微生物学报

Name	Main origin	Main ingredients	Main fermented products	Main microorganisms
Daqu^[4,6,14]	China	Wheat, barley, pea	Daqu Baijiu, vinegar	Aspergillus, Mucor, Rhizopus, Lichtheimia, Thermomyces, Rhizomucor, Monascus, Pichia, Saccharomycopsis, Bacillus, Lactobacillus, Weissella, Thermoactinomyces, Kroppenstedtia, Saccharopolyspora
Xiaoqu^[6,14]	China	Rice, soybean	Xiaoqu Baijiu, Huangjiu	Aspergillus, Rhizopus, Saccharomycopsis, Pichia, Candida, Lactobacillus, Pediococcus, Weissella
Fuqu^[6,14]	China	Wheat bran	Fuqu Baijiu	Aspergillus, Rhizopus, Saccharomyces, Bacillus
Koji^[15]	Japan	Rice, barley	Saké (fermented wine), Miso (fermented soybean paste), Shoyu (soy sauce)	Aspergillus, Wickerhamomyces, Candida, Ochrobactrum, Lactobacillus
Nuruk^[16]	Korea	Wheat, rice	Makgeolli (fermented wine), Yakju (fermented wine), Soju (distilled liquor)	Aspergillus, Rhizomucor, Lichtheimia, Mucor, Saccharomycopsis, Pichia, Debaryomyces
Loog-pang^[17]	Thailand	Rice, herb, spice	Khao-maak (alcoholic food), Sato (fermented wine)	Amylomyces, Rhizopus, Aspergillus, Mucor, Absidia, Saccharomycopsis, Pichia, Saccharomyces, Pediococcus
Bánh men^[18]	Vietnam	Rice, herb, spice	Ruou nep than (fermented wine), Ruou de (distilled liquor)	Rhizopus, Absidia, Amylomyces, Saccharomycopsis, Saccharomyces, Issatchenkia, Pediococcus, Lactobacillus, Weissella, Bacillus
Bubod^[17]	Philippines	Rice, herb	Basi (fermented wine)	Mucor, Rhizopus, Saccharomyces, Saccharomycopsis
Hamei^[19-20]	India	Rice, herb	Atingba (fermented wine), Yu (distilled liquor)	Mucor, Rhizopus, Penicillium, Saccharomyces, Pichia, Trichosporon
Xaj-pitha^[21]	India	Rice, herb	Rohi (fermented wine), Xaj (fermented wine)	Rhizopus, Mucor, Meyerozyma, Wickerhamomyces, Saccharomyces, Candida, Lactobacillus, Leuconostoc, Weissella
Marcha^[17,19]	India, Bhutan, Nepal, Xizang of China	Rice, herb, spice	Kodo ko jaanr (alcoholic beverage), Bhaati jaanr (alcoholic beverage), Raksi (distilled liquor)	Aspergillus, Mucor, Rhizopus, Penicillium, Bjerkandera, Saccharomycopsis, Pichia, Pediococcus
Phab^[17]	India, Bhutan, Nepal, Xizang of China	Wheat, herb	Chhang (alcoholic beverage)	Penicillium, Aspergillus, Saccharomycopsis
Ragi^[17]	Indonesia	Rice, millet, cassava, herb, spice	Tapé (alcoholic food)	Rhizopus, Mucor, Amylomyces, Aspergillus, Saccharomycopsis, Candida, Saccharomyces, Pichia, Pediococcus

Name	Main origin	Main ingredients	Main fermented products	Main microorganisms
Daqu^[4,6,14]	China	Wheat, barley, pea	Daqu Baijiu, vinegar	Aspergillus, Mucor, Rhizopus, Lichtheimia, Thermomyces, Rhizomucor, Monascus, Pichia, Saccharomycopsis, Bacillus, Lactobacillus, Weissella, Thermoactinomyces, Kroppenstedtia, Saccharopolyspora
Xiaoqu^[6,14]	China	Rice, soybean	Xiaoqu Baijiu, Huangjiu	Aspergillus, Rhizopus, Saccharomycopsis, Pichia, Candida, Lactobacillus, Pediococcus, Weissella
Fuqu^[6,14]	China	Wheat bran	Fuqu Baijiu	Aspergillus, Rhizopus, Saccharomyces, Bacillus
Koji^[15]	Japan	Rice, barley	Saké (fermented wine), Miso (fermented soybean paste), Shoyu (soy sauce)	Aspergillus, Wickerhamomyces, Candida, Ochrobactrum, Lactobacillus
Nuruk^[16]	Korea	Wheat, rice	Makgeolli (fermented wine), Yakju (fermented wine), Soju (distilled liquor)	Aspergillus, Rhizomucor, Lichtheimia, Mucor, Saccharomycopsis, Pichia, Debaryomyces
Loog-pang^[17]	Thailand	Rice, herb, spice	Khao-maak (alcoholic food), Sato (fermented wine)	Amylomyces, Rhizopus, Aspergillus, Mucor, Absidia, Saccharomycopsis, Pichia, Saccharomyces, Pediococcus
Bánh men^[18]	Vietnam	Rice, herb, spice	Ruou nep than (fermented wine), Ruou de (distilled liquor)	Rhizopus, Absidia, Amylomyces, Saccharomycopsis, Saccharomyces, Issatchenkia, Pediococcus, Lactobacillus, Weissella, Bacillus
Bubod^[17]	Philippines	Rice, herb	Basi (fermented wine)	Mucor, Rhizopus, Saccharomyces, Saccharomycopsis
Hamei^[19-20]	India	Rice, herb	Atingba (fermented wine), Yu (distilled liquor)	Mucor, Rhizopus, Penicillium, Saccharomyces, Pichia, Trichosporon
Xaj-pitha^[21]	India	Rice, herb	Rohi (fermented wine), Xaj (fermented wine)	Rhizopus, Mucor, Meyerozyma, Wickerhamomyces, Saccharomyces, Candida, Lactobacillus, Leuconostoc, Weissella
Marcha^[17,19]	India, Bhutan, Nepal, Xizang of China	Rice, herb, spice	Kodo ko jaanr (alcoholic beverage), Bhaati jaanr (alcoholic beverage), Raksi (distilled liquor)	Aspergillus, Mucor, Rhizopus, Penicillium, Bjerkandera, Saccharomycopsis, Pichia, Pediococcus
Phab^[17]	India, Bhutan, Nepal, Xizang of China	Wheat, herb	Chhang (alcoholic beverage)	Penicillium, Aspergillus, Saccharomycopsis
Ragi^[17]	Indonesia	Rice, millet, cassava, herb, spice	Tapé (alcoholic food)	Rhizopus, Mucor, Amylomyces, Aspergillus, Saccharomycopsis, Candida, Saccharomyces, Pichia, Pediococcus

