食品工业科技

指标	对照组		补氮组
乙醇（%，v/v）	8.02±0.48^e	13.14±0.15^a	11.12±0.45^cd	12.03±0.17^ab	10.78±0.25^d	12.75±0.42^a	11.08±0.11^bc	11.79±0.23^ab
残糖（g/L）	43.97±3.52^a	3.00±0.32^c	2.07±1.41^c	2.83±0.30^c	10.59±2.15^b	3.06±1.36^c	2.16±0.29^c	1.92±0.04^c
甘油（g/L）	13.88±0.53^c	9.30±0.21^f	18.63±0.32^a	12.76±0.06^d	17.04±0.89^b	9.92±0.41^f	17.32±0.08^b	11.47±0.39^e
总酸（g/L）	5.16±0.11^bcd	5.30±0.10^bc	5.53±0.12^ab	4.72±0.36^de	4.93±0.15^cd	4.70±0.00^de	5.90±0.62^a	4.43±0.10^e
pH	3.51±0.01^c	3.56±0.01^b	3.53±0.01^c	3.53±0.01^c	3.65±0.15^a	3.66±0.02^a	3.57±0.01^b	3.66±0.01^a

指标	对照组		补氮组
乙醇（%，v/v）	8.02±0.48^e	13.14±0.15^a	11.12±0.45^cd	12.03±0.17^ab	10.78±0.25^d	12.75±0.42^a	11.08±0.11^bc	11.79±0.23^ab
残糖（g/L）	43.97±3.52^a	3.00±0.32^c	2.07±1.41^c	2.83±0.30^c	10.59±2.15^b	3.06±1.36^c	2.16±0.29^c	1.92±0.04^c
甘油（g/L）	13.88±0.53^c	9.30±0.21^f	18.63±0.32^a	12.76±0.06^d	17.04±0.89^b	9.92±0.41^f	17.32±0.08^b	11.47±0.39^e
总酸（g/L）	5.16±0.11^bcd	5.30±0.10^bc	5.53±0.12^ab	4.72±0.36^de	4.93±0.15^cd	4.70±0.00^de	5.90±0.62^a	4.43±0.10^e
pH	3.51±0.01^c	3.56±0.01^b	3.53±0.01^c	3.53±0.01^c	3.65±0.15^a	3.66±0.02^a	3.57±0.01^b	3.66±0.01^a

化合物	香气阈值（mg/L）^[32−34]	香气描述^[32−34]	对照组		补氮组
异丁醇	40.00	醇味	0.98	0.57	1.41	0.68	1.72	0.81	1.33	0.94
左旋-2,3-丁二醇	150.00	黄油味，奶酪味	0.21	0.19	0.21	0.2	0.16	0.18	0.18	0.16
正己醇	8.00	生青味	1.06	1.55	1.26	1.13	1.08	0.92	1.36	1.35
苯乙醇	10.00	花香，花粉味	1.18	1.34	0.95	0.73	1.61	0.86	1.11	0.71
乙酸乙酯	7.50	甜香，果香	6.05	3.76	5.75	3.29	7.68	6.08	6.19	3.69
乙酸异戊酯	0.20	香蕉香	0	6.76	1.15	0	0	7.02	2.24	0
乙酸苯乙酯	0.25	花香，果香，木香	0	0.24	0.2	0.21	0	0.2	0.2	0.22
丁酸乙酯	0.40	果香	0	0.4	0.11	0.25	0.1	0.32	0.23	0.38
己酸乙酯	0.01	菠萝香	0	11.24	0.16	3.08	0	16.95	0.38	2.93
辛酸乙酯	0.20	甜香，柑橘香	0.41	2.51	0.44	3.38	0.45	2.72	0.34	3.35
癸酸乙酯	0.20	果香，白兰地味	0.54	5.03	0.55	0.86	0.54	2.57	0.67	1.14

化合物	香气阈值（mg/L）^[32−34]	香气描述^[32−34]	对照组		补氮组
异丁醇	40.00	醇味	0.98	0.57	1.41	0.68	1.72	0.81	1.33	0.94
左旋-2,3-丁二醇	150.00	黄油味，奶酪味	0.21	0.19	0.21	0.2	0.16	0.18	0.18	0.16
正己醇	8.00	生青味	1.06	1.55	1.26	1.13	1.08	0.92	1.36	1.35
苯乙醇	10.00	花香，花粉味	1.18	1.34	0.95	0.73	1.61	0.86	1.11	0.71
乙酸乙酯	7.50	甜香，果香	6.05	3.76	5.75	3.29	7.68	6.08	6.19	3.69
乙酸异戊酯	0.20	香蕉香	0	6.76	1.15	0	0	7.02	2.24	0
乙酸苯乙酯	0.25	花香，果香，木香	0	0.24	0.2	0.21	0	0.2	0.2	0.22
丁酸乙酯	0.40	果香	0	0.4	0.11	0.25	0.1	0.32	0.23	0.38
己酸乙酯	0.01	菠萝香	0	11.24	0.16	3.08	0	16.95	0.38	2.93
辛酸乙酯	0.20	甜香，柑橘香	0.41	2.51	0.44	3.38	0.45	2.72	0.34	3.35
癸酸乙酯	0.20	果香，白兰地味	0.54	5.03	0.55	0.86	0.54	2.57	0.67	1.14

