微生物学报

酿酒酵母高级醇代谢的转录调控研究进展

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陈璐 ¹ , 刘延琳 ¹ , 秦义 ¹^,²^,³^,^*

微生物学报 | 综述 2024,64(12): 4669-4680

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微生物学报 | 综述 2024, 64(12): 4669-4680

酿酒酵母高级醇代谢的转录调控研究进展

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陈璐¹, 刘延琳¹, 秦义¹^,²^,³^,^*

作者信息

1 西北农林科技大学葡萄酒学院, 陕西杨凌 712100

2 国家林业和草原局葡萄与葡萄酒工程技术研究中心, 陕西杨凌 712100

3 西北农林科技大学合阳葡萄试验示范站, 陕西渭南 714000

Transcriptional regulation of higher alcohol metabolism in Saccharomyces cerevisiae

Lu CHEN¹, Yanlin LIU¹, Yi QIN¹^,²^,³^,^*

Affiliations

1 College of Enology, Northwest A&F University, Yangling 712100, Shaanxi, China

2 National Forestry and Grassland Administration Engineering Research Center for Viti-viniculture, Yangling 712100, Shaanxi, China

3 Heyang Viti-viniculture Station, Northwest A&F University, Weinan 714000, Shaanxi, China

出版时间: 2024-09-26 doi: 10.13343/j.cnki.wsxb.20240305

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摘要

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高级醇是酿酒酵母在葡萄酒酿造过程中产生的主要代谢副产物，其代谢受到多层次调控体系的精细调节。酿酒酵母高级醇代谢途径中的酶系及其编码基因已基本明确，但酿酒酵母高级醇代谢的转录调控机制仍不清晰。本文在总结酵母高级醇代谢途径及其代谢调控策略的基础上，重点综述了参与酵母高级醇代谢调控的转录因子Aro80p、GATA和Leu3p及其作用机制。旨在为系统理解酵母高级醇代谢转录调控机制，以及选育高级醇产量适中的酵母菌种提供理论参考。

关键词

酿酒酵母 / 高级醇 / 转录因子 / 转录调控

Abstract

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Higher alcohols, major metabolic byproducts produced by Saccharomyces cerevisiae during winemaking, are intricately regulated by a multilevel system. While the enzymatic machinery and their encoding genes involved in the metabolic pathways of higher alcohols in S. cerevisiae have been largely elucidated, the transcriptional regulation underlying this process remains poorly understood. This paper, building upon a summary of the metabolic pathways and regulatory strategies of higher alcohols metabolism in yeast, focuses on the transcription factors Aro80p, GATA and Leu3p implicated in the regulation of higher alcohols metabolism in yeast and their mechanisms of action. The review aims to give theoretical insights into a comprehensive understanding of the transcriptional regulation of higher alcohols metabolism in yeast and facilitate the breeding of yeast strains with moderate production of higher alcohols.

Key words

Saccharomyces cerevisiae / higher alcohols / transcription factors / transcriptional regulation

引用本文

陈璐, 刘延琳, 秦义. 酿酒酵母高级醇代谢的转录调控研究进展. 微生物学报, 2024 , 64 (12) : 4669 -4680 . DOI: 10.13343/j.cnki.wsxb.20240305

Lu CHEN, Yanlin LIU, Yi QIN. Transcriptional regulation of higher alcohol metabolism in Saccharomyces cerevisiae[J]. Acta Microbiologica Sinica, 2024 , 64 (12) : 4669 -4680 . DOI: 10.13343/j.cnki.wsxb.20240305