菌种 Strains	来源 Sources	应用 Applications	主要结果与影响(相对于对照组) Main results and effects (compared with the control group)
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲 Fortifying Daqu^[71]	(1) Bacillus、Clavispora和Aspergillus丰度增加，Pichia、Saccharomycopsis丰度降低 (1) The abundances of Bacillus, Clavispora, and Aspergillus increased, while those of Pichia and Saccharomycopsis decreased (2) 液化力、糖化力显著增加，酯化力显著降低 (2) The liquefying power and saccharifying power increased significantly, while the esterifying power decreased significantly (3) 吡嗪类和芳香族类化合物含量增加 (3) The contents of pyrazines and aromatic compounds increased
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[72]	(1) 前期Lactobacillus基因转录量降低，后期Kazachstania、Naumovozyma、Saccharomyces基因转录量增加 (1) The gene transcription level of Lactobacillus decreased in the early stage, while those of Kazachstania, Naumovozyma, and Saccharomyces increased in the later stage (2) 后期Saccharomyces乙醇脱氢酶、乙醛脱氢酶和乳酸脱氢酶基因转录量增加 (2) The gene transcription levels of alcohol dehydrogenase, aldehyde dehydrogenase, and lactate dehydrogenase of Saccharomyces increased in the later stage
Bacillus subtilis Bacillus velezensis	大曲 Daqu	强化大曲 Fortifying Daqu^[73]	(1) Bacillus、Lactobacillus和Candida丰度增加 (1) The abundances of Bacillus, Lactobacillus, and Candida increased (2) 液化力、糖化力、酯化力显著增加 (2) The liquefying power, saccharifying power, and esterifying power increased significantly (3) 酯类、吡嗪类和醇类化合物含量显著增加 (3) The contents of esters, pyrazines, and alcohols increased significantly
Monascus purpureus Aspergillus oryzae Rhizopus arrhizus	中温大曲 Medium-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[74]	(1) 酒精度提高8% (1) The alcohol content increased by 8% (2) 风味物质数量和含量均增加 (2) Both the quantity and content of flavor substances increased
Rhizopus oryzae Bacillus velezensis	中温大曲 Medium-temperature Daqu	强化麸曲 Fortifying Fuqu^[75]	(1) 通过平板对峙实验，确认两者无拮抗作用 (1) Through the plate confrontation experiment, it was confirmed that there was no antagonistic effect between the two strains (2) 通过单因素和响应面实验，确定复合麸曲的最佳制备工艺 (2) Through single-factor and response surface experiments, the optimal preparation process of the composite Fuqu was determined
Bacillus amyloliquefaciens Saccharomycopsis fibuligera Absidia corymbifera	低温大曲 Low-temperature Daqu	强化大曲 Fortifying Daqu^[76]	(1) 淀粉酶活力提高 (1) The amylase activity increased (2) 微生物群落结构差异较小，丰富度提高 (2) There was little difference in the microbial community structure, and the richness increased (3) 真菌强化组中的乙醇、异丁醇、异戊醇含量增加 (3) The contents of ethanol, isobutanol, and isoamyl alcohol in the fungi-fortified group increased

菌种 Strains	来源 Sources	应用 Applications	主要结果与影响(相对于对照组) Main results and effects (compared with the control group)
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲 Fortifying Daqu^[71]	(1) Bacillus、Clavispora和Aspergillus丰度增加，Pichia、Saccharomycopsis丰度降低 (1) The abundances of Bacillus, Clavispora, and Aspergillus increased, while those of Pichia and Saccharomycopsis decreased (2) 液化力、糖化力显著增加，酯化力显著降低 (2) The liquefying power and saccharifying power increased significantly, while the esterifying power decreased significantly (3) 吡嗪类和芳香族类化合物含量增加 (3) The contents of pyrazines and aromatic compounds increased
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[72]	(1) 前期Lactobacillus基因转录量降低，后期Kazachstania、Naumovozyma、Saccharomyces基因转录量增加 (1) The gene transcription level of Lactobacillus decreased in the early stage, while those of Kazachstania, Naumovozyma, and Saccharomyces increased in the later stage (2) 后期Saccharomyces乙醇脱氢酶、乙醛脱氢酶和乳酸脱氢酶基因转录量增加 (2) The gene transcription levels of alcohol dehydrogenase, aldehyde dehydrogenase, and lactate dehydrogenase of Saccharomyces increased in the later stage
Bacillus subtilis Bacillus velezensis	大曲 Daqu	强化大曲 Fortifying Daqu^[73]	(1) Bacillus、Lactobacillus和Candida丰度增加 (1) The abundances of Bacillus, Lactobacillus, and Candida increased (2) 液化力、糖化力、酯化力显著增加 (2) The liquefying power, saccharifying power, and esterifying power increased significantly (3) 酯类、吡嗪类和醇类化合物含量显著增加 (3) The contents of esters, pyrazines, and alcohols increased significantly
Monascus purpureus Aspergillus oryzae Rhizopus arrhizus	中温大曲 Medium-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[74]	(1) 酒精度提高8% (1) The alcohol content increased by 8% (2) 风味物质数量和含量均增加 (2) Both the quantity and content of flavor substances increased
Rhizopus oryzae Bacillus velezensis	中温大曲 Medium-temperature Daqu	强化麸曲 Fortifying Fuqu^[75]	(1) 通过平板对峙实验，确认两者无拮抗作用 (1) Through the plate confrontation experiment, it was confirmed that there was no antagonistic effect between the two strains (2) 通过单因素和响应面实验，确定复合麸曲的最佳制备工艺 (2) Through single-factor and response surface experiments, the optimal preparation process of the composite Fuqu was determined
Bacillus amyloliquefaciens Saccharomycopsis fibuligera Absidia corymbifera	低温大曲 Low-temperature Daqu	强化大曲 Fortifying Daqu^[76]	(1) 淀粉酶活力提高 (1) The amylase activity increased (2) 微生物群落结构差异较小，丰富度提高 (2) There was little difference in the microbial community structure, and the richness increased (3) 真菌强化组中的乙醇、异丁醇、异戊醇含量增加 (3) The contents of ethanol, isobutanol, and isoamyl alcohol in the fungi-fortified group increased

大曲糖化剂在酿酒工业中的应用及其糖化功能研究进展

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杨阳 , 马吾霞 , 刘晓彤 , 毛雪婷 , 汤涵岚 , 杨儒洁 , 胡永芯 , 汪茜 , 沈才洪 , 王松涛

微生物学报 | 综述 2025,65(10): 4357-4373

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微生物学报 | 综述 2025, 65(10): 4357-4373

大曲糖化剂在酿酒工业中的应用及其糖化功能研究进展

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杨阳, 马吾霞, 刘晓彤, 毛雪婷, 汤涵岚, 杨儒洁, 胡永芯, 汪茜, 沈才洪, 王松涛

作者信息

泸州品创科技有限公司，国家固态酿造工程技术研究中心，四川泸州

Application of Daqu as a saccharification agent in the brewing industry and research advances in its saccharifying function

Yang YANG, Wuxia MA, Xiaotong LIU, Xueting MAO, Hanlan TANG, Rujie YANG, Yongxin HU, Qian WANG, Caihong SHEN, Songtao WANG

Affiliations

National Engineering Research Center of Solid-state Brewing, Luzhou Pinchuang Technology Co. , Ltd. , Luzhou, Sichuan, China

出版时间: 2025-09-04 doi: 10.13343/j.cnki.wsxb.20250241

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摘要

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作为中国传统酿酒工艺的核心糖化剂，大曲凭借其复杂的微生物群落和多功能酶系在白酒生产中发挥着不可替代的作用。本文概述了东西方酿酒工业中糖化剂的应用场景与特点，进而基于微生物学、基因组学和蛋白质组学等方面的研究重点阐述了大曲在糖化功能微生物及酶多样性、糖化机制等方面的特性，并提出了运用多组学技术揭示大曲在酿酒过程中的功能机制、深化机理研究与生产实践相结合等策略，为酿酒工业中糖化剂的创新发展提供了参考。

关键词

大曲 / 糖化剂 / 酿酒工业 / 多组学 / 微生物群落 / 功能

Abstract

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As the core saccharification agent in traditional Chinese brewing processes, Daqu plays an irreplaceable role in Baijiu production due to its complex microbial community and multifunctional enzyme system. This review begins by outlining the application scenarios and characteristics of saccharification agents in both Eastern and Western brewing industries. Subsequently, based on the research of microbiology, genomics, proteomics, etc., this review elaborates on the diversity of microorganisms and enzymes with saccharifying function in Daqu, along with the roles of Daqu in saccharification. Finally, strategic approaches are proposed, including utilizing multi-omics technologies to elucidate the functional mechanisms of Daqu during brewing and enhancing the integration of mechanism research with industrial practice, which may provide references for the innovative development of saccharification agents in the brewing industry.