氮源调控与接种策略对杆状小星裂酵母与酿酒酵母混合发酵葡萄酒风味品质的影响

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钟轲 , 李蕊蕊 , 杨东岳 , 汤晓宏 , 李志宇 , 丁燕 , 韩晓梅 , 孙玉霞 ^*

食品工业科技 | 特邀主编专栏—食品风味感知前沿与应用（客座主编：刘源、孙金沅、寇兴然、程焕；产业顾问：陈丽） 2026,47(18): 35-43

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食品工业科技 |特邀主编专栏—食品风味感知前沿与应用（客座主编：刘源、孙金沅、寇兴然、程焕；产业顾问：陈丽） 2026 , 47 (18) : 35 -43

氮源调控与接种策略对杆状小星裂酵母与酿酒酵母混合发酵葡萄酒风味品质的影响

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钟轲（1996−），女，硕士，助理工程师，研究方向：葡萄酒酿造及酿酒微生物，E-mail：zk17854117992@163.com

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钟轲（1996−），女，硕士，助理工程师，研究方向：葡萄酒酿造及酿酒微生物，E-mail：zk17854117992@163.com

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钟轲, 李蕊蕊, 杨东岳, 汤晓宏, 李志宇, 丁燕, 韩晓梅, 孙玉霞^*

作者信息

山东省葡萄研究院山东省酿酒葡萄与葡萄酒技术创新中心，山东济南 250100

通讯作者:

孙玉霞（1973−），女，硕士，研究员，研究方向：葡萄酒果酒风味物质研究，E-mail：sunyuxia1230@163.com。

作者简介:

钟轲（1996−），女，硕士，助理工程师，研究方向：葡萄酒酿造及酿酒微生物，E-mail：zk17854117992@163.com

Effect of Nitrogen Source Regulation and Inoculation Strategy on the Flavor Quality of Starmerella bacillaris Mixed Fermented Wine

Ke ZHONG, Ruirui LI, Dongyue YANG, Xiaohong TANG, Zhiyu LI, Yan DING, Xiaomei HAN, Yuxia SUN^*

Affiliations

Shandong Academy of Grape Shandong Technology Innovation Center of Wine Grape and Wine, Jinan 250100, China

出版时间: 2026-09-15 doi: 10.13386/j.issn1002-0306.2025110250

文章导航

摘要

收起

为明确氮源管理和接种策略对杆状小星裂酵母（Starmerella bacillaris）与酿酒酵母（Saccharomyces cerevisiae）混合发酵葡萄酒风味品质的改善潜力，本研究评估了在两种可同化氮水平（160和300 mg/L）和三种接种方式（单独接种、与酿酒酵母同时接种和顺序接种）对葡萄酒发酵动力学、理化指标、香气成分及感官特性的影响。结果表明，氮源补充能促进酵母的糖代谢与发酵进程，并将挥发性组分总量提升3.23%~35.98%，其中高级醇与乙酸酯的升高较显著（P<0.05）。不同接种方式影响了葡萄酒的甘油含量、香气化合物的构成和感官表达。S. bacillaris的参与显著提高了甘油含量，可达11.47~18.63 g/L（P<0.05），提升了葡萄酒的甜润感与饱满度。主成分分析显示，S. cerevisiae单独发酵或与S. bacillaris同时接种时，葡萄酒以乙酸异戊酯、癸酸乙酯等多种酯类为主，果香特征明显；S. bacillaris单独发酵或与S. cerevisiae顺序接种时，葡萄酒中苯乙醇、乙酸乙酯、异丁醇等含量较高，带来花香与醇香。此外，补充氮源与顺序接种策略之间存在协同增效作用，在氮源充足的条件下进行顺序发酵，苯乙醇含量提升了29.73%，使酒体具有鲜明的玫瑰花香，同时在香气协调性与酒体结构完整性方面表现最优。感官评价证实，补氮结合顺序发酵能最大程度地优化愉悦香气并改善口感，整体品质最为突出。

关键词

杆状小星裂酵母 / 顺序接种 / 同时接种 / 可同化氮 / 挥发性香气化合物

Abstract

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To clarify the potential of nitrogen source management and inoculation strategies for improving the flavor quality of wine fermented by a mixture of Starmerella bacillaris (S. bacillaris) and Saccharomyces cerevisiae (S. cerevisiae), this study evaluated the effects of two levels of assimilable nitrogen (160 and 300 mg/L) and three inoculation methods (pure, simultaneous, and sequential inoculations of S. bacillaris and S. cerevisiaeon) on wine fermentation kinetics, physicochemical parameters, volatile composition, and sensory characteristics. Results showed that nitrogen source supplementation can significantly promote the sugar metabolism and fermentation process of yeast, and increase the total amount of volatile compounds by 3.23% to 35.98% (P<0.05), with notable increases in higher alcohols and acetates. Different inoculation methods affect the glycerol content, composition of aroma compounds, and sensory expression of wine. S. bacillaris significantly increased the glycerol content, reaching 11.47~18.63 g/L (P<0.05), enhancing the sweetness and fullness of the wine. Principal component analysis revealed that different inoculation strategies exhibit distinct aroma characteristics. Both S. cerevisiae pure culture and simultaneous inoculations were dominated by isoamyl acetate and ethyl decanoate, enhancing the fruity aroma of wine. Both S. bacillaris pure culture and sequential inoculations were dominated by phenylethyl alcohol, ethyl acetate and isobutanol, imparting floral and mellow aromas to wine. There was a significant synergistic effect between the supplementation of nitrogen sources and the sequential inoculation strategy. The sequential fermentation of the nitrogen-supplemented group was particularly outstanding in terms of aroma harmony and structural integrity, with a 29.73% increase in phenylethyl alcohol content, imparting a rose-like fragrance to the wine. Sensory evaluation indicated that nitrogen supplementation combined with sequential inoculation can significantly enhance the aroma and taste of wine, resulting in the best overall quality.

