高度影响了温度和草场环境，同时也影响了微生物群落组成，从而对酸奶的风味产生影响，这与董闯等[15]研究发现不同海拔高度的牦牛乳拉拉关键香气成分存在差异的结果一致。此外，不同地区酸奶样品的风味还与制作工艺、来源等相关。
2.2　酸奶样品中蛋白质含量分析
酸奶样品中的蛋白质含量如图2所示。根据标准GB 25190-2010规定，每100 mL牛奶蛋白质含量不得低于2.9 g。已有研究表明，牦牛酸奶能够提供满足人体每日膳食所需要的蛋白质，且其中含有人体所需的18种氨基酸[17]。经测定，酸奶样品中的蛋白质含量符合国家标准，其蛋白质含量高于市售的普通酸奶，且不同样品中蛋白质含量有所不同。其中，L28号样品中蛋白质含量最高，为16.58%；L25号样品中蛋白质含量最低，为3.28%。
Liu等[18]研究发现高海拔牦牛奶的营养物质更加丰富；叶丹[22]对采自西藏林芝市巴宜区（3000 m）、拉萨市堆龙德庆（3700 m）、那曲市班戈县（4700 m）的三个不同海拔高度的牦牛乳样品中的蛋白质进行研究，发现来自不同海拔高度的牦牛乳样品中的蛋白质含量差异极显著，且牦牛乳中的蛋白质含量随着海拔的升高含量增多；席斌等[23]对采自3个不同地区的牦牛乳样品进行分析发现，海拔高度会对牦牛乳中的乳蛋白含量以及乳脂肪含量产生影响，且样品中的乳蛋白和乳脂肪含量随海拔升高而增加。而L25号样品采自海拔3012 m，其余样品均采自海拔4000 m以上，说明海拔高度确实对酸奶样品中的蛋白质含量有影响，与上述研究结果一致。
由于酸奶样品的来源不同，其蛋白质含量也有所差异。此外，西藏不同地区酸奶中蛋白质含量的差异还受到不同地区的草场类型、饲养管理以及牦牛品种等因素的影响。这些因素相互作用，共同决定了牦牛酸奶中蛋白质的含量和品质。
2.3　酸奶样品中有机酸含量分析
酸奶样品中的有机酸含量如表3所示。在酸奶发酵过程中，乳酸通过乳酸菌将乳糖分解而产生，它是最主要的有机酸。乳酸的含量通常较高，能够显著降低酸奶的pH，使其具有酸味，并且有助于抑制有害微生物的生长[24]。由表3可知，乳酸是酸奶样品中最主要的有机酸，其含量远远高于其他有机酸。乳酸的酸味温和适中，是常见的酸味剂，有很强的防腐保鲜功效。其中，L28号样品中乳酸含量最高（6984.52 mg/L），L7号样品中乳酸含量最低（4986.71 mg/L）。乙酸具有特殊的醋酸味，但如果含量过高，可能会使酸奶产生不可接受的醋味[25]。苹果酸具有柔和的天然果香气味，容易被人体吸收，且对人体内部循环具有重要健康功效[26]。其含量会因发酵菌株和发酵条件的不同而有所差异，它可以增加酸奶的酸味和风味复杂性。其中，L28样品苹果酸含量最高（177.91 mg/L），L18样品含量最低（6.83 mg/L）。酸奶样品中含量最低的有机酸为柠檬酸，所有样品含量均在0~50 mg/L。柠檬酸可以增强酸奶的风味，使酸奶的味道更加丰富和立体[27]。
乳酸和乙酸是酸奶酸味的主要来源，而柠檬酸和苹果酸则可以增加酸奶的风味层次。不同有机酸的比例和含量会影响酸奶的整体风味，使其具有独特的口感和香气。有机酸在酸奶发酵过程中还可以促进其他有益成分的生成，如B族维生素等，从而提高酸奶的营养价值和健康功能[28]。有机酸的形成与酸奶的质地密切相关，适当的酸度可以使酸奶达到理想的稠度和弹性。例如，乳酸的积累可以促进酸奶的凝固，使其具有更好的质地。同时乳酸等有机酸的积累可以降低酸奶的pH，抑制有害微生物的生长繁殖，从而延长酸奶的保质期[29]。
2.4　酸奶样品中微生物多样性分析
2.4.1　属水平菌群结构分析