Key words

Daqu / saccharification agent / brewing industry / multi-omics / microbial community / function

引用本文

杨阳, 马吾霞, 刘晓彤, 毛雪婷, 汤涵岚, 杨儒洁, 胡永芯, 汪茜, 沈才洪, 王松涛. 大曲糖化剂在酿酒工业中的应用及其糖化功能研究进展. 微生物学报, 2025 , 65 (10) : 4357 -4373 . DOI: 10.13343/j.cnki.wsxb.20250241

Yang YANG, Wuxia MA, Xiaotong LIU, Xueting MAO, Hanlan TANG, Rujie YANG, Yongxin HU, Qian WANG, Caihong SHEN, Songtao WANG. Application of Daqu as a saccharification agent in the brewing industry and research advances in its saccharifying function[J]. Acta Microbiologica Sinica, 2025 , 65 (10) : 4357 -4373 . DOI: 10.13343/j.cnki.wsxb.20250241

正文

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糖化作用是一类关键的生化反应，涉及淀粉、纤维素、半纤维素等多糖类物质逐步水解为寡糖、单糖的过程^[1-2]，该过程可由不同类型糖化功能酶催化完成。这类反应不仅为微生物生长提供了发酵基质，还促进了后续其他生化反应的进行，因此在酿造工业中得到了广泛应用^[3]。

糖化剂的使用历史悠久，可追溯至人类早期文明，至今仍在现代酿酒和食品加工领域占据重要地位。大曲作为东方传统糖化发酵剂的代表，凭借其复杂的微生物群落和丰富的酶系，在糖化、发酵和风味形成等关键环节展现出显著优势^[4]。尽管大曲在发酵和风味形成等方面的研究日益丰富，其糖化功能的相关研究仍主要集中在糖化力和液化力2类基础指标的检测上^[5]。对于其深层次机制和功能，以及除淀粉以外多糖(如纤维素、半纤维素、果胶等)的广义糖化功能，目前尚缺乏系统探讨及综述总结。

近年来，随着现代组学技术的快速发展，大曲研究取得显著进展，揭示了其在微生物组成和酶系功能等方面的独特优势^[6]。基于此，本文从酿酒工业糖化剂的整体应用背景出发，系统探讨大曲糖化剂在酿酒工业中的应用特点，并以微生物组成及功能为切入点，深入综述其糖化功能研究的最新进展，以期为传统糖化剂的优化和升级提供理论依据。

1 酿酒工业中的糖化剂

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1.1 糖化剂的应用场景及特点

全球酒类品种繁多，制作原料丰富，但其基本原理均为利用微生物将可发酵糖转化为乙醇。如图1所示，根据原料是否经过糖化过程，酿酒模式可分为单式发酵和复式发酵，后者根据糖化和发酵是否同时进行可细分为单行复式发酵和并行复式发酵^[7]。

单式发酵的典型例子包括葡萄酒和苹果酒，这类酒的制作不涉及糖化步骤，可发酵糖直接来源于成熟果实。用水果制作的白兰地、以龙舌兰根茎酿造的龙舌兰酒以及由糖蜜加工制成的朗姆酒，也都属于先单式发酵再经蒸馏制成的酒类产品。单行复式发酵常见于啤酒生产中，该过程需先使用麦芽糖化剂促进原料糖化，之后人工接种酵母进行发酵。类似地，在威士忌、伏特加、金酒的酿造过程中也普遍采用麦芽进行谷物的酶解，糖化与发酵环节界限分明。

白酒和黄酒是中国传统酒文化的瑰宝，也是并行复式发酵的代表。先民们经过千百年的生产实践，塑造出“以曲行酿”的东方酿酒形态。传统酒曲以谷物经自然发酵制成，兼具糖化、促酵、投粮、生香等功能^[4,6,8]，被誉为“中国第五大发明”和“酒之骨”。白酒和黄酒均以蒸煮的粮食与水为原料，用酒曲作为糖化发酵剂，糖化反应和乙醇发酵同步进行。这种酿酒模式能创造出风味独特、口感丰富且市场接受度高的酒精饮料产品。然而，由于制曲和酿酒这两大核心环节具有半开放式和自然发酵的特性，产品的品质及产量容易受到地域、气候、季节等环境因素的影响^[6,9]。

1.2 麦芽糖化剂及糖化酶剂

麦芽糖化剂是单行复式发酵中的关键辅助物质，通常以大麦为主要原料^[10]。麦芽制作是一个促使谷物种子强制萌芽的过程。首先，干燥谷物经过预除杂处理后反复浸泡2-4次以充分吸收水分。随后，胚芽开始恢复代谢活动并合成赤霉酸。赤霉酸的扩散促进多类水解酶的生物合成和释放，这些酶的合成数量与赤霉酸水平及糊粉层对激素的响应度相关，直接影响着麦芽的酿造性能^[11]。当胚乳中的水解酶和可溶性糖积累到一定程度后采用分步式烘干法对麦芽进行处理，即先在50-60 ℃烘干，然后逐步升温至65-75 ℃，以尽可能减少酶的变性^[12]。接下来，麦芽需先被粉碎并置于热水中，再与糊化后的淀粉质原辅料混合。糖化结束后还需经过滤、淋洗、整合、煮沸、冷却等一系列操作，最终加入酵母进行酿酒。麦芽糖化剂中的一系列水解酶负责将谷物中的淀粉和非淀粉多糖分解。淀粉水解酶包括α-淀粉酶、β-淀粉酶、极限糊精酶和α-葡萄糖苷酶等，与糖化力的呈现有关；非淀粉多糖水解酶主要包括内切木聚糖酶、α-l-阿拉伯呋喃糖苷酶等阿拉伯木聚糖酶和β-1,3-1,4-葡聚糖酶等β-葡聚糖酶，与麦芽的易碎性和麦汁的黏稠性相关^[12]。

在现代酿酒工业中α-淀粉酶、葡萄糖淀粉酶等糖化酶剂逐渐成为麦芽糖化剂的重要补充，它们能高效地将淀粉分解为可发酵糖，提高糖化效率和酒精产量，弥补现有糖化剂酶系在某些条件下的不足(如高温、低pH等)^[13]。尽管如此，糖化酶剂仅具有单一的糖化功能，无法像传统糖化剂那样为生产过程赋予额外的复杂风味。

1.3 多菌混合糖化剂

在西方酿酒工业中，麦芽糖化过程中的菌群和酶系活动通常会因煮沸环节而终止。在中国及其他亚洲国家的传统酿造实践中，当人们利用谷物等粮食制作含酒精的饮料和食品时常采用经固态发酵技术制备的“多菌多酶”混合培养物作为糖化发酵剂，具有鲜明的民族特色(表1)。

尽管某些多菌混合糖化剂的生产已从手工规模扩展至工业级，但依然遵循自然发酵的工艺框架。这些糖化剂主要以小麦或大米为原料，从环境中自然富集霉菌、酵母、乳酸菌等多种微生物，制成干燥而致密的块状、球状、饼状或不规则状成品，展现出良好的储存稳定性^[17]。为了稳定糖化剂的制备质量，有时也会采用之前制备的成品进行接种。中国的大曲和小曲、印度的Marcha、泰国的Loog-pang、韩国的Nuruk、菲律宾的Budod和越南的Bánh men等均属于这类传统糖化剂的范畴。