Key words

Starmerella bacillaris / sequential inoculation / simultaneous inoculation / assimilatable nitrogen / volatile aroma compounds

引用本文

钟轲, 李蕊蕊, 杨东岳, 汤晓宏, 李志宇, 丁燕, 韩晓梅, 孙玉霞. 氮源调控与接种策略对杆状小星裂酵母与酿酒酵母混合发酵葡萄酒风味品质的影响. 食品工业科技, 2026 , 47 (18) : 35 -43 . DOI: 10.13386/j.issn1002-0306.2025110250

Ke ZHONG, Ruirui LI, Dongyue YANG, Xiaohong TANG, Zhiyu LI, Yan DING, Xiaomei HAN, Yuxia SUN. Effect of Nitrogen Source Regulation and Inoculation Strategy on the Flavor Quality of Starmerella bacillaris Mixed Fermented Wine[J]. Science and Technology of Food Industry, 2026 , 47 (18) : 35 -43 . DOI: 10.13386/j.issn1002-0306.2025110250

正文

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酿酒酵母作为葡萄酒发酵的核心微生物，能够将葡萄汁中的糖类转化为乙醇、二氧化碳及其他次生代谢产物^[1−2]。为应对当前单一酵母发酵导致的葡萄酒同质化问题，采用非酿酒酵母与酿酒酵母的混合发酵策略已成为行业重要发展方向，非酿酒酵母在调控和增强葡萄酒香气结构与复杂性方面展现出独特潜力^[3]。例如，有孢汉逊酵母（Hanseniaspora spp.）和梅奇酵母（Metschnikowia spp.）能够合成酯类与高级醇^[4]；顺序接种戴尔有孢圆酵母（Torulaspora delbrueckii）与酿酒酵母，可提升葡萄酒的果香与复杂度，而异常毕赤酵母（Pichia anomala）与酿酒酵母混合发酵，则能增加3-甲基乙酸丁酯与乙基酯的含量^[5]。目前，戴尔有孢圆酵母、美极梅奇酵母（Metschnikowia pulcherrima）和克鲁维毕赤酵母（Pichia kluyveri）等部分非酿酒酵母已实现商业化，作为发酵剂应用于酿酒实践^[6]。杆状小星裂酵母（Starmerella bacillaris）最初于2003年从匈牙利托卡伊产区的贵腐葡萄中被分离，该菌株因独特代谢特性备受关注，其特点是能够耐受高糖、高酸，且高产甘油能增加葡萄酒的圆润度，在口感上增强对甜味的感知，同时低产挥发酸，保持葡萄酒的清新感和香气纯净度^[7−10]。研究表明，S. bacillaris与酿酒酵母的共培养可带来多方面风味提升包括提升乙酸异丁酯、异丁醇等风味物质含量^[9]；提高赤霞珠葡萄酒中己酸乙酯和乙酸己酯的浓度^[11]；提高长相思白葡萄酒中酯类和硫醇含量^[12]，以及提升巴贝拉葡萄酒中甘油和酯类的含量^[13]。在共发酵过程中，接种策略、菌株组合及酵母可同化氮（Yeast Assimilatable Nitrogen，YAN）等因素共同影响发酵动力学与挥发性有机物组成，其中YAN对酵母生长与代谢具有关键调控作用。

葡萄汁中的氮浓度对酵母的发酵效率及葡萄酒的香气特征具有关键影响。当可同化氮含量低于140~150 mg/L时，葡萄汁通常被视为氮源不足，可能导致酒精发酵迟缓甚至停滞，并促进硫化氢、甲硫醇和乙硫醇等不良还原性化合物的生成^[14−17]。为弥补氮的缺乏，酿酒师常在发酵过程中进行氮源补充。许多芳香化合物的合成与酵母的氮代谢密切相关^[18]。SU等^[17]发现，在模拟葡萄汁中添加铵态氮可提高乙酯产量；WANG等^[19]的研究表明，添加磷酸二铵会加快氮的消耗速率，但在总YAN较低而磷酸二铵比例较高时，反而会引起发酵迟缓。ENGLEZOS等^[20]发现，在葡萄酒发酵中，S. bacillaris对氨基酸的吸收速率较慢，而铵离子则被完全利用。此外，LOLA的研究显示，在氮添加量为300 mg/L的条件下，发酵所得的总萜烯含量是150 mg/L条件下的两倍^[18]。目前，关于酵母发酵过程中添加不同类型氮源的研究已较为丰富，但在不同接种策略下（如顺序接种、同时接种）如何优化氮源添加的研究仍相对有限。

在葡萄酒生产中，优化酵母接种策略与合理补充氮源是提升风味复杂度与感官品质的关键。本研究旨在探究可同化氮剂量对酿酒酵母与S. bacillaris混合发酵的影响，系统分析发酵动力学、香气组成及感官特性。结果表明，氮源补充能有效调控混合发酵进程，并塑造葡萄酒的芳香轮廓与感官特征，为优化发酵工艺提供了重要依据。

