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Article(id=1218130663195202065, tenantId=1146029695717560320, journalId=1146031591421210625, issueId=1218130661861409543, articleNumber=null, orderNo=21, doi=10.3981/j.issn.1000-7857.2025.05.00157, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1748188800000, receivedDateStr=2025-05-26, revisedDate=1764172800000, revisedDateStr=2025-11-27, acceptedDate=null, acceptedDateStr=null, onlineDate=1768354581878, onlineDateStr=2026-01-14, pubDate=1766851200000, pubDateStr=2025-12-28, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1768147200000, onlineIssueDateStr=2026-01-12, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1768354581878, creator=13701087609, updateTime=1774080473837, updator=sys-migrate, issue=Issue{id=1218130661861409543, tenantId=1146029695717560320, journalId=1146031591421210625, year='2025', volume='43', issue='24', pageStart='1', pageEnd='119', issueExtLink='null', onlineDate='null', pubDate='1766851200000', pubDateStr='2025-12-28', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1768354581561, creator='13701087609', updateTime=1774330540257, updator='13041195026', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1243195649395634850, tenantId=1146029695717560320, journalId=1146031591421210625, issueId=1218130661861409543, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1243195649399829155, tenantId=1146029695717560320, journalId=1146031591421210625, issueId=1218130661861409543, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=92, endPage=100, ext={EN=ArticleExt(id=1218130663501386258, articleId=1218130663195202065, tenantId=1146029695717560320, journalId=1146031591421210625, language=EN, title=Schizosaccharomyces pombe: An emerging platform for efficient ergothioneine production, columnId=1150494644690366681, journalTitle=Science & Technology Review, columnName=Papers, runingTitle=null, highlight=null, articleAbstract=

Ergothioneine (EGT) is a potent natural sulfur−containing antioxidant with broad application potential in the pharmaceutical, cosmetic, and nutraceutical industries. However, its traditional production methods are inefficient and fail to meet market demand. Recent advancements in synthetic biology offer promising avenues for the efficient bio−manufacturing of EGT. This review systematically summarizes the research progress on using Schizosaccharomyces pombe as a promising chassis organism for EGT production. It focuses on its endogenous biosynthetic capabilities and metabolic engineering strategies, such as promoter engineering, nutrient stress regulation, and mutagenesis screening. We also discuss the major challenges hindering the industrial application of S. pombe, including gaps in fundamental knowledge, unclear physiological functions of EGT, and a lack of standardized analytical methods. Finally, future research directions are proposed, including elucidating the metabolic regulatory network, integrating green production processes, and establishing standardized evaluation systems. This review aims to provide a theoretical foundation for the further development and optimization of S. pombe as a robust platform for EGT synthesis.

, authors=null, authorsList=Zhouqing LUO, Kejing FENG, Dong XU, Mijia DING, Yuhe LIN, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=All rights reserved. Unauthorized reproduction is prohibited., 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=1218130665837613602, articleId=1218130663195202065, tenantId=1146029695717560320, journalId=1146031591421210625, language=CN, title=裂殖酵母：麦角硫因高效生产的潜力宿主, columnId=1146540929516700224, journalTitle=科技导报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

麦角硫因（ergothioneine, EGT）是一种天然高效的含硫抗氧化剂，在医药、化妆品和功能性食品等领域具有广泛的应用前景。然而，其传统生产方法效率低下，难以满足市场需求。近年来，合成生物学技术的发展为EGT的高效生物制造提供了新思路。介绍了裂殖酵母（Schizosaccharomyces pombe）作为潜力宿主在EGT生产中的研究进展，重点分析了其内源合成能力、代谢工程改造策略（如启动子优化、营养胁迫调控和诱变筛选）。探讨了裂殖酵母在工业化应用中面临的主要挑战，包括相关领域存在空白，EGT生理功能不明确和缺乏标准化检测方法等，并提出解析代谢调控网络、整合绿色生产工艺和建立标准化评价体系等未来研究方向，例如，为裂殖酵母作为EGT生产底盘的进一步开发和优化提供参考。

, authors=

罗周卿，教授，研究方向为基因组设计与合成、合成生物学及代谢工程，电子信箱：

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罗周卿，教授，研究方向为基因组设计与合成、合成生物学及代谢工程，电子信箱：