酒曲在中国拥有悠久的历史，但直到1892年，法国著名细菌学家和免疫学家Albert Calmette才首次对其菌群进行了系统的科学研究。他在中国的地方曲中分离出包括淀粉霉属(Amylomyces)、毛霉属(Mucor)、曲霉属(Aspergillus)以及野生酵母和细菌在内的多种微生物^[17]，并将其中一株糖化能力强的鲁氏淀粉霉(Amylomyces rouxii)用于液态法酿酒，开创了以纯种发酵为基础的酿酒生产路线，即阿米露法(amylo process)^[22]。后来，在日本Koji和中国麸曲等糖化剂制备过程中也采用了纯菌接种技术，体现了传统酿造工艺的现代化尝试^[23-24]。尽管现代化糖化剂具备发酵周期短、成本低等优势，但传统糖化剂因其在生产成分复杂、风味丰富的酒精饮料和发酵食品上具有明显优势，在许多地区仍然备受重视和广泛应用。

1.4 大曲糖化剂

由于原料和制作工艺的不同，中国酒曲主要被分为大曲、小曲和麸曲3大类，它们在白酒酿造中均能发挥糖化发酵剂的作用^[6]。其中，大曲微生物种类最为丰富，虽然其出酒率偏低，但由它参与酿造的白酒产品质量最佳。在1952-1989年期间，中国5届名酒评选共选出17种国家级名白酒，全部为大曲白酒，由此可见大曲在中国酒曲大家族中的重要地位。准确解析大曲的功能属性并有效地扬长避短，对中国白酒的传承和创新具有重要意义。

在大曲发酵过程中微生物代谢活动产生的热量对生境温度的变化起着关键作用，介导了其“前缓、中挺、后缓落”的变化模式^[25]。如图2所示，根据曲心发酵的最高温度(顶温)，大曲可分为低温大曲(顶温为40-50 ℃)、中温大曲(顶温为50-60 ℃)和高温大曲(顶温为60-65 ℃) 3种类型^[26]。顶温差异主要受安曲密度、覆盖方式、强制通风、人工翻曲等因素影响。低温大曲以大麦、豌豆为主要原料，发酵周期约为28 d，主要用于酿造清香型白酒；中温大曲以小麦为主要原料，发酵周期约为30 d，主要用于酿造浓香型白酒；高温大曲同样以小麦为主要原料，发酵周期常为30-60 d，主要用于酿造酱香型白酒^[6,27]。

2 大曲微生物组成及功能研究基础

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2.1 大曲微生物组成研究基础

经过数十天的自然发酵过程大曲中富集了大量微生物，这在很大程度上塑造了大曲在后续酿酒中的糖化发酵特性。深入了解大曲微生物群落物种组成及其种群之间的关系，一直是大曲研究的重要内容。图3列举了目前大曲菌群研究的主要技术、研究重点及相关应用方向。

研究者采用常规培养、选择培养、差异培养等技术，结合形态学、生理生化及分子测序等鉴定手段，从大曲中分离获得多种微生物的纯培养，其中不乏大曲高温放线菌(Thermoactinomyces daqus)、大曲热黄微菌(Thermoflavimicrobium daqui)、大曲土壤芽孢杆菌(Solibacillus daqui)等多个新种^[28-30]，凸显大曲菌库资源的丰富性。在细菌中主要的可培养类群包括芽孢杆菌属(Bacillus)和乳杆菌属(Lactobacillus)；在酵母中可培养类群以复膜孢酵母菌属(Saccharomycopsis)、威克汉姆酵母菌属(Wickerhamomyces)和毕赤酵母菌属(Pichia)为主；在丝状真菌方面则以Aspergillus和根霉菌属(Rhizopus)为主^[31-33]。

传统培养技术仅能捕捉到环境中不足1%的微生物^[34]，这种低覆盖率限制了对大曲群落组成的全面理解。近年来，扩增子测序、宏基因组、宏转录组等现代研究手段的发展将大曲微生态研究推向高通量模式和系统层面。多项研究表明，不同顶温类型大曲的微生物群落呈现出显著的特异性。Kang等^[35]通过扩增子测序发现，低温大曲以泛菌属(Pantoea)、葡萄球菌属(Staphylococcus)等细菌及Saccharomycopsis等真菌为主导；中温大曲以糖多孢菌属(Saccharopolyspora)、Staphylococcus、Bacillus等细菌及Saccharomycopsis、Aspergillus等真菌为主导；高温大曲则主要富集Saccharopolyspora、Bacillus、Staphylococcus、高温放线菌属(Thermoactinomyces)、克罗彭斯特德菌属(Kroppenstedtia)及Saccharomycopsis、丝衣霉菌属(Byssochlamys)、嗜热丝孢菌属(Thermomyces)、Aspergillus等耐/嗜热类群。张清玫等^[36]基于宏基因组技术结合线性判别分析进一步鉴定出不同类型大曲的特征菌群，低温大曲以Lactobacillus、明串珠菌属(Leuconostoc)、魏斯氏菌属(Weissella)等乳酸菌和Pichia为主；中温大曲以Bacillus、横梗霉菌属(Lichtheimia)和酵母菌属(Saccharomyces)为主；高温大曲则以串链孢菌属(Desmospora)、Saccharopolyspora、Byssochlamys和Aspergillus为主。值得注意的是，即便在相同类型大曲内部，温、湿度波动等因素介导的微环境异质性仍可驱动群落结构发生分化。Hou等^[37]通过宏基因组技术解析了不同发酵温度(通过曲块颜色深浅表征)的高温大曲微生物群落，发现真菌相对丰度随温度升高而显著增加；在此项研究中，谢瓦曲霉(Aspergillus chevalieri, 25.65%)和象牙色克罗彭斯特德菌(Kroppenstedtia eburnea, 21.07%)分别为真菌和细菌类群中的关键物种。

目前，大曲微生态研究主要聚焦于特定制曲条件下的微生物群落组成特征。除前文提到的温度影响外还涵盖了原辅料^[38]、压曲方式^[39]、强化方法^[40]等因素，以及在空间(如产区^[41]、曲房内位置^[42]、曲块内位置^[43])和时间(如季节^[44]、发酵阶段^[45]、发酵批次^[46])上的差异分布。此外，关于生物和非生物因素的驱动作用研究也在逐步深入，揭示了物种间及物种与环境因素之间的相互作用对微生物群落构建的影响^[6,14,33]。上述研究不仅深化了对大曲核心菌群组成、演替规律和发酵特性的理解，也为微生物功能研究提供了重要的生物学基础。通过指导大曲的菌株强化发酵和驱动因素调控，这些成果可为未来的合成群落设计^[47]和酿酒工艺优化提供重要参考。

2.2 大曲微生物功能研究基础

在大曲菌库资源中具有特定代谢功能的菌株因在酿酒工业中的应用潜力而备受关注，主要分为产香类(乙酸乙酯、丁酸乙酯、戊酸乙酯、己酸乙酯、辛酸乙酯、正丙醇、苄硫醇、三甲基吡嗪、四甲基吡嗪等)和产酶类(淀粉酶、纤维素酶、木聚糖酶、单宁酶、果胶酶、酯酶、蛋白酶、氧化还原酶、裂解酶、异构酶等)^[33,48-50]。传统的大曲微生物功能研究主要依赖于对大曲来源菌株产酶特性的分析，但这种方法不仅耗时耗力，且无法涵盖绝大多数微生物，也难以复现其在大曲环境中的原位表达模式。