1　材料与方法

收起

1.1　材料与仪器

马瑟兰葡萄（糖240 g/L，YAN 160 mg/L，pH4.1，总酸4.26 g/L）　2024年9月采摘于山东烟台中粮长城酒厂的一个商业葡萄园；酵母菌株：商业酿酒酵母FX10　购自法国Laffort公司；杆状小星裂酵母S. bacillaris酵母（SIVE8902）　由本实验室筛选自山东烟台君顶酒庄蛇龙珠葡萄自然发酵醪中，保藏于国典型培养物保藏中心（CCTCC NO：M20241637）；氯化钠（分析纯）、偏重亚硫酸钾（分析纯）　上海麦克林生化科技有限公司；酵母浸出粉胨葡萄糖液体培养基（yeast extract peptone dextrose，YEPD）　青岛海博生物技术有限公司；二碳酸二甲酯（dimethyl dicarbonate，DMDC）　德国朗盛特殊化学品集团；果胶酶（酶活100 U/mg）、NUTRISTART™　购自法国Laffort公司；4-甲基-2-戊醇　色谱纯，美国Sigma公司；PRIMARY AMINO NITROGEN（PAN）试剂盒、AMMONIA（Rapid）试剂盒　爱尔兰Megazyme公司。

pHSJ-4A测定仪　上海科学仪器材料有限公司；BX-1手持折光计　德国SCHMIDT+HAENSCH公司；57329-U萃取头　美国Supelco公司；PAL RSI85自动进样器　瑞士CTC公司；Agilent1260 Infinity液相色谱仪、Agilent GC 7890B/MS 5977A气相色谱-质谱联用仪（gas chromatography mass spectrometry，GC-MS）、DB-WAX色谱柱（60 m×0.25 mm，0.25 μm）　美国Agilent公司。

1.2　实验方法

1.2.1　马瑟兰葡萄酒发酵

将采摘后的马瑟兰葡萄精心挑选，迅速运送到实验室进行酿酒实验。参考段亚茹等^[21]、刘红艳^[22]研究及欧洲食品安全局对DMDC在饮料中应用的安全标准（200~250 mg/L）^[23]，在马瑟兰葡萄去梗破碎后，添加240 mg/L DMDC，室温放置4 h以灭活天然酵母。于4 ℃浸渍过夜后过滤，将葡萄汁平均分为两组，参考LOLA等^[18]关于发酵氮需求的研究，将160 mg/L设为对照组（C组），代表避免发酵风险的最低营养边界；同时设置300 mg/L为补氮组（H组），模拟生产中经氮源补充后的充足状态，构建从临界至充足的清晰营养梯度。

取300 g葡萄汁于500 mL发酵瓶中开展发酵实验，实验具体流程与设计如下：接种前，将酵母菌株于30 ℃以200 r/min条件下振荡培养24 h进行活化。培养后的菌液经6000 r/min离心5 min收集菌体，并使用0.85%无菌NaCl溶液洗涤两次备用^[24]。采用血细胞计数法将菌悬液浓度调整至标准接种密度：S. bacillaris为1.0×10⁷ cells/mL，酿酒酵母FX10为1.0×10⁶ cells/mL，两者接种比例为10:1。依据不同接种方式设置以下四种处理组合，并在C、H两种氮水平下进行：SB：单菌接种杆状小星裂酵母S. bacillaris；SC：单菌接种酿酒酵母FX10；SE（顺序接种）：先接种杆状小星裂酵母S. bacillaris，48 h后再接种酿酒酵母FX10；SI（同时接种）：将两种酵母同时接入发酵基质。各组依“氮组-接种方式”分别标记为C-SB、C-SC、C-SE、C-SI及H-SB、H-SC、H-SE、H-SI。每组实验均设置三个生物学重复。

所有发酵均在25 ℃恒温下进行。通过每48 h称量发酵瓶的重量损失（反映CO₂释放量）来监测发酵进程。当连续两次称重损失小于0.1 g时，判定发酵结束。发酵结束后，样品于4 ℃、8000×g离心10 min，收集上清酒液并添加40 mg/L SO₂，置于4 ℃冷藏保存，供后续分析测定。

1.2.2　基本理化指标

参照汤晓宏等^[25]的研究方法，通过高效液相色谱法测定甘油。样品在分析前通过0.22 μm有机膜过滤，色谱工作条件：300×7.7 mm Hi-Plex H色谱柱，流动相为0.05 mol/L硫酸溶液，流速为0.6 mL/min。色谱柱温度为70 ℃，示差检测器检测温度为30 ℃，进样量为20 μL。精密称取0.2089 g甘油于10 mL容量瓶中，超纯水定容，即得20.89 g/L的标准溶液，按一定浓度做梯度稀释，制作标准曲线。乙醇、残糖、总酸参考GB/T 15038-2006《葡萄酒、果酒通用分析方法》。