正文

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全球气候变暖给葡萄酒产业带来了深刻影响，其中最为突出的是葡萄采收时的含糖量在过去20年里逐年攀升^[1]。近年我国西北葡萄酒产区主栽红色酿酒葡萄的含糖量普遍超过260 g/L。葡萄的高含糖量不仅使发酵困难、酒度升高，还会导致葡萄酒高级醇含量显著升高。高级醇也称为“杂醇油”，指含有3个及以上碳原子的一元醇，是酿酒酵母的主要代谢副产物^[2-5]。研究表明，葡萄酒中的高级醇含量小于300 mg/L时，能够赋予葡萄酒令人愉悦的风味，当超过400 mg/L则会对葡萄酒带来辛辣、腐臭等不愉悦的杂味，从而导致葡萄酒香气不纯净、感官品质劣变等问题^[6]。更重要的是，高级醇含量过高易引发头痛、口渴、醒酒慢等不良症状，对饮用者构成安全隐患^[7]。随着葡萄原料含糖量逐年增加，这一问题表现得越来越突出，已成为阻碍我国西北产区葡萄酒质量飞跃的重要问题。因此，亟须在葡萄酒酿造过程中合理控制高级醇含量。

葡萄酒中高级醇的产生与多种发酵环境和工艺条件相关，如发酵基质的同化氮含量和类型^[7]、糖含量^[8]和添加方法、发酵温度、发酵基质的初始pH值及溶氧量等^[9-10]。面对复杂的发酵环境条件和多样的工艺措施，酿酒酵母可通过基因转录、转录后翻译和翻译后加工等多层次的调控体系，精细调节高级醇的合成代谢，其中转录水平的调控发生在基因表达的初期阶段，通常是代谢调控的主要方式之一。目前酿酒酵母的高级醇代谢途径已梳理清楚，代谢途径中的酶系及其编码基因已基本明确^[11]，针对酿酒酵母高级醇代谢的转录调控研究较少，在酿酒酵母高级醇合成代谢中发挥关键作用的转录因子及其转录调控机制尚不全面，这也导致对酿酒酵母的定向代谢工程育种存在代谢调控效果不理想等诸多问题。本文综述了酿酒酵母高级醇代谢途径和参与高级醇代谢的相关转录因子，旨在系统了解酿酒酵母高级醇代谢及转录调控机制，为改良和选育高级醇产量适中的优良酿酒酵母菌种提供理论支持。

1 高级醇代谢途径

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葡萄酒中高级醇的主体组分为正丙醇、异戊醇、异丁醇、苯乙醇和活性戊醇等，约占高级醇总量的70%。酿酒酵母的高级醇主要经由糖合成代谢途径(Harris途径)和氨基酸分解代谢途径(Ehrlich途径)合成(图1)^[12]。Harris途径主要指葡萄糖/果糖的糖酵解终产物α-酮酸，经2-酮基酸脱羧酶(KDCs)脱羧、脱氢酶(ADHs)脱氢还原成活性戊醇、异丁醇、异戊醇等高级醇；Ehrlich途径是氨基酸在氨基转移酶(BATs)的催化作用下生成α-酮酸，α-酮酸在脱羧酶的作用下生成相应的醛和CO₂，醛经还原生成相应的醇。酿酒酵母所产生的高级醇约75%来自Harris途径，约25%来自Ehrlich途径^[7]。

酵母的高级醇合成代谢是多基因控制的复杂性生物学过程，涉及多个调控因子。现有研究采用了过量表达/敲除高级醇类物质的代谢途径基因^[13]，删除/增强竞争途径^[14]，改变辅因子水平^[15]，或在线粒体或细胞质重构代谢途径^[16]等策略，调控了酵母的高级醇代谢模式。其中，研究人员通过删除非特异性氨基酸转运蛋白编码基因GAP1、AGP1^[10]以及支链氨基酸分解途径的BAT2^[17]，敲除α-酮酸合成代谢途径的LEU1、LEU2、GDH1^[18]，删除α-酮酸分解代谢途径的HOM2、PAD1、QCR2、SPE1、ALD6^[19]，过量表达乙酸酯代谢基因ATF1或同时敲除IAH1基因^[20]等，均在一定程度上降低了酵母的高级醇产量。

对碳、氮等代谢通路上的单(多)基因敲除或过量表达，在削弱酿酒酵母高级醇合成途径代谢能力的同时，可能会导致乙酸等代谢副产物的增加，这些代谢副产物将严重影响微生物细胞生长和损伤葡萄酒感官品质，以及导致工程菌株发酵特性的改变。因此，通过对高级醇合成途径转录调控的精细优化，可能实现在基因组层面上定向进化或同时改变多个相关基因群，实现精准弱化高级醇合成能力的目的。