在这种背景下，宏组学技术成为突破传统局限的新途径。基于中温大曲发酵过程的宏基因组分析表明，Weissella、Lactobacillus和Pediococcus等乳杆菌目(Lactobacillales)类群，Lichtheimia等毛霉菌目(Mucorales)类群，以及Aspergillus、Rasamsonia和Byssochlamys等散囊菌目(Eurotiales)类群在发酵过程中依次成为优势菌群，并参与底物裂解及风味前体/化合物生成等系列反应^[51]。另一项研究通过宏基因组技术对比低温、中温、高温大曲微生物群落功能，发现3类大曲虽均以碳水化合物和氨基酸代谢为核心生物学功能，但其功能谱和代谢通路存在显著分化；例如，低温大曲和中温大曲的EC 3.1.1.1 (羧酸酯酶)和EC 3.1.1.3 (三酰基甘油脂肪酶)基因呈现较高丰度特征，而EC 2.3.1.85 (脂肪酸合成酶)基因仅在高温大曲中检出，反映3类大曲在脂肪酸和酯类合成潜力上的差异^[52]。宏基因组分析主要反映微生物群落的潜在功能，宏转录组学分析则能从基因表达层面揭示群落中活跃的类群和代谢途径。Xu等^[53]采用宏转录组技术对比单曲(中温大曲)和多曲(中温与高温大曲)对兼香型白酒发酵的影响，研究表明在单曲发酵中Lactobacillus转录活性的增强促进了乳酸乙酯合成；在多曲发酵中醋乳杆菌属(Acetilactobacillus)、Saccharomyces和哈萨克斯坦酵母菌属(Kazachstania)的转录活性增强协同促进了乙酸酯类和中链脂肪酸乙酯的合成。

相较于基因组成解析和转录动态监测的间接预测特性，宏蛋白质组学则能够直接解析样本中蛋白质的功能，实现对不同条件下的蛋白质组的定性和定量比较^[54]。在酒曲研究中，Wang等^[55]通过非标记(label-free)定量宏蛋白质组技术对2种小曲酶系进行分析，共鉴定出1 421种真菌蛋白质和183种细菌蛋白质，发现饼曲相较于散曲主要上调了碳水化合物代谢相关蛋白质表达，涉及糖酵解/糖异生、磷酸戊糖途径、柠檬酸循环、丙酮酸代谢等关键途径。Yang等^[56]从3种颜色的高温大曲中共鉴定出1 030种蛋白质，系统揭示了粗糙脉孢霉(Neurospora crassa)等4种微生物通过调控五元环氨基酸代谢诱导大曲微生态分化的分子机制。柳习月等^[57]聚焦大曲中的苯丙氨酸代谢通路，发现Aspergillus、Bacillus、Byssochlamys、Kroppenstedtia、假单胞菌属(Pseudomonas)、Saccharopolyspora等微生物可通过合成EC 1.2.1.5/EC 1.2.1.39 (醛脱氢酶)、EC 3.5.1.4 (酰胺酶)、EC 2.6.1.1/EC 2.6.1.21 (转氨酶)、EC 4.1.1.28 (氨基酸脱羧酶)等功能酶协同催化苯丙氨酸转化为苯乙酸、苯乙醛等芳香族风味物质，为大曲提供花果香。在宏蛋白质组数据处理中蛋白质的鉴定通常依赖于搜库分析，而宏基因组数据是构建搜库参考数据库的重要来源。总的来说，构建基于微生物学、基因组学、蛋白质组学和生物信息学等多技术联用的分析路线，在解析和挖掘大曲体系中特定功能微生物和酶资源方面展现出巨大潜力，也为进一步研究大曲糖化功能提供了重要的理论基础和技术支持。

3 大曲糖化功能研究进展

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3.1 大曲糖化功能生化指标分析

在大曲质量评价体系中糖化力和液化力是评估其淀粉水解能力的重要指标，均以可溶性淀粉为检测底物。糖化力表示从淀粉分子释放还原糖的能力，常以3,5-二硝基水杨酸法或菲林试剂法检测^[58]；液化力表示将淀粉从高分子状态水解为较低分子状态(糊精等)的能力，常以碘液显色法检测^[5]。两者均代表大曲中多种酶的综合能力，有研究者将糖化力和液化力分别简单归因于葡萄糖淀粉酶和α-淀粉酶的作用，这种观点并不准确。作为内切酶，α-淀粉酶随机切割淀粉分子内部的α-1,4-糖苷键，主要生成的麦芽糖等短链寡糖同样具有还原性，能反映在糖化力指标上；作为外切酶，葡萄糖淀粉酶从淀粉分子末端依次切割α-1,4-糖苷键，也能使淀粉分子逐渐变短，消除与碘液的颜色反应，从而反映在液化力指标上。此外，β-淀粉酶、异淀粉酶、α-葡萄糖苷酶、麦芽糖淀粉酶等也可参与淀粉水解释放还原糖。总的来说，糖化力和液化力所指征的生化过程本质上密不可分，共同关联着大曲在酿酒过程中的淀粉糖化效率。

大曲的淀粉水解能力与发酵温度密切相关。研究表明不同顶温类型的成熟大曲在相关指标上呈现出显著差异，低温大曲和中温大曲的糖化力及液化力通常高于高温大曲^[35,39]。在不同颜色(颜色越深，顶温越高)的高温大曲中白曲、黄曲、黑曲的糖化力和液化力依次递减^[59]。这种差异可能与酶的热稳定性密切相关：在发酵温度较高、高温持续时间较长等因素影响下一些热不稳定的酶活性受损严重，从而削弱大曲性能。大曲的淀粉糖化性能指标同样也受到原辅料、贮存期、产地、季节等多重因素的影响。例如，有研究发现中温大曲成熟前后液化力无显著变化，而糖化力则显著降低^[60]。赵亮等^[61]通过模拟生产试验发现绿茶添加量为10%-30%的高温大曲在发酵结束时糖化力均显著大于未添加绿茶的大曲。

除了淀粉糖化功能评价外，部分研究也关注了广义糖化所涵盖的非淀粉多糖水解过程。Jiang等^[44]研究发现春季生产的大曲相较于夏季生产的大曲表现出更高的纤维素酶活力。刘建文等^[62]监测了特香型大曲中多种水解酶在1年贮存期内的活力变化，发现纤维素酶和木聚糖酶的活力均呈现先升高后下降的趋势，且在贮存3-4个月时达到最优状态。

3.2 大曲糖化功能组学研究进展

现代组学技术的兴起深化了对酿造体系微生物群落的研究，也为大曲糖化功能研究提供了新的视角。Huang等^[63]基于宏转录组分析在中温大曲升温和顶温阶段观察到糖酵解、淀粉代谢和丙酮酸代谢等途径中关键酶基因的表达上调，并从顶温阶段样本中鉴定出932种潜在表达的碳水化合物活性酶；在这项研究中功能微生物分类大多定位在科级或更高分类级别，主要包括发菌科(Trichocomaceae)、芽孢杆菌目(Bacillales)以及散囊菌亚纲(Eurotiomycetidae)等。

随着技术手段和数据库的持续完善，近期大曲研究已能更准确地判别关键功能物种。Zheng等^[64]研究表明，EC 3.2.1.1 (α-淀粉酶)和EC 3.2.1.3 (葡萄糖淀粉酶)是低温大曲中最主要的淀粉糖化功能酶，两者蛋白质丰度占比超过95%，主要由Lichtheimia贡献^[64]。Xia等^[65]基于宏蛋白质组学分析发现Kroppenstedtia、Aspergillus、根毛霉菌属(Rhizomucor)、Byssochlamys、Thermomyces是大曲中主要的淀粉酶贡献菌群，嗜热子囊菌属(Thermoascus)是主要的纤维素酶贡献菌群。Zhu等^[66]通过多组学分析研究了6轮成熟高温大曲中的糖化功能菌群，发现23种真菌和5种细菌能够分泌与主要碳水化合物代谢相关的13种糖苷水解酶；这些微生物主要包括扣囊复膜孢酵母(Saccharomycopsis fibuligera)、米曲霉(Aspergillus oryzae)、紫红曲霉(Monascus purpureus)和壮观丝衣霉(Byssochlamys spectabilis)等。Yang等^[67]归纳了中温和高温大曲中23种糖化功能相关酶，涉及淀粉、纤维素、半纤维素、几丁质和果胶等多糖水解酶系；宏基因组数据表明较高的发酵温度主要通过促进Bacillales同时抑制Lactobacillales和Mucorales，影响大曲的糖化功能酶基因丰度，而Eurotiales在2种顶温模式下无显著差异；宏蛋白质组学结果进一步证实了B. spectabilis、Monascus purpureus、埃默森罗萨氏菌(Rasamsonia emersonii)和疏棉状嗜热丝孢菌(Thermomyces lanuginosus)等Eurotiales类群在糖化功能中的优势贡献。