1.2.3　可同化氮的测定

使用PRIMARY AMINO NITROGEN（PAN）试剂盒检测有机氮含量，使用AMMONIA（Rapid）试剂盒检测无机氮含量。

1.2.4　挥发性组分的测定

根据LI等^[26]的方法，通过顶空固相微萃取结合气相色谱-质谱（HS-SPME-GC-MS）对挥发性组分进行定量分析。色谱分析在配备有5977A MSD的Agilent 7890B气相色谱仪上进行。挥发性组分用VF-WAXms（30 m×0.32 mm，0.25 μm）毛细管气相色谱柱分离。以4-甲基-2-戊醇为内标（2.00 g/L样品溶液20 μL），使用构建的定量校准曲线计算浓度。对于缺乏标准品的挥发性组分，根据具有最相似化学结构或碳原子数的标准化合物的校准曲线估算浓度。具体操作：取5 mL葡萄酒样品，加入2.0 g氯化钠和20 μL 2.00 g/L的4-甲基-2-戊醇（内标物）置于15 mL顶空瓶中，放置于45 ℃的可加热磁力搅拌器上预热10 min，萃取50 min，萃取过程中顶空瓶在搅拌器中连续振荡，进样口250 ℃解吸10 min进样。结束后，色谱柱初始温度40 ℃，保持2 min，然后以6 ℃/min的速率升到230 ℃，保持15 min。载气为氦气，平均线速率为25 cm/s。采用不分流进样模式。所有质谱在电子电离源（EI，70eV），质量扫描范围为30~400 u，在每秒3.9次扫描速率条件下获得。香气活度值（odor activity value，OAV）计算公式如下：

$ \text{OAV=}\frac{\text{挥发性香气化合物质量浓度}}{\text{挥发性香气化合物香气阈值}} $

1.2.5　感官评定

感官评价参照国家标准GB/T 15038-2006《葡萄酒、果酒通用分析方法》并借鉴崔文文等^[27]描述的方法，做了一些修改。感官分析由10名专业的评估人员组成的感官小组进行（6名女性和4名男性，年龄在26~54岁之间，平均年龄为35.9岁）。以随机顺序将20 mL葡萄酒样品放入酒杯中呈现给评估人员。葡萄酒感官评价分为9个属性，包括花香、果香、甜味、草本、溶剂、强度、平衡、酸度和酒体。小组成员被要求使用9分制来衡量葡萄酒参数的强度（1=极低，5=中等强度，9=极高），每个感官属性的最终得分是10名评估者的平均值。

1.3　数据处理

所有指标均重复测定三次，采用SPSS 27.0软件进行数据统计分析Duncan’s多重比较检验，并采用Origin2018进行折线图、OAV分析图、PCA分析图绘制；CNSplot进行双形态堆积柱状图、雷达图绘制。

2　结果与分析

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2.1　不同氮浓度对酵母发酵动力学的影响

在酒精发酵过程中，二氧化碳的释放速率和糖的消耗速率是评估乙醇生产和发酵动力学的重要指标。图1显示了在两种氮浓度下采用不同接种策略的酵母发酵动力学。总体而言，除S. bacillaris单菌发酵外，其余样品在发酵至14 d残糖量均降至4 g/L以下，表明发酵基本完全。

在相同接种策略下，补氮组在发酵初期即表现出较快的糖消耗速率，而对照组则相对缓慢。发酵至2 d，对照组（C）的残糖含量介于154.00 mg/L（C-SI）至189.21 mg/L（C-SB）之间，而补氮组（H）的残糖范围为143 mg/L（H-SI）至172 mg/L（H-SB），显示出补氮对糖代谢的促进作用。该结果与LOLA等^[18]的研究一致，其报道指出在低氮条件（150 mg/L）发酵8 d后，残糖量比高氮组高出9.06%。尽管补充氮源可在一定程度上提升酵母发酵性能，S. bacillaris的单菌发酵仍未能实现糖的完全利用。ROCA-MESA等^[16]研究发现，无论在无机氮、有机氮或混合氮源条件下，S. bacillaris均无法独立完成全部发酵过程。

2.2　不同氮浓度下葡萄酒的基本理化指标

在不同接种方式与氮浓度条件下，所得葡萄酒的基本参数见表1。与酿酒酵母相比，非酿酒酵母的酒精发酵能力相对较差，难以独立完成发酵。在S. bacillaris单菌发酵中，补氮组（H-SB，10.78%）较对照组（C-SB，8.02%）显著（P<0.05）提升了酒精含量，同时降低了残糖含量（P<0.05），结果表明，氮源补充能提升S. bacillaris的发酵动力与糖代谢能力。甘油作为酵母在酒精发酵中的第二大代谢产物，能够增强葡萄酒的甜味感知与口感柔顺度，并有助于提升香气与风味的整体复杂性，从而改善葡萄酒质量^[28−29]。本研究中，S. bacillaris表现出较强的甘油合成能力，其纯培养条件下的甘油产量分别达13.88 g/L（C-SB）和17.04 g/L（H-SB）。进一步比较接种方式可见，顺序发酵（SE）的甘油产量较同步发酵（SI）显著提高，在对照组和补氮组中分别提升了46.00%和51.00%（P<0.05）。这可能是由于在顺序发酵的前48 h内，由高产甘油的S. bacillaris主导代谢，而同步发酵中该菌株难以与发酵力更强的酿酒酵母竞争，导致甘油合成受限。在酸度方面，补氮处理降低了除H-SE外所有样品的总酸含量，并相应提高了pH。