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罗周卿，教授，研究方向为基因组设计与合成、合成生物学及代谢工程，电子信箱：

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（a） 在细菌和真菌中，EGT的有氧合成途径；（b） EGT无氧合成途径

, figureFileSmall=ZeBYb96DMhLsTj2aAt0h8Q==, figureFileBig=m5zZaG2csw+zG6sHvus8wA==, tableContent=null), ArticleFig(id=1242146818830709723, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=EN, label=null, caption=null, figureFileSmall=U9VQk7QMsrr5b9PdaU/fTA==, figureFileBig=BWF3QRn/5UdlR/IYqsmI7g==, tableContent=null), ArticleFig(id=1242146818906207196, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=CN, label=图3, caption=EGT化学合成法的经典技术路线, figureFileSmall=U9VQk7QMsrr5b9PdaU/fTA==, figureFileBig=BWF3QRn/5UdlR/IYqsmI7g==, tableContent=null), ArticleFig(id=1242146818981704669, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=EN, label=null, caption=null, figureFileSmall=uhgczsZTagXTu1YO7sHW+g==, figureFileBig=8KN1dQ0PpltoIHVd+CumYw==, tableContent=null), ArticleFig(id=1242146819048813534, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=CN, label=图4, caption=裂殖酵母作为底盘生产EGT的优势, figureFileSmall=uhgczsZTagXTu1YO7sHW+g==, figureFileBig=8KN1dQ0PpltoIHVd+CumYw==, tableContent=null), ArticleFig(id=1242146819111728095, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
方法发明人 合成路线 反应条件 总收率/% 特点
Erdelmeier等[26]
 　以组氨酸三甲基内盐为原料，经半胱氨酸硫化和高温裂解2步反应获得麦角硫因 　液溴、3−甲基丙烯酸、盐酸等溶剂，加热 40 　模仿生物合成路线，产物易于分离纯化，但原料价格昂贵
Khonde和Jardine[27] 　以N−叔丁氧羰基−1−苄基−L−组氨酸为原料，经溴化脱苄基、半胱氨酸硫化、氧化、化学或酶促裂解4步反应得到麦角硫因 　N,N−二甲基甲酰胺等溶剂，过氧化氢等氧化剂 70 　模仿生物合成路线，总收率较高，但原料价格昂贵，且需多步层析柱纯化
马晓雪等[25] 　以组氨酸为原料，经还原、甲基化、半胱氨酸硫化、高温裂解4步反应得到麦角硫因 　钯碳催化，液溴、甲醇等溶剂，加热 47 　模仿生物合成路线，反应条件较温和，但反应效率较低
), ArticleFig(id=1242146819183031264, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=CN, label=表1, caption=

EGT化学合成法不同路线的比较

, figureFileSmall=null, figureFileBig=null, tableContent=
方法发明人 合成路线 反应条件 总收率/% 特点
Erdelmeier等[26]
 　以组氨酸三甲基内盐为原料，经半胱氨酸硫化和高温裂解2步反应获得麦角硫因 　液溴、3−甲基丙烯酸、盐酸等溶剂，加热 40 　模仿生物合成路线，产物易于分离纯化，但原料价格昂贵
Khonde和Jardine[27] 　以N−叔丁氧羰基−1−苄基−L−组氨酸为原料，经溴化脱苄基、半胱氨酸硫化、氧化、化学或酶促裂解4步反应得到麦角硫因 　N,N−二甲基甲酰胺等溶剂，过氧化氢等氧化剂 70 　模仿生物合成路线，总收率较高，但原料价格昂贵，且需多步层析柱纯化
马晓雪等[25] 　以组氨酸为原料，经还原、甲基化、半胱氨酸硫化、高温裂解4步反应得到麦角硫因 　钯碳催化，液溴、甲醇等溶剂，加热 47 　模仿生物合成路线，反应条件较温和，但反应效率较低
), ArticleFig(id=1242146819258528737, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
物种关键策略发酵时间/h产量生产效率/
（mg·L−1·h−1）
大肠杆菌[29]　引入EgtB/EgtD/EgtE；过表达CysE/SerA/YdeD；敲除MetJ192657 mg·L−13.4
大肠杆菌[16]　引入EgtD/EgtE和截短Egt1；随机突变筛选EgtD和Egt1；补料分批发酵965.4 g·L−156.3
新金色分枝杆菌[30]　 Mn_3042基因敲除；EgtABCDE/metK/hisG基因过表达；补料分批发酵2161.56 g·L−17.2
谷氨酸棒状杆菌[31]　 Egt1/Egt2基因引入；CysE/CysK/CysR基因过表达；SdaA基因敲除；强化硫同化与磷酸戊糖途径36264 mg·L−17.3
枯草芽孢杆菌[32]　 EgtABCDE基因引入；培养基优化60568 mg·L−19.5
米曲霉[33]　引入Egt1/Egt2120231 mg·kg−11.9
圆红冬孢酵母[34]　 RtEGT1/RtEGT2/SAM2/SAH1基因过表达；培养基优化168267 mg·L−11.6
解脂耶氏酵母[35]　 Egt1/Egt2基因引入；磷酸盐限制策略；补料分批发酵2201.63 g·L−17.4
酿酒酵母[36]　 Egt1/Egt2基因引入；培养基优化；补料分批发酵84598 mg·L−17.1
酿酒酵母[37]　 Egt1/Egt2基因引入；MET14基因过表达；spe2基因敲除；组氨酸高产菌株的诱变与筛选1602.39 g·L−115.0
), ArticleFig(id=1242146819447272418, tenantId=1146029695717560320, journalId=1146031591421210625, articleId=1218130663195202065, language=CN, label=表2, caption=