多糖的高效水解通常依赖不同酶之间的协同作用。内切型水解酶随机切割多糖的内部糖苷键，产生新的非还原链末端，为外切型水解酶提供了更多的作用底物。大曲中典型的协同作用组合有很多，例如，水解淀粉的α-淀粉酶、葡萄糖淀粉酶、EC 3.2.1.10 (寡聚-1,6-葡萄糖苷酶)和EC 3.2.1.20 (α-葡萄糖苷酶)；水解纤维素的EC 3.2.1.4 (内切-β-1,4-葡聚糖酶)、EC 3.2.1.91 (外切-β-1,4-葡聚糖酶)和EC 3.2.1.21 (β-葡萄糖苷酶)；水解几丁质的EC 3.2.1.14 (内切几丁质酶)和EC 3.2.1.52 (β-N-乙酰基氨基葡萄糖苷酶)；水解木聚糖的EC 3.2.1.8 (内切木聚糖酶)和EC 3.2.1.37 (β-木糖苷酶)等^[67]。酶的协同作用在一定程度上展现了大曲在糖化过程中的高效性。此外，大曲糖化功能酶的微生物来源也具有多样性。在大曲的微生物群落构建中某些类群尤其是丝状真菌的糖化功能基因呈现冗余特性，这意味着它们可能通过互补的方式维持大曲糖化功能的整体稳定性。综合上述多项研究，大曲作为糖化剂的优势得以进一步凸显。

3.3 大曲糖化功能微生物及功能酶挖掘

3.3.1 大曲糖化功能微生物筛选及应用

大曲糖化功能主要依赖于糖化功能菌群的富集和产酶活动。丝状真菌因其强大的多糖水解酶分泌能力被视为重要的糖化功能微生物，可培养类群主要包括Aspergillus、Rhizopus、Mucor、红曲霉菌属(Monascus)、犁头霉菌属(Absidia)和青霉属(Penicillium)等^[31,68-69]。其中，Absidia和Penicillium可能对发酵状态和最终产品的风味产生负面影响，因此在酿酒实践中少有应用^[68]。此外，Bacillus也被认为是大曲中重要的糖化功能细菌^[70]。这些菌株除了分泌淀粉酶、纤维素酶等糖化功能酶之外，还可以产生蛋白酶、脂肪酶等多种酶类，进一步促进原料的降解和风味的形成。

近年来，研究者从大曲中筛选出多种糖化功能微生物，并验证了它们在酿酒工业中的多重应用效果(表2)。此外，针对非淀粉多糖水解筛选出的功能菌株也表现出较强的应用潜力。例如，有研究针对性地以桔皮粉为唯一碳源，从中温大曲中筛选到高产果胶酶的鸡肠球菌(Enterococcus gallinarum)和橙色嗜热子囊菌(Thermoascus aurantiacus)菌株，其中E. gallinarum所产果胶酶在30-50 ℃具有良好的热稳定性^[77]。Zhang等^[78]从大曲中筛选出的微生物群落FA12能稳定分泌纤维素酶和木聚糖酶，其酶活力分别位于0.235 0-0.447 0 U/mL和0.191 7-0.307 8 U/mL区间，还能够发酵酒糟释放阿魏酸。Liu等^[79]将高产纤维素酶的枯草芽孢杆菌(Bacillus subtilis)用于大曲强化，显著提升了大曲糖化力、液化力及优势活性菌群相对丰度(85.08%，对照组为63.42%)，同时上调了淀粉和蔗糖代谢相关酶的表达。

值得注意的是，从生态相互作用的角度来看，功能菌株不仅拥有特定的功能，还具备多样的代谢特性，这导致其在实际应用中的效果存在着一些不确定性。功能菌株可能对产量和风味产生积极影响，但也可能与其他微生物发生竞争或拮抗，从而影响生产系统的效能和稳定性^[80-81]。

3.3.2 大曲糖化功能酶鉴定及表达

以前挖掘环境样品中的未知酶基因资源常采用黏粒(cosmid)、F黏粒(fosmid)文库筛选法。若将基因组学与生物信息学技术相结合可进一步提升功能酶的筛选效率^[82]。Huang等^[63]基于对大曲发酵过程宏转录组数据的分析和挖掘，从顶温阶段(62 ℃)样本中筛选、克隆了多种新型糖化酶基因，并通过异源表达系统表征了它们的酶学性质以及底物谱，拓展了大曲糖化功能的研究途径。所涉及的酶种包括α-淀粉酶^[83]、α-葡萄糖苷酶^[84]及EC 3.2.1.73 (β-1,3-1,4-葡聚糖酶)^[85]等。此外，另有研究基于白酒发酵过程酒醅的多组学数据从大曲DNA中克隆了8种淀粉糖化功能酶基因，其中成功表达的细菌来源葡萄糖淀粉酶KeGA5和真菌来源α-淀粉酶RpAM11的最适反应温度分别为55 ℃和75 ℃，两者在多种含有α-1,4-葡萄糖苷键的底物体系和高粱孵育体系中均展现出显著的协同作用^[86]。鉴于大曲蕴含着丰富的酶学资源，相信未来将有更多高效的糖化功能酶被鉴定、表达及应用。

3.4 大曲糖化功能在白酒酿造过程中的作用

经过蒸煮处理后高粱中的淀粉实现了部分糊化。向摊晾后的高粱原料中添加曲粉便启动了酿酒工艺中的糖化发酵阶段^[87]。大曲中的糖化酶在原料糖化及后续的乙醇与风味物质代谢中扮演着关键角色，这一点已得到业界的广泛认可。近期，一项基于宏蛋白质组技术的研究表明，在白酒发酵的初始阶段低温大曲贡献了90%的α-淀粉酶和99%的葡萄糖淀粉酶，这些酶在糖化过程中发挥了关键作用^[64]。Wang等^[88]从小曲白酒的发酵酒醅中鉴定出51种碳水化合物水解酶，其中约80%来源于小曲，并发现α-淀粉酶和葡萄糖淀粉酶在乙醇生成过程中具有协同效应。

生产经验表明，固态酿造白酒酒醅中淀粉浓度的下降速度和幅度受到曲的质量、发酵温度和升酸状况等因素的制约^[68]。实验发现，添加外源的淀粉水解相关糖化酶可有效提升出酒率并减少高级醇的生成^[89-90]。然而，现有的商业糖化酶制剂仍无法取代大曲，主要原因在于大曲不仅含有多种糖化酶，还富含其他酶类和微生物，能够实现更复杂的糖化及发酵过程。此外，糖化酶的直接添加容易引起底物快速分解和微生物过度发酵，从而导致温度的迅速上升，对发酵体系和白酒产品品质产生不利影响^[91]。

非淀粉多糖的水解在白酒酿造过程中的作用常常被忽视，但同样值得探究。例如，几丁质不仅是大曲丝状真菌和白酒酿造过程中酵母等真菌细胞壁的主要成分，还是成熟大曲贮存期间曲虫外壳的主要结构。几丁质经酶促降解后的主要产物为氨基葡萄糖，其氨基可与羰基化合物(如醛、酮类)发生席夫碱反应，形成具有风味的含氮化合物^[92]，并参与美拉德反应或其他热处理过程中的复杂反应生成棕色化合物，可能对发酵产品的风味和色泽产生显著贡献。未来在淀粉多糖底物的研究基础上有必要针对非淀粉多糖底物在酿造过程中的降解及其对风味的贡献开展更深入的研究。