在细菌属水平上（图3a），乳杆菌属（Lactobacillus）、链球菌属（Streptococcus）、乳球菌属（Lactococcus）是最主要的优势菌属，此外还含有少量的梭菌属（Clostridium）、双歧杆菌属（Bifidobacterium）、无氧芽孢杆菌属（Anoxybacillus）、醋杆菌属（Acetobacter）、明串珠菌属（Leuconostoc）等。在样本L18、L25、L28中，乳杆菌属（Lactobacillus）的平均相对丰度达到了90%以上，它可以将乳糖分解为葡萄糖和半乳糖，进一步通过厌氧发酵将其转化为乳酸，并且在发酵过程中产生乙醛、双乙酰等挥发性化合物，从而对酸奶风味有很大影响。样本L5中链球菌属（Streptococcus）的平均相对丰度为96.95%，链球菌属中的嗜热链球菌（Streptococcus thermophilus）是酸奶发酵中不可或缺的关键菌种之一，嗜热链球菌能够快速发酵乳糖，降低pH，促进酪蛋白凝结，这种凝结作用可以使酸奶形成稳定的凝胶结构，赋予酸奶良好的质地和持水性[30]。样本L32中乳球菌属（Lactococcus）的平均相对丰度为87.83%，其中乳酸乳球菌（Lactococcus lactis）在发酵过程中能够产生多种风味物质，如乙醛、双乙酰（2,3-丁二酮）和乙偶姻等，这些代谢物参与形成发酵乳制品的特征性风味和香气[31]。
在真菌属水平上（图3b），酵母菌属（Saccharomyces）、克鲁维酵母属（Kluyveromyces）、地霉属（Geotrichum）、双足囊菌属（Dipodascus）等是最主要的优势菌属，此外还含有少量的毕赤酵母属（Pichia）、锥毛壳属（Coniochaeta）等。在样本L14、L18、L20、L28中，酵母菌属（Saccharomyces）的平均相对丰度较高，达到了50%以上。酵母菌属（Saccharomyces）在发酵过程中能够代谢乳糖和乳酸，产生多种挥发性和非挥发性风味物质，并与乳酸菌协同发酵，显著提升了酸奶的整体品质和风味。在样本L7、L25、L30、L32中，地霉属（Geotrichum）的平均相对丰度较高。地霉属菌株在发酵过程中可以产生酯类、醇类和有机酸等物质，这些物质对酸奶的风味有重要贡献。在样本L3、L5、L10、L23中，克鲁维酵母属（Kluyveromyces）的平均相对丰度较高。克鲁维酵母属菌株能够发酵乳糖，降低酸奶中的乳糖含量，而乳糖的水解可以减少乳糖不耐受症状，比如马克斯克鲁维酵母能够利用乳糖作为碳源，产生β-半乳糖苷酶，从而将乳糖分解成葡萄糖和半乳糖[32]。各样本中的真菌菌群结构差异较大，其优势菌属各不相同，说明酸奶样品具有丰富的真菌多样性。
2.4.2　 α多样性分析
OTU（Operational taxonomic unit）即操作分类单元，将序列按照彼此的相似性归为若干聚类单元，每个聚类单元为一个OTU，样本中OTU的数目能够初步反映其中微生物物种的丰富程度。各样品的细菌和真菌OUT数结果统计和alpha多样性指数如表所示。
本研究在97%相似性水平下进行OTU聚类。如表4所示，各发酵乳样本中细菌测序样本中OTU数目介于5~46之间。通过比较各发酵乳样本中细菌OTU数目可知，L32中细菌OTU数目最多，L18中细菌OTU数目最少；如表5所示，各发酵乳样本中真菌测序样本中OTU数目介于17~143之间。通过比较各发酵乳样本中真菌OTU数目可知，L20中真菌OTU数目最多，L32中真菌OTU数目最少。
Alpha多样性分析是指单样本内物种多样性分析，通过计算几种多样性指数值来反映样本内微生物物种的丰富度和多样性程度，alpha多样性指数包括ACE、Chao、Shannon、Simpson和Coverage等。在生态学研究中用ACE和Chao指数来评估物种总数。ACE和Chao指数反映样本内微生物物种的丰富度，计算时仅考虑群落中物种数量，不考虑各物种的丰度，这两个指数越高，样本内微生物物种的丰富度越高。Shannon和Simpson指数反映样本内微生物物种的多样性程度，计算时考虑样本中物种丰富度和物种均匀度两方面。在物种丰富度相同时，样本内各物种的均匀度越高，则多样性程度越高。Coverage指数反映样本文库的覆盖率，用于评估测序结果能否反映样本内微生物的实际情况[33]。
如表4及表5所示，本研究中所有细菌与真菌测序样本计算得到的Coverage指数值均高于99%，表明测序结果的文库覆盖率高，能够反映所有样本内微生物的实际情况。