2.3　葡萄酒的挥发性成分

葡萄酒的香气与风味是评价其品质和区分风格的关键。在全部8款葡萄酒中共鉴定出33种挥发性组分，分别为9种高级醇、3种脂肪酸、4种乙酸酯、6种乙基酯、4种其他酯类、3种萜烯以及4种醛酮类化合物（附表1，详见本刊官网 http://www.spgykj.com/文章的网络版）。需要指出的是，并非所有检出的挥发性物质都对葡萄酒香气有实质影响。气味活性值（odor activity value，OAV）是评估单一成分在复杂混合物中香气贡献的关键指标^[30−31]。如表2所示，33种化合物中有11种的OAV>0.1，这些物质主要为高级醇和酯类。它们能够直接或通过协同作用塑造葡萄酒的整体香气轮廓，因此被认为是构成样品香气特征的主体成分^[9,30]。

如图2和附表1（详见本刊官网http://www.spgykj.com/文章的网络版），在相同接种方式下，补氮组（H组）挥发性组分总含量较对照组（C组）提高了3.23%（H-SE）至35.98%（H-SB），该增长主要源于高级醇与乙酸酯类含量的提升。高级醇在多数样品中占主导地位，作为酵母代谢的关键产物，其总量低于300 mg/L时可为葡萄酒带来甜润与果香，而过高则易产生杂醇味^[35−36]。本研究样品中高级醇总量均低于该阈值。

在香气贡献方面，共有四种高级醇的气味活性值（OAV）>0.1（表2和图3），分别为异丁醇（醇味）、正己醇（生青味）、左旋-2,3-丁二醇（黄油味，奶酪味）以及苯乙醇（花香，花粉味），它们是葡萄酒中主要呈香的高级醇。如图4所示，异丁醇含量最高，占高级醇总量的44.81%（C-SC）至69.53%（H-SB）。与对照组相比，补氮使异丁醇含量提高了3.90%（H-SE）至74.71%（H-SB）。除异丁醇外，含量最高的三种挥发性物质依次为左旋-2,3-丁二醇、苯乙醇和正丙醇。相较于对照组，补氮组左旋-2,3-丁二醇含量下降1.47%（H-SC）至22.48%（H-SB）。而在补氮条件下，H-SB和H-SE的苯乙醇含量分别较H-SC显著增加87.61%与29.73%（P<0.05），这说明S. bacillaris能赋予较高的玫瑰香气。综上，补氮处理提高了葡萄酒高级醇的含量，这可能是因为补充处理后，酵母通过Ehrlich途径代谢增强，促使更多氨基酸（如缬氨酸、亮氨酸和苯丙氨酸）经转氨生成α-酮酸，进一步脱羧、还原，转化为异丁醇、异戊醇和苯乙醇等高级醇^[37−38]。

脂肪酸通常与酵母的代谢特征密切相关，在葡萄酒中，低浓度的脂肪酸可赋予酒体脂肪与奶酪般的气息，而含量过高则往往对整体香气产生负面影响^[39]。如附表1（详见本刊官网http://www.spgykj.com/文章的网络版）所示，本研究共鉴定出三种脂肪酸（异丁酸、辛酸和己酸），其总含量范围为345.22 μg/L（H-SC）至697.58 μg/L（C-SE）。补氮处理中，H-SC和H-SE己酸含量分别降低了55.23%和20.11%。所有样品己酸含量均维持在其感官阈值以下，未产生脂肪或肥皂等不悦气味。

在本研究中共检出14种酯类化合物。与对照组相比，补氮处理的总酯含量提高了0.14%（H-SI）至21.62%（H-SB）。在乙酸酯方面，共检测到4种。补氮组总乙酸酯含量较对照组提升7.87%（H-SE）至58.53%（H-SC）。乙酸酯是由相应醇类与乙酰辅酶A在醇乙酰转移酶催化下缩合形成，补充氮源后可上调编码乙酸酯合成酶基因ATF1/2的表达，从而促进乙酸酯的生成^[40−41]。具体来看，乙酸乙酯和乙酸异丁酯分别增加7.58%（H-SE）至61.93%（H-SC）与6.40%（H-SE）至52.20%（H-SB）。如图3和表2所示，乙酸乙酯的OAV>1，可直接贡献果香；而OAV超过0.1的乙酸异戊酯（香蕉香）与乙酸苯乙酯（草莓香）也对葡萄酒香气具有重要作用。乙酸异丁酯、乙酸异戊酯和乙酸苯乙酯及其对应的高级醇前体（异丁醇、异戊醇、苯乙醇）均为Ehrlich途径相关氨基酸代谢所产生的主要挥发性物质^[37]。

乙基酯虽然在葡萄酒中浓度通常低于乙酸酯，但由于其种类多样且气味阈值较低，在增强香气复杂性方面具有关键作用^[24]。如图5和附表1（详见本刊官网http://www.spgykj.com/文章的网络版）所示，本研究检测到6种乙基酯，总含量介于1613.63 μg/L至4741.62 μg/L之间。乙基酯主要在乙醇乙酰转移酶催化下，由乙醇与酰基辅酶A合成，是构成葡萄酒果香酯类的关键成分^[42]。补氮对不同接种方式的影响差异较大：H-SB和H-SE中乙基酯总量分别下降43.36%和20.52%，而H-SC和H-SI则分别上升17.47%和6.33%。这一差异可能与S. bacillaris合成乙基酯能力较弱有关，其在顺序发酵和纯发酵前期占主导，导致整体乙基酯含量降低。己酸乙酯和辛酸乙酯具有果香与花香，因其阈值低，在微量下即可贡献香气；同时，OAV>0.1的丁酸乙酯（香蕉与草莓香）和癸酸乙酯（果香与愉悦气息）也对整体香韵具有重要影响。此外，研究还检出其他4种酯类，补氮组中该类酯总量及碳酸二甲酯的含量普遍高于对照组（H-SI除外）。