4 总结与展望

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大曲作为优质白酒生产中最常用的糖化发酵剂，凭借开放的发酵模式、相对稳定的发酵效果以及丰富的微生物和功能酶系统成为亚洲传统多菌混合糖化剂研究的理想模型。本文通过对传统培养方法与现代组学技术相关文献的综述，系统总结了大曲在糖化功能微生物富集、糖化功能酶系组成及糖化机制等方面的研究进展及优势，为传统工艺的科学阐释提供了关键证据。

从当前研究现状来看，尽管多种现代生物技术的应用已揭示了部分信息，但仍存在亟待突破的瓶颈。(1) 在构建“大曲制备-白酒发酵”全链条关联模型时，准确量化大曲成品糖化性能对白酒发酵菌群演替及风味代谢的影响仍需多组学技术串联的深入研究和大量的实验验证。(2) 现有研究多聚焦于外源淀粉水解酶的直接应用，而对大曲来源淀粉水解酶及非淀粉多糖水解酶缺少系统挖掘和应用探究。(3) 以功能微生物为核心构建人工群落时需解决菌株竞争抑制、代谢通路兼容性等问题，同时也可尝试探索其与糖化功能酶制剂的协同增效策略。(4) 在现有的糖化力、液化力指标基础上需建立更加科学合理的大曲糖化功能评价指标体系，并结合机器学习算法实现对大曲糖化功能的智能预测和工艺参数调控。深入解析以上科学问题可提升生产效率和产品质量的稳定性，并推动传统酿酒工艺与现代科技的深度融合。

基金

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泸州市科技计划(2024JYJ102)

参考文献

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文献

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doi: 10.13343/j.cnki.wsxb.20250241

接收时间：2025-03-25
首发时间：2025-11-03
出版时间：2025-09-04

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泸州品创科技有限公司，国家固态酿造工程技术研究中心，四川泸州

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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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Name	Main origin	Main ingredients	Main fermented products	Main microorganisms
Daqu^[4,6,14]	China	Wheat, barley, pea	Daqu Baijiu, vinegar	Aspergillus, Mucor, Rhizopus, Lichtheimia, Thermomyces, Rhizomucor, Monascus, Pichia, Saccharomycopsis, Bacillus, Lactobacillus, Weissella, Thermoactinomyces, Kroppenstedtia, Saccharopolyspora
Xiaoqu^[6,14]	China	Rice, soybean	Xiaoqu Baijiu, Huangjiu	Aspergillus, Rhizopus, Saccharomycopsis, Pichia, Candida, Lactobacillus, Pediococcus, Weissella
Fuqu^[6,14]	China	Wheat bran	Fuqu Baijiu	Aspergillus, Rhizopus, Saccharomyces, Bacillus
Koji^[15]	Japan	Rice, barley	Saké (fermented wine), Miso (fermented soybean paste), Shoyu (soy sauce)	Aspergillus, Wickerhamomyces, Candida, Ochrobactrum, Lactobacillus
Nuruk^[16]	Korea	Wheat, rice	Makgeolli (fermented wine), Yakju (fermented wine), Soju (distilled liquor)	Aspergillus, Rhizomucor, Lichtheimia, Mucor, Saccharomycopsis, Pichia, Debaryomyces
Loog-pang^[17]	Thailand	Rice, herb, spice	Khao-maak (alcoholic food), Sato (fermented wine)	Amylomyces, Rhizopus, Aspergillus, Mucor, Absidia, Saccharomycopsis, Pichia, Saccharomyces, Pediococcus
Bánh men^[18]	Vietnam	Rice, herb, spice	Ruou nep than (fermented wine), Ruou de (distilled liquor)	Rhizopus, Absidia, Amylomyces, Saccharomycopsis, Saccharomyces, Issatchenkia, Pediococcus, Lactobacillus, Weissella, Bacillus
Bubod^[17]	Philippines	Rice, herb	Basi (fermented wine)	Mucor, Rhizopus, Saccharomyces, Saccharomycopsis
Hamei^[19-20]	India	Rice, herb	Atingba (fermented wine), Yu (distilled liquor)	Mucor, Rhizopus, Penicillium, Saccharomyces, Pichia, Trichosporon
Xaj-pitha^[21]	India	Rice, herb	Rohi (fermented wine), Xaj (fermented wine)	Rhizopus, Mucor, Meyerozyma, Wickerhamomyces, Saccharomyces, Candida, Lactobacillus, Leuconostoc, Weissella
Marcha^[17,19]	India, Bhutan, Nepal, Xizang of China	Rice, herb, spice	Kodo ko jaanr (alcoholic beverage), Bhaati jaanr (alcoholic beverage), Raksi (distilled liquor)	Aspergillus, Mucor, Rhizopus, Penicillium, Bjerkandera, Saccharomycopsis, Pichia, Pediococcus
Phab^[17]	India, Bhutan, Nepal, Xizang of China	Wheat, herb	Chhang (alcoholic beverage)	Penicillium, Aspergillus, Saccharomycopsis
Ragi^[17]	Indonesia	Rice, millet, cassava, herb, spice	Tapé (alcoholic food)	Rhizopus, Mucor, Amylomyces, Aspergillus, Saccharomycopsis, Candida, Saccharomyces, Pichia, Pediococcus

Name

Main origin

Main ingredients

Main fermented products

Main microorganisms

Daqu^[4,6,14]

China

Wheat, barley, pea

Daqu Baijiu, vinegar

Aspergillus, Mucor, Rhizopus, Lichtheimia, Thermomyces, Rhizomucor, Monascus, Pichia, Saccharomycopsis, Bacillus, Lactobacillus, Weissella, Thermoactinomyces, Kroppenstedtia, Saccharopolyspora

Xiaoqu^[6,14]

China

Rice, soybean

Xiaoqu Baijiu, Huangjiu

Aspergillus, Rhizopus, Saccharomycopsis, Pichia, Candida, Lactobacillus, Pediococcus, Weissella

Fuqu^[6,14]

China

Wheat bran

Fuqu Baijiu

Aspergillus, Rhizopus, Saccharomyces, Bacillus

Koji^[15]

Japan

Rice, barley

Saké (fermented wine), Miso (fermented soybean paste), Shoyu (soy sauce)

Aspergillus, Wickerhamomyces, Candida, Ochrobactrum, Lactobacillus

Nuruk^[16]

Korea

Wheat, rice

Makgeolli (fermented wine), Yakju (fermented wine), Soju (distilled liquor)

Aspergillus, Rhizomucor, Lichtheimia, Mucor, Saccharomycopsis, Pichia, Debaryomyces

Loog-pang^[17]

Thailand

Rice, herb, spice

Khao-maak (alcoholic food), Sato (fermented wine)

Amylomyces, Rhizopus, Aspergillus, Mucor, Absidia, Saccharomycopsis, Pichia, Saccharomyces, Pediococcus

Bánh men^[18]

Vietnam

Rice, herb, spice

Ruou nep than (fermented wine), Ruou de (distilled liquor)

Rhizopus, Absidia, Amylomyces, Saccharomycopsis, Saccharomyces, Issatchenkia, Pediococcus, Lactobacillus, Weissella, Bacillus

Bubod^[17]

Philippines

Rice, herb

Basi (fermented wine)

Mucor, Rhizopus, Saccharomyces, Saccharomycopsis

Hamei^[19-20]

India

Rice, herb

Atingba (fermented wine), Yu (distilled liquor)