如表4所示，通过比较细菌alpha多样性指数可知，L20、L32、L7的细菌ACE和Chao指数较高，表明其细菌物种丰富度较高，说明高海拔地区（如L20：4860 m、L31：4850 m、L7：4565 m）影响细菌物种的丰富度，可能原因是高海拔地区低温、强紫外线等极端环境促使微生物产生适应性分化，形成独特的群落结构。根据细菌Shannon和Simpson指数排序可知，L30（定日县协格尔镇）的细菌多样性程度最高，但其海拔（4425 m）并非最高，分析原因可能因其位于珠峰北坡，独特的地形加上冰川融水补给为细菌提供了多样化生态位[34]。
如表5所示，通过比较真菌alpha多样性指数可知，真菌高ACE和Chao指数较高的样本L20、L23同样位于高海拔4860 m、4678 m；而真菌多样性最高的L31海拔达4850 m，推测是因为高海拔可能通过低氧、强辐射等条件筛选出适应性强的真菌类群。
分析结果也可以看出，高纬度地区，如那曲地区31°N以上的样本L3、L5、L7、L10，细菌和真菌多样性指数普遍低于日喀则南部28°~29°N的样本L20、L23、L31，可能是因为纬度降低伴随温度升高或降水模式变化，促进微生物代谢活动及物种共存。而经度对微生物多样性同样有影响，东部如那曲地区88°~90°E和阿里地区81°~82°E样本的微生物多样性普遍低于西部日喀则市85°~88°E，可能因日喀则地处喜马拉雅北麓，受印度洋暖湿气流影响更显著，水热条件相对优越，支持更复杂的微生物群落[35]。
2.4.3　 β多样性分析
使用基于Bray-Curtis距离算法的主坐标分析不同样本之间细菌和真菌的β多样性。图4a为细菌测序样本主坐标分析图，图4a中PCoA1和PCoA2对细菌菌群结构差异的贡献率分别为59.18%和32.87%。PCoA1的贡献率高于50%，表明所分析样本之间细菌菌群结构的差异主要可以通过PCoA1方向上的距离来评价。在PCoA1方向上，L30、L32与其他样本距离较大，表明其与其他样本中的细菌群落结构存在较大差异。图4b为真菌测序样本主坐标分析图，该图中PCoA1和PCoA2对真菌菌群结构差异的贡献率分别为57.45%和28.08%。PCoA1的贡献率高于50%，表明所分析样本之间真菌菌群结构的差异主要可以通过PCoA1方向上的距离来评价。在PCoA1方向上，L25、L7、L30、L32距离较近，L18、L20、L14、L31、L28也距离较近，表明这几组样本的真菌群落相似度较高；而其他几组样本距离较远，说明菌群差异性较大。
2.5　酸奶样品中微生物菌群结构与挥发性风味相关性分析
香气活力值（odor activity value，OAV）是挥发性风味物质含量与其气味阈值之比，可以作为评价某个风味成分对整体风味贡献大小的依据[36]。OAV>1的风味物质被认为是对整体风味有贡献的成分，风味物质OAV越大则对风味贡献越大[37]。通过Spearman相关系数，将测序得到的微生物种类与OAV>1的特征风味物质进行相关性分析，结果如图5所示。
细菌属与挥发性风味的相关性如图5a所示，Lactobacillus、Acetobacter、Leuconostoc等是与风味物质相关性较高的细菌菌属。Lactobacillus与辛酸呈显著正相关（P<0.05），使酸奶具有焦糖香气，对酸奶的风味具有突出贡献。Acetobacter与乙酸、丁二醇、L（-）-乳酸乙酯和己酸呈较强的正相关（P<0.05），主要是因为Acetobacter在发酵过程中能够代谢乳酸，将其转化为乙酸和二氧化碳，能够丰富酸奶的风味层次。Leuconostoc与丁二醇、丁酸、乙酸和L（-）-乳酸乙酯呈正相关（P<0.05），与辛酸和苯甲酸呈负相关，Leuconostoc可与Lactobacillus和Streptococcus进行协同作用，共同影响发酵过程。Jiang等[4]也发现Lactobacillus与苯甲醛、2,3-戊二酮、乙醇和乙酸乙酯等挥发性化合物表现出显著相关性，而Streptococcus和Lactococcus丰度相对较高的样品表现出相对较低的挥发性化合物含量。