本研究共鉴定出三类萜烯类化合物（芳樟醇、香茅醇、D-柠檬烯）以及四类醛酮类化合物（2,6-二甲基-4-庚酮、苯甲醛、乙醛、异戊醛）。萜烯通常可为葡萄酒带来愉悦的花香与果香；而醛酮类物质多呈现苹果或坚果的香气，有助于丰富香气的层次感^[40,43]。在所有样品中，芳樟醇为含量最丰富的萜烯化合物，其存在为葡萄酒增添了令人愉悦的甜香调。

2.4　香气的PCA分析

主成分分析（Principal Component Analysis，PCA）被用于可视化11种OAV>0.1的挥发性有机物，并识别不同处理组间具有区分作用的关键香气成分（图6）。前两个主成分累计贡献了73.4%，其中PC1占55.0%，PC2占18.4%。PCA得分图显示，葡萄酒样品依接种方式被明显区分：SC与SI位于PC1正半轴，其主要贡献成分为酯类物质，包括乙酸异戊酯、癸酸乙酯、己酸乙酯、丁酸乙酯、乙酸苯乙酯和辛酸乙酯；而SB与SE则分布于PC1负半轴，其香气特征主要由苯乙醇、乙酸乙酯、异丁醇及左旋-2,3-丁二醇所主导。值得注意的是，在补氮条件下，H-SB与H-SE位于PC2正半轴，其香气轮廓主要受异丁醇、乙酸乙酯和苯乙醇的影响。这表明氮源补充能调控S. bacillaris纯发酵及顺序发酵过程中挥发性物质的组成，有助于增强葡萄酒中愉悦的果香与玫瑰香气。

2.5　感官评价

葡萄酒的感官评价由10名专业品评员完成。结果如图7所示，在两种氮浓度下，同步发酵与顺序发酵样品在愉悦香气和酒体结构方面评分普遍较高。与对照组相比，补氮处理进一步提升了顺序发酵与同步发酵葡萄酒的愉悦香气得分，该结果与样品中苯乙醇、乙酸乙酯、乙酸异戊酯、己酸乙酯及癸酸乙酯含量较高相符。在补氮组中，H-SE样品展现出浓郁的花香与突出的结构感，能改善葡萄酒的整体口感。得益于S. bacillaris高产甘油的特性，在相同氮源条件下，其单菌发酵、同步发酵及顺序发酵所产酒样的甜润度与饱满度均优于FX10单菌发酵，且酸度感知更低。此外，所有样品的溶剂与草本类香气评分较低，表明不良气味得到有效控制。综上，补充氮源有助于增强葡萄酒的花果香气，特别是在H-SE处理中，能够改善愉悦香气的复杂度与酒体结构完整性，从而提升葡萄酒的整体品质。

3　结论

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本研究系统解析了氮源水平与接种策略对S. bacillaris参与葡萄酒混合发酵风味形成的协同调控机制与效应。在发酵动力学方面，补充氮源（300 mg/L）显著提高了酵母的糖消耗速率与发酵效率，尤其促进了顺序发酵与同步发酵的顺利进行（P<0.05）。S. bacillaris虽无法独立完成酒精发酵，但其在纯培养及混菌体系中均表现出突出的甘油合成能力（11.47~18.63 g/L），为葡萄酒带来甜润与饱满口感。在香气组成上，补氮处理使挥发性组分总量提升了3.23%~35.98%，特别是增强了高级醇和乙酸酯类的合成。主成分分析进一步表明，不同接种方式具有鲜明的香气特征：SC与SI以酯类为主导，呈现浓郁果香；SB与SE则以高级醇为主，赋予酒样更丰富的花香与醇香。补氮组中的顺序发酵（H-SE）在酯类与高级醇之间达到良好平衡，香气表现尤为协调。感官评价表明，补氮处理提升了顺序发酵与同步发酵样品的愉悦香气与结构完整性，其中H-SE在花香强度、口感结构与整体接受度上均表现最优，表明该组合能有效提升葡萄酒的整体感官品质与风格完整性。综上，氮源补充与顺序发酵策略的结合，不仅优化了发酵动力学与香气组成，更通过协同调控甘油与挥发性代谢物的合成，有效提升了葡萄酒的感官品质与风格典型性，为相关酿造工艺的改进提供了理论依据与实践参考。在未来的研究中，可进一步结合转录组与代谢组学技术，深入解析S. bacillaris与酿酒酵母在氮源竞争利用中的分子互作机制，阐明氮调控下混菌发酵的关键代谢通路网络。同时，还需在中试及规模化生产上，系统考察该工艺在不同葡萄品种与酿造风格中的普适性与稳定性，以推动其在实际生产中的有效应用。

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2026年第47卷第18期

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doi: 10.13386/j.issn1002-0306.2025110250

接收时间：2025-11-24
首发时间：2026-10-24
出版时间：2026-09-15

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收稿日期：2025-11-24

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山东省葡萄研究院山东省酿酒葡萄与葡萄酒技术创新中心，山东济南 250100

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孙玉霞（1973−），女，硕士，研究员，研究方向：葡萄酒果酒风味物质研究，E-mail：sunyuxia1230@163.com。