Mucor, Rhizopus, Penicillium, Saccharomyces, Pichia, Trichosporon

Xaj-pitha^[21]

India

Rice, herb

Rohi (fermented wine), Xaj (fermented wine)

Rhizopus, Mucor, Meyerozyma, Wickerhamomyces, Saccharomyces, Candida, Lactobacillus, Leuconostoc, Weissella

Marcha^[17,19]

India, Bhutan, Nepal, Xizang of China

Rice, herb, spice

Kodo ko jaanr (alcoholic beverage), Bhaati jaanr (alcoholic beverage), Raksi (distilled liquor)

Aspergillus, Mucor, Rhizopus, Penicillium, Bjerkandera, Saccharomycopsis, Pichia, Pediococcus

Phab^[17]

India, Bhutan, Nepal, Xizang of China

Wheat, herb

Chhang (alcoholic beverage)

Penicillium, Aspergillus, Saccharomycopsis

Ragi^[17]

Indonesia

Rice, millet, cassava, herb, spice

Tapé (alcoholic food)

Rhizopus, Mucor, Amylomyces, Aspergillus, Saccharomycopsis, Candida, Saccharomyces, Pichia, Pediococcus

菌种 Strains	来源 Sources	应用 Applications	主要结果与影响(相对于对照组) Main results and effects (compared with the control group)
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲 Fortifying Daqu^[71]	(1) Bacillus、Clavispora和Aspergillus丰度增加，Pichia、Saccharomycopsis丰度降低 (1) The abundances of Bacillus, Clavispora, and Aspergillus increased, while those of Pichia and Saccharomycopsis decreased (2) 液化力、糖化力显著增加，酯化力显著降低 (2) The liquefying power and saccharifying power increased significantly, while the esterifying power decreased significantly (3) 吡嗪类和芳香族类化合物含量增加 (3) The contents of pyrazines and aromatic compounds increased
Bacillus licheniformis	高温大曲 High-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[72]	(1) 前期Lactobacillus基因转录量降低，后期Kazachstania、Naumovozyma、Saccharomyces基因转录量增加 (1) The gene transcription level of Lactobacillus decreased in the early stage, while those of Kazachstania, Naumovozyma, and Saccharomyces increased in the later stage (2) 后期Saccharomyces乙醇脱氢酶、乙醛脱氢酶和乳酸脱氢酶基因转录量增加 (2) The gene transcription levels of alcohol dehydrogenase, aldehyde dehydrogenase, and lactate dehydrogenase of Saccharomyces increased in the later stage
Bacillus subtilis Bacillus velezensis	大曲 Daqu	强化大曲 Fortifying Daqu^[73]	(1) Bacillus、Lactobacillus和Candida丰度增加 (1) The abundances of Bacillus, Lactobacillus, and Candida increased (2) 液化力、糖化力、酯化力显著增加 (2) The liquefying power, saccharifying power, and esterifying power increased significantly (3) 酯类、吡嗪类和醇类化合物含量显著增加 (3) The contents of esters, pyrazines, and alcohols increased significantly
Monascus purpureus Aspergillus oryzae Rhizopus arrhizus	中温大曲 Medium-temperature Daqu	强化大曲后酿酒 Brewing after fortifying Daqu^[74]	(1) 酒精度提高8% (1) The alcohol content increased by 8% (2) 风味物质数量和含量均增加 (2) Both the quantity and content of flavor substances increased
Rhizopus oryzae Bacillus velezensis	中温大曲 Medium-temperature Daqu	强化麸曲 Fortifying Fuqu^[75]	(1) 通过平板对峙实验，确认两者无拮抗作用 (1) Through the plate confrontation experiment, it was confirmed that there was no antagonistic effect between the two strains (2) 通过单因素和响应面实验，确定复合麸曲的最佳制备工艺 (2) Through single-factor and response surface experiments, the optimal preparation process of the composite Fuqu was determined
Bacillus amyloliquefaciens Saccharomycopsis fibuligera Absidia corymbifera	低温大曲 Low-temperature Daqu	强化大曲 Fortifying Daqu^[76]	(1) 淀粉酶活力提高 (1) The amylase activity increased (2) 微生物群落结构差异较小，丰富度提高 (2) There was little difference in the microbial community structure, and the richness increased (3) 真菌强化组中的乙醇、异丁醇、异戊醇含量增加 (3) The contents of ethanol, isobutanol, and isoamyl alcohol in the fungi-fortified group increased

菌种

Strains

来源

Sources

应用

Applications

主要结果与影响(相对于对照组)

Main results and effects (compared with the control group)

Bacillus licheniformis

高温大曲

High-temperature Daqu

强化大曲

Fortifying Daqu^[71]

(1) Bacillus、Clavispora和Aspergillus丰度增加，Pichia、Saccharomycopsis丰度降低

(1) The abundances of Bacillus, Clavispora, and Aspergillus increased, while those of Pichia and Saccharomycopsis decreased

(2) 液化力、糖化力显著增加，酯化力显著降低

(2) The liquefying power and saccharifying power increased significantly, while the esterifying power decreased significantly

(3) 吡嗪类和芳香族类化合物含量增加

(3) The contents of pyrazines and aromatic compounds increased

Bacillus licheniformis

高温大曲

High-temperature Daqu

强化大曲后酿酒

Brewing after fortifying Daqu^[72]

(1) 前期Lactobacillus基因转录量降低，后期Kazachstania、Naumovozyma、Saccharomyces基因转录量增加

(1) The gene transcription level of Lactobacillus decreased in the early stage, while those of Kazachstania, Naumovozyma, and Saccharomyces increased in the later stage

(2) 后期Saccharomyces乙醇脱氢酶、乙醛脱氢酶和乳酸脱氢酶基因转录量增加

(2) The gene transcription levels of alcohol dehydrogenase, aldehyde dehydrogenase, and lactate dehydrogenase of Saccharomyces increased in the later stage

Bacillus subtilis

Bacillus velezensis

大曲

Daqu

强化大曲

Fortifying Daqu^[73]

(1) Bacillus、Lactobacillus和Candida丰度增加

(1) The abundances of Bacillus, Lactobacillus, and Candida increased

(2) 液化力、糖化力、酯化力显著增加

(2) The liquefying power, saccharifying power, and esterifying power increased significantly

(3) 酯类、吡嗪类和醇类化合物含量显著增加

(3) The contents of esters, pyrazines, and alcohols increased significantly

Monascus purpureus

Aspergillus oryzae

Rhizopus arrhizus

中温大曲

Medium-temperature Daqu

强化大曲后酿酒

Brewing after fortifying Daqu^[74]

(1) 酒精度提高8%

(1) The alcohol content increased by 8%

(2) 风味物质数量和含量均增加

(2) Both the quantity and content of flavor substances increased

Rhizopus oryzae

Bacillus velezensis

中温大曲

Medium-temperature Daqu

强化麸曲

Fortifying Fuqu^[75]

(1) 通过平板对峙实验，确认两者无拮抗作用

(1) Through the plate confrontation experiment, it was confirmed that there was no antagonistic effect between the two strains

(2) 通过单因素和响应面实验，确定复合麸曲的最佳制备工艺

(2) Through single-factor and response surface experiments, the optimal preparation process of the composite Fuqu was determined

Bacillus amyloliquefaciens

Saccharomycopsis fibuligera

Absidia corymbifera

低温大曲

Low-temperature Daqu

强化大曲

Fortifying Daqu^[76]

(1) 淀粉酶活力提高

(1) The amylase activity increased

(2) 微生物群落结构差异较小，丰富度提高

(2) There was little difference in the microbial community structure, and the richness increased

(3) 真菌强化组中的乙醇、异丁醇、异戊醇含量增加

(3) The contents of ethanol, isobutanol, and isoamyl alcohol in the fungi-fortified group increased