真菌属与挥发性风味的相关性如图5b所示，Apiotrichum、Dipodascaceae、Pichia等是与风味物质相关性较高的真菌菌属。Kluyveromyces与己酸、乙酸、异戊醇和L（-）-乳酸乙酯呈负相关；Saccharomyces与丁二醇呈负相关（P<0.01）。而Dipodascaceae与己酸、丁酸、乙酸和丁二醇呈正相关（P<0.05）；Pichia与β-苯乙醇、甲酸乙烯酯和苯甲酸呈正相关（P<0.05），卢灏泽等[38]也发现Pichia是与西藏牦牛奶酪中风味物质相关性较高的真菌菌属。
由此可见，各类微生物的作用机制有所差异，它们之间形成了复杂的相互作用的生态网络，进而对酸奶样品中代谢产物与营养物质产生影响，形成了独特的风味。本研究揭示了西藏传统发酵乳中的微生物结构及代谢的关联，这为进一步基于假设的靶向研究与途径分析相结合提供了基础，这将有利于西藏传统发酵乳制品行业的产业化和标准化生产。
3　结论
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本研究对西藏不同地区的传统发酵乳中的微生物菌群组成和挥发性风味物质进行了综合分析。在13种酸奶样本中主要检测出了35种主要的挥发性风味成分，主要包括酯类14种、酸类10种、醇类4种、酮类3种、烷烃类3种和其他化合物1种。研究发现，地理位置对微生物多样性有着重要影响，不同地区的海拔、气候、土壤等因素共同塑造了微生物群落的结构和多样性。日喀则市在细菌和真菌的物种丰富度及多样性程度上都有较为突出的表现，可能是一个微生物资源丰富且具有特殊生态价值的地区；那曲市和阿里地区虽然在某些微生物指标上不是最突出的，但也具有独特的微生物群落特征，其中细菌以链球菌属（Streptococcus）为主导，真菌则以地霉属（Geotrichum）和克鲁维酵母属（Kluyveromyces）为主导，因此还需要进一步深入研究来探索其潜在的微生物资源和生态意义。此外，通过Spearman相关系数与OAV>1的特征风味物质进行相关性分析，与风味物质相关性较高的细菌菌属主要是Lactobacillus、Acetobacter、Leuconostoc；与风味物质相关性较高的真菌菌属主要是Apiotrichum、Dipodascaceae、Pichia。研究结果为开发功能性微生物发酵菌剂与发酵乳制品风味品质提供理论基础。

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2026年第47卷第12期
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doi: 10.13386/j.issn1002-0306.2025050095
  • 接收时间：2025-05-14
  • 首发时间：2026-10-22
  • 出版时间：2026-06-15
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  • 收稿日期：2025-05-14
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    1西北农林科技大学葡萄酒学院，陕西杨凌 712100
    2中国食品发酵工业研究院有限公司，北京 100125
    3国家酒类品质与安全国际联合研究中心，北京 100125
    4西藏自治区农牧科学院农产品开发与食品科学研究所，西藏拉萨 850030

通讯作者:

薛洁（1974−），女，博士，教授级高级工程师，研究方向：发酵工程，E-mail：。
刘延琳（1966−），女，博士，教授，研究方向：酿酒微生物，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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