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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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指标	对照组		补氮组
乙醇（%，v/v）	8.02±0.48^e	13.14±0.15^a	11.12±0.45^cd	12.03±0.17^ab	10.78±0.25^d	12.75±0.42^a	11.08±0.11^bc	11.79±0.23^ab
残糖（g/L）	43.97±3.52^a	3.00±0.32^c	2.07±1.41^c	2.83±0.30^c	10.59±2.15^b	3.06±1.36^c	2.16±0.29^c	1.92±0.04^c
甘油（g/L）	13.88±0.53^c	9.30±0.21^f	18.63±0.32^a	12.76±0.06^d	17.04±0.89^b	9.92±0.41^f	17.32±0.08^b	11.47±0.39^e
总酸（g/L）	5.16±0.11^bcd	5.30±0.10^bc	5.53±0.12^ab	4.72±0.36^de	4.93±0.15^cd	4.70±0.00^de	5.90±0.62^a	4.43±0.10^e
pH	3.51±0.01^c	3.56±0.01^b	3.53±0.01^c	3.53±0.01^c	3.65±0.15^a	3.66±0.02^a	3.57±0.01^b	3.66±0.01^a

指标

对照组

补氮组

C-SB

C-SC

C-SE

C-SI

H-SB

H-SC

H-SE

H-SI

乙醇（%，v/v）

8.02±0.48^e

13.14±0.15^a

11.12±0.45^cd

12.03±0.17^ab

10.78±0.25^d

12.75±0.42^a

11.08±0.11^bc

11.79±0.23^ab

残糖（g/L）

43.97±3.52^a

3.00±0.32^c

2.07±1.41^c

2.83±0.30^c

10.59±2.15^b

3.06±1.36^c

2.16±0.29^c

1.92±0.04^c

甘油（g/L）

13.88±0.53^c

9.30±0.21^f

18.63±0.32^a

12.76±0.06^d

17.04±0.89^b

9.92±0.41^f

17.32±0.08^b

11.47±0.39^e

总酸（g/L）

5.16±0.11^bcd

5.30±0.10^bc

5.53±0.12^ab

4.72±0.36^de

4.93±0.15^cd

4.70±0.00^de

5.90±0.62^a

4.43±0.10^e

3.51±0.01^c

3.56±0.01^b

3.53±0.01^c

3.65±0.15^a

3.66±0.02^a

3.57±0.01^b

3.66±0.01^a

化合物	香气阈值（mg/L）^[32−34]	香气描述^[32−34]	对照组		补氮组
异丁醇	40.00	醇味	0.98	0.57	1.41	0.68	1.72	0.81	1.33	0.94
左旋-2,3-丁二醇	150.00	黄油味，奶酪味	0.21	0.19	0.21	0.2	0.16	0.18	0.18	0.16
正己醇	8.00	生青味	1.06	1.55	1.26	1.13	1.08	0.92	1.36	1.35
苯乙醇	10.00	花香，花粉味	1.18	1.34	0.95	0.73	1.61	0.86	1.11	0.71
乙酸乙酯	7.50	甜香，果香	6.05	3.76	5.75	3.29	7.68	6.08	6.19	3.69
乙酸异戊酯	0.20	香蕉香	0	6.76	1.15	0	0	7.02	2.24	0
乙酸苯乙酯	0.25	花香，果香，木香	0	0.24	0.2	0.21	0	0.2	0.2	0.22
丁酸乙酯	0.40	果香	0	0.4	0.11	0.25	0.1	0.32	0.23	0.38
己酸乙酯	0.01	菠萝香	0	11.24	0.16	3.08	0	16.95	0.38	2.93
辛酸乙酯	0.20	甜香，柑橘香	0.41	2.51	0.44	3.38	0.45	2.72	0.34	3.35
癸酸乙酯	0.20	果香，白兰地味	0.54	5.03	0.55	0.86	0.54	2.57	0.67	1.14

化合物

香气阈值（mg/L）^[32−34]

香气描述^[32−34]

对照组

补氮组

C-SB

C-SC

C-SE

C-SI

H-SB

H-SC

H-SE

H-SI

异丁醇

40.00

醇味

0.98

0.57

1.41

0.68

1.72

0.81

1.33

0.94

左旋-2,3-丁二醇

150.00

黄油味，奶酪味

0.21

0.19

0.21

0.2

0.16

0.18

0.16

正己醇

8.00

生青味

1.06

1.55

1.26

1.13

1.08

0.92

1.36

1.35

苯乙醇

10.00

花香，花粉味

1.18

1.34

0.95

0.73

1.61

0.86

1.11

0.71

乙酸乙酯

7.50

甜香，果香

6.05

3.76

5.75

3.29

7.68

6.08

6.19

3.69

乙酸异戊酯

0.20

香蕉香

6.76

1.15

7.02

2.24

乙酸苯乙酯

0.25

花香，果香，木香

0.24

0.2

0.21

0.2

0.22

丁酸乙酯

0.40

果香

0.4

0.11

0.25

0.1

0.32

0.23

0.38

己酸乙酯

0.01

菠萝香

11.24

0.16

3.08

16.95

0.38

2.93

辛酸乙酯

0.20

甜香，柑橘香

0.41

2.51

0.44

3.38

0.45

2.72

0.34

3.35

癸酸乙酯

0.20

果香，白兰地味

0.54

5.03

0.55

0.86

0.54

2.57

0.67

1.14