各物种中EGT产量

, figureFileSmall=null, figureFileBig=null, tableContent=
物种关键策略发酵时间/h产量生产效率/
（mg·L−1·h−1）
大肠杆菌[29]　引入EgtB/EgtD/EgtE；过表达CysE/SerA/YdeD；敲除MetJ192657 mg·L−13.4
大肠杆菌[16]　引入EgtD/EgtE和截短Egt1；随机突变筛选EgtD和Egt1；补料分批发酵965.4 g·L−156.3
新金色分枝杆菌[30]　 Mn_3042基因敲除；EgtABCDE/metK/hisG基因过表达；补料分批发酵2161.56 g·L−17.2
谷氨酸棒状杆菌[31]　 Egt1/Egt2基因引入；CysE/CysK/CysR基因过表达；SdaA基因敲除；强化硫同化与磷酸戊糖途径36264 mg·L−17.3
枯草芽孢杆菌[32]　 EgtABCDE基因引入；培养基优化60568 mg·L−19.5
米曲霉[33]　引入Egt1/Egt2120231 mg·kg−11.9
圆红冬孢酵母[34]　 RtEGT1/RtEGT2/SAM2/SAH1基因过表达；培养基优化168267 mg·L−11.6
解脂耶氏酵母[35]　 Egt1/Egt2基因引入；磷酸盐限制策略；补料分批发酵2201.63 g·L−17.4
酿酒酵母[36]　 Egt1/Egt2基因引入；培养基优化；补料分批发酵84598 mg·L−17.1
酿酒酵母[37]　 Egt1/Egt2基因引入；MET14基因过表达；spe2基因敲除；组氨酸高产菌株的诱变与筛选1602.39 g·L−115.0
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裂殖酵母：麦角硫因高效生产的潜力宿主
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罗周卿 , 冯科景 , 徐冬 , 丁汨佳 , 林宇禾
科技导报 | 研究论文 2025,43(24): 92-100
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裂殖酵母：麦角硫因高效生产的潜力宿主
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罗周卿，教授，研究方向为基因组设计与合成、合成生物学及代谢工程，电子信箱：

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罗周卿，教授，研究方向为基因组设计与合成、合成生物学及代谢工程，电子信箱：

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罗周卿 , 冯科景, 徐冬, 丁汨佳, 林宇禾
作者信息
  • 厦门大学生命科学学院，厦门 361102
Schizosaccharomyces pombe: An emerging platform for efficient ergothioneine production
Zhouqing LUO , Kejing FENG, Dong XU, Mijia DING, Yuhe LIN
Affiliations
  • School of Life Sciences, Xiamen University, Xiamen 361102, China
出版时间: 2025-12-28 doi: 10.3981/j.issn.1000-7857.2025.05.00157
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摘要
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麦角硫因（ergothioneine, EGT）是一种天然高效的含硫抗氧化剂，在医药、化妆品和功能性食品等领域具有广泛的应用前景。然而，其传统生产方法效率低下，难以满足市场需求。近年来，合成生物学技术的发展为EGT的高效生物制造提供了新思路。介绍了裂殖酵母（Schizosaccharomyces pombe）作为潜力宿主在EGT生产中的研究进展，重点分析了其内源合成能力、代谢工程改造策略（如启动子优化、营养胁迫调控和诱变筛选）。探讨了裂殖酵母在工业化应用中面临的主要挑战，包括相关领域存在空白，EGT生理功能不明确和缺乏标准化检测方法等，并提出解析代谢调控网络、整合绿色生产工艺和建立标准化评价体系等未来研究方向，例如，为裂殖酵母作为EGT生产底盘的进一步开发和优化提供参考。