2 高级醇转录调控

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2.1 转录因子结构

转录因子(transcription factor, TF)通过识别并结合到DNA上的特定序列(顺式作用元件、增强子或沉默子)，激活或抑制相关基因的转录，进而影响高级醇的合成。根据与DNA结合结构域类型^[21]，TF可分为3类，包括锌簇(Zn²⁺)稳定型、螺旋-旋转-螺旋型和拉链型。其中，Zn²⁺稳定型的DNA结合结构域在各种生物体中普遍存在。它的功能域模型如图2A所示，整个DNA结合结构域(DNA-binding domain, DBD)分为3个区域：锌指区、连接体区和二聚区。锌簇蛋白有一个DBD，其中包含2个Zn²⁺，并以6个半胱氨酸(Cys)残基作为配体^[22]。在多数锌簇TF的结构中存在一个显著特征，即DBD，位于TF的氨基端(N端)。这一结构域负责识别并结合特定的DNA序列，从而调控基因的表达。与此同时，与DNA序列直接相互作用并参与特异性结合的残基，则位于TF的羧基端(C端)^[23] (图2B)。这种结构使TF能够以高效且特异的方式与DNA相互作用，进而调控转录过程。

2.2 与高级醇代谢相关的转录因子

目前已知的与高级醇代谢相关的转录因子有Aro80p^[24]、GATA^[25]和Leu3p^[26-29]。

2.2.1 Aro80p

Aro80p属于Zn₂Cys₆蛋白家族，由ARO80基因编码，其能够对芳香族氨基酸作出响应，并促进ARO9和ARO10基因的表达^[30]。ARO8和ARO9编码的芳香族转氨酶I和II催化氨基酸形成相应的α-酮酸类似物，在ARO10编码的芳香族脱羧酶的作用下，进一步转化为2-苯乙醇、色醇和酪醇^[31] (图1)。

Lee等^[32]研究发现，ARO9和ARO10基因的表达会被Aro80p激活，从而促进2-苯乙醇的合成；ARO80基因的缺失显著降低2-苯乙醇的合成。然而同时也发现，利用AgTEF1启动子过表达ARO80，会导致支链氨基酸分解代谢衍生的异戊醇和异丁醇水平增加2.5倍，而2-苯乙醇的水平基本不变^[33]。

Aro80p的N末端具有1个DBD，能够与靶基因启动子序列上一段特定的36个碱基对(bp)长度的激活序列相结合。这一激活序列最初在ARO9和ARO10基因的启动子序列中被发现，它由4组CCG序列组成，每组CCG之间相隔7 bp (图3)；此外，在ARO80基因的启动子序列中也证实了这一激活序列的存在^[30]。若要Aro80p与ARO9和ARO10启动子上的特定重复序列CCG相结合，还需芳香族氨基酸协助触发诱导信号。当培养基中氮源充足时，通过氮代谢物的阻遏抑制氨基酸通透酶编码基因GAP1表达，导致质膜上Gap1p失活。然而，当氮源只有l-苯丙氨酸存在时，Gap1p的转运活性将恢复，使得芳香族氨基酸在Gap1p的协助下进入细胞并参与代谢过程^[34-35]，进而促使Aro80p与靶基因启动子的特定重复序列相结合发挥作用(图4)。这种特定的结合触发了ARO9与ARO10基因的表达活性，从而实现对2-苯乙醇的有效调控。因此，可以看出，Aro80p不仅直接参与了ARO9、ARO10和ARO80的调控，也间接影响了氨基酸转运酶。

2.2.2 GATA转录因子

GATA家族作为一类具有IV型锌指结构的转录调控因子，具有特异性识别并结合GATA基序(motif)的能力^[36]。在酿酒酵母中，氮代谢相关基因的表达由4个GATA家族转录因子共同调控，它们是Gat1p、Gln3p、Gzf3p和Dal80p^[37-38]。这4个转录因子具有一个共同的特征性锌指部分，使其能够结合到上游激活序列上，该区域位于受氮调控的基因上游约数百个碱基对处，并激活(Gat1p和Gln3p)或抑制(Gzf3p和Dal80p)转录的启动。