麦角硫因  /  裂殖酵母  /  合成生物学  /  微生物发酵  /  代谢工程

Ergothioneine (EGT) is a potent natural sulfur−containing antioxidant with broad application potential in the pharmaceutical, cosmetic, and nutraceutical industries. However, its traditional production methods are inefficient and fail to meet market demand. Recent advancements in synthetic biology offer promising avenues for the efficient bio−manufacturing of EGT. This review systematically summarizes the research progress on using Schizosaccharomyces pombe as a promising chassis organism for EGT production. It focuses on its endogenous biosynthetic capabilities and metabolic engineering strategies, such as promoter engineering, nutrient stress regulation, and mutagenesis screening. We also discuss the major challenges hindering the industrial application of S. pombe, including gaps in fundamental knowledge, unclear physiological functions of EGT, and a lack of standardized analytical methods. Finally, future research directions are proposed, including elucidating the metabolic regulatory network, integrating green production processes, and establishing standardized evaluation systems. This review aims to provide a theoretical foundation for the further development and optimization of S. pombe as a robust platform for EGT synthesis.

ergothioneine  /  Schizosaccharomyces pombe  /  synthetic biology  /  microbial fermentation  /  metabolic engineering
罗周卿, 冯科景, 徐冬, 丁汨佳, 林宇禾. 裂殖酵母：麦角硫因高效生产的潜力宿主. 科技导报, 2025 , 43 (24) : 92 -100 . DOI: 10.3981/j.issn.1000-7857.2025.05.00157
Zhouqing LUO, Kejing FENG, Dong XU, Mijia DING, Yuhe LIN. Schizosaccharomyces pombe: An emerging platform for efficient ergothioneine production[J]. Science & Technology Review, 2025 , 43 (24) : 92 -100 . DOI: 10.3981/j.issn.1000-7857.2025.05.00157
正文
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麦角硫因（ergothioneine，EGT）是一种天然的含硫组氨酸衍生物，于1909年由Charles从麦角菌Ergot（Clavieps purpurea Tul.）中分离，并以此菌命名[1]。1956年，Garay通过实验证明，EGT能够消除过氧化氢对麦角菌孢子萌发的抑制[2]。此后半个多世纪，EGT作为一种天然抗氧化剂的生理功能和应用潜力逐渐得到广泛的研究和开发。随着EGT市场需求快速增长，现有生产方法因效率低、产量不足而难以满足需求，亟需开发高效且经济的新型生产策略[3]。近年来，随着EGT生物合成途径的探明和合成生物学技术的发展，研究热点集中在对微生物进行改造用于发酵生产EGT，其中对裂殖酵母（Schizosaccharomyces pombe，S. pombe）的改造包含了多种策略并取得了显著的成果。然而，与大肠杆菌（Escherichia coli）和酿酒酵母（Saccharomyces cerevisiae）等模式生物相比，裂殖酵母的遗传改造研究仍较匮乏，且缺乏高效的基因操作工具。这对未来进一步提高EGT产量的研究构成了挑战。展望未来，随着合成生物学技术的持续进步，裂殖酵母有望成为EGT工业生产的关键宿主，为EGT的商业化应用开辟新的道路。
1 EGT的生理功能和市场应用
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1.1 EGT的生理功能
EGT已知的作用机制包括直接清除活性氧（ROS）、螯合金属阳离子以抑制ROS产生、提高抗氧化基因表达、调节抗氧化相关酶活性等[4]。广泛的抗氧化机制使EGT能够保护细胞免受氧化损伤，因此对多种疾病都具有一定疗效，如：在炎症治疗中，EGT可调节细胞炎症因子水平，减少细胞氧化损伤，发挥抗炎作用[5]；在癌症治疗中，EGT能够增强巨噬细胞反应，并调节肿瘤微环境以增强癌症疫苗的作用效果[5]；EGT还可调节神经元分化和神经发生，降低神经病变，对抑郁和认知障碍等神经系统疾病也具有一定效果[6]。
同时，EGT强大的抗氧化能力使其在化妆品和功能食品领域也扮演着重要角色。EGT可通过清除ROS、抑制光老化等机制显著减轻紫外线诱导的皮肤损伤与老化，兼具抗紫外线、美白、抗皱及延缓衰老等多重功效，是极具潜力的化妆品原料[6]；同时，EGT作为一种人体不能合成的膳食来源抗氧化剂，几乎存在于人体所有组织和细胞中，被认为可能是一种维生素，可制成膳食补充剂，参与保护易受氧化损伤的组织细胞，如眼组织和心血管等[7]。
1.2 EGT的市场应用
目前，EGT已凭借其卓越的抗氧化性能，在多个领域展现出广阔的应用前景（图1）[4−7]，是当下热门的原料产品。近年来，全球对EGT的需求呈现出稳定增长态势，根据QYResearch（北京恒州博智国际信息咨询有限公司）的统计，2022年全球EGT市场销售额达到了4.63亿元，预计2029年将达到8.93亿元，年复合增长率为14.46%（2023—2029年）；地区层面来看，中国市场在过去几年变化较快，2022年市场规模为2237万元，约占全球的4.83%，预计2029年将达到18580万元，届时全球占比将达到20.80%。未来几年，中国EGT市场规模有望持续扩大，其经济价值也将不断攀升[8]。
2014年，EGT被中国国家市场监督管理总局正式列入化妆品原料清单[9]。2017年，欧盟委员会批准EGT作为一种新的食品添加成分，同年，美国食品和药物管理局还授予EGT GRAS认证（Generally Recognized as Safe），这一决定使产品制造商能够将EGT应用至各种消费品中，包括化妆品、食品、饮料和保健品[10]。多项安全认证加速了EGT在市场上的应用，目前全球已有多家药企和膳食补充剂品牌推出含有EGT的功能性食品，多家著名化妆品企业也推出了含有EGT的护肤品，包括雅诗兰黛、迪奥、倩碧[11]。随着经济的持续增长以及人们健康意识的不断提高，EGT的需求将稳定增长，市场前景愈发繁荣。
2 EGT的生物合成途径和工业生产方法
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2.1 EGT的生物合成途径
已知EGT的生物合成途径可分为有氧合成途径与无氧合成途径2类[12−14]。其中，有氧合成途径进一步细分为细菌途径（涉及EgtA至EgtE酶系）和真菌途径（涉及Egt1与Egt2酶）（图2（a））。至于无氧合成途径，其分布相对广泛，主要涉及的酶为EanA与EanB（图2（b））。此外，部分厌氧细菌和古细菌则通过编码金属蝶呤依赖性EGT合成酶（metallopterin−dependent ergothioneine synthase，MES）来替代EanB酶，从而实现EGT的合成[3,15]。
在细菌合成途径中，甲基转移酶EgtD催化S−腺苷甲硫氨酸（SAM）的甲基转移至组氨酸，形成组氨酸三甲基内盐，此为EGT合成的起始步骤；同时，谷氨酸−半胱氨酸连接酶EgtA将谷氨酸和半胱氨酸转化为γ−谷氨酰半胱氨酸（γ−glutamyl−cysteine）。随后，以二价铁和氧气作为辅助因子，EgtB酶催化γ−谷氨酰半胱氨酸与组氨酸三甲基内盐反应生成γ−谷氨酰−组氨酸三甲基内盐基半胱氨酸亚砜，EgtC酶进一步催化产生组氨酸三甲基内盐基半胱氨酸亚砜。最后，β−裂解酶EgtE与磷酸吡哆醛（PLP）结合催化组氨酸三甲基内盐基半胱氨酸亚砜脱去丙酮酸和氨，最终生成EGT[3]。
在真菌途径中，Egt1酶同时具备甲基转移酶和亚砜合酶2个结构域，能够替代细菌途径里EgtB、EgtC、EgtD 3种酶的功能，无需谷氨酸参与合成中间产物，直接催化组氨酸与SAM