当敲除酿酒酵母GAT1和GLN3基因后，与高级醇代谢相关的基因表达水平显著降低^[39]。Wang等^[17]发现，GAT1缺失菌株中高级醇总含量增加28.36%，达到615.73 mg/L。GLN3的缺失可增加酵母对异丁醇的耐受性，异丁醇产量提高了4.9倍^[40]。此外，尽管目前暂未发现Dal81p是否参与了酵母的高级醇代谢调控，但有研究显示Dal80p可能通过调节尿素代谢参与了氨基甲酸乙酯的调控^[41]。Dal80p的同系物Dal81p与前者的作用明显不同，Dal81p是从尿囊素、尿素和亮氨酸获取氮的正调节因子^[42]。

GATA家族Gat1p和Gln3p作为转录激活因子调控酿酒酵母中的氮分解代谢物抑制(nitrogen catabolism repression, NCR)。当微生物处于含有多种氮源的环境中时，它们会优先选择利用易于同化且能量效率高的氮源(如氨、谷氨酸等)，而抑制利用成本较高或效率较低的氮源相关基因的表达^[43]。GLN3基因的分离和测序结果表明，在306−330区域，Gln3p有1个锌指结构，与高等生物的GATA转录因子具有显著同源性^[30]。免疫沉淀实验显示，Gln3p与GLN1基因启动子上游5′-GATAA/G-3′序列相结合^[44]。在酵母中，除了已知的Gln3p调控因子外，还发现了氮代谢途径中的另一个重要激活因子Gat1p。该因子与Gln3p都具有GATA锌指基序，对于NCR敏感基因的表达至关重要，并呈现出轻微的转录激活潜力^[45]。Gat1p能够与目标基因的启动子中位于310−334区域的特定序列“5′-GATAA/G-3′”进行结合^[44]。在氮源充足的情况下，Tor1p及其他未知的磷酸激酶被激活，致使Gat1p和Gln3p发生磷酸化，并被核膜排斥至细胞质中，阻碍其参与靶基因的转录调控^[46]。当Gat1p和Gln3p处于非活化状态时，多个基因的转录受到抑制，包括DUR1、DUR2 (编码脲基酰胺酶)、DUR3 (编码尿素渗透酶)、CAR1 (编码精氨酸酶)；而在氮源匮乏时，Gat1p和Gln3p进入细胞核，并与NCR敏感基因启动子上的GATA序列结合，进而激活转录^[47] (图5)。

2.2.3 Leu3p

Leu3p作为转录调控因子，可能参与了包括异戊醇和异丁醇等高级醇类物质的代谢调控^[48-49]。Leu3p是一个Zn(II)₂Cys₆锌簇蛋白转录因子，结合DNA翻转重复序列CCG-N4-CGG，LEU3基因的表达受Gcn4介导的通用氨基酸控制(general amino acid control, GAAC)；Leu3p自身的激活程度取决于细胞内Leu3p浓度，也受酵母细胞内亮氨酸/异戊醇生物合成途径的中间代谢物α-异丙基苹果酸(α-isopropylmalate, α-IPM)的调控；α-IPM是Leu3p的共激活因子，当α-IPM充足时，形成Leu3-α-IPM复合物，发挥激活因子功能，反之Leu3p则为转录阻遏物^[50-51] (图6)。

转录因子Leu3p对异戊醇合成途径编码基因BAT1和LEU4的调控机制相对清晰。BAT1编码线粒体支链氨基酸转氨酶，催化Ehrlich途径第1步反应(转氨作用)；Bat1p主要参与支链氨基酸的生物合成^[51]，在对数生长期内表现为快速表达并大量积累，进入稳定期后其表达受到抑制。在生物合成阶段BAT1的表达主要受到Leu3-α-IPM的激活调控，其中，Leu3p结合于无核小体区域(nucleosome-free regions, NFR)^[26]的BAT1侧翼区。LEU4编码的α-异丙基苹果酸合酶(Leu4p)是亮氨酸/异戊醇生物合成途径的关键调控酶，受亮氨酸反馈抑制^[52]。LEU4启动子的主要调控元件是Leu3p结合元件(upstream active sequence, UAS)和2个一般控制响应Gcn4p结合元件(genetic code expansions, GCEs)^[53]。亮氨酸对LEU4的调节是通过α-IPM进行的。当细胞内亮氨酸供应不足时，其对α-异丙基苹果酸合酶的反馈抑制作用减弱，引起α-IPM水平升高并与Leu3p相互作用，从而激活LEU4表达^[52]。

尽管转录因子Leu3p对BAT1和LEU4的作用机制相对清晰，但通过YEASTRACT数据库^[54]分析发现，Leu3p与异戊醇和异丁醇代谢途径的16个基因也有直接或间接的关系。其中，Leu3p激活OAC1、ALD5、BAP2、LEU1/2、ILV5、PDC5/6、THI3、ADH2/6、GAP1的表达(或者共表达)，激活或抑制BAT2、ILV2/3表达，抑制ILV6的表达(图7)。

3 展望

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高级醇是包括葡萄酒在内的饮料酒重要风味物质和潜在健康危害因子，主要由酿酒酵母代谢产生。由于酿酒酵母高级醇代谢转录调控机制不明确，导致葡萄酒酿造中针对高级醇含量的控制缺乏有效策略。因此，需要进一步在转录、转录后、翻译和翻译后加工等水平探究高级醇代谢的精细调控。尽管目前已有实验证据表明转录因子Aro80p、GATA和Leu3p与酿酒酵母高级醇代谢有关，同时借助转录组学、基因组学等组学技术预测了更多与高级醇代谢具有潜在关系的转录因子，但是这些转录因子是在无约束条件下的分析结果，无法明确指出哪些转录因子是调控酿酒酵母高级醇合成代谢的关键转录因子。因此，在全基因组范围内挖掘和验证调控酿酒酵母高级醇的关键转录因子，是首要解决的关键科学问题；其次，通过关键转录因子靶向基因的解析，回答在不同发酵阶段关键转录因子通过何种方式调控酿酒酵母高级醇的合成代谢能力，从而揭示酿酒酵母高级醇合成代谢的调控机制，是需要解决的第2个关键科学问题。

对调控酿酒酵母关键转录因子的挖掘及其作用机制的解析，对于深刻系统理解酿酒酵母的高级醇代谢调控机制，为酿酒酵母高级醇代谢的精细调控提供新的思路，也为选育能够产出适量高级醇、适用于包括葡萄酒在内的各类饮料酒生产的优质酿酒酵母菌种，提供了重要的科学依据和实践指导，具有深远的意义。

基金

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昌吉州科技支撑产业高质量发展专项(2022Z03)
宁夏回族自治区重点研发计划(2023BCF01027)
国家现代农业产业技术体系项目(CARS-29-jg-03)
合阳葡萄试验示范站共建专项(2023WNXNZX-2)

参考文献

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2024年第64卷第12期

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文章信息

doi: 10.13343/j.cnki.wsxb.20240305

接收时间：2024-05-16
首发时间：2026-03-21
出版时间：2024-09-26

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收稿日期：2024-05-16
录用日期：2024-09-25

基金

Changji State Science and Technology Support High-quality Industrial Development Project(2022Z03)

昌吉州科技支撑产业高质量发展专项(2022Z03)

Ningxia Hui Autonomous Region Key Research and Development Project(2023BCF01027)

宁夏回族自治区重点研发计划(2023BCF01027)

National Modern Agricultural Industry Technology System Project(CARS-29-jg-03)

国家现代农业产业技术体系项目(CARS-29-jg-03)

Heyang Viti-Viniculture Station Joint Construction Project(2023WNXNZX-2)

合阳葡萄试验示范站共建专项(2023WNXNZX-2)

作者信息

1 西北农林科技大学葡萄酒学院, 陕西杨凌 712100

2 国家林业和草原局葡萄与葡萄酒工程技术研究中心, 陕西杨凌 712100

3 西北农林科技大学合阳葡萄试验示范站, 陕西渭南 714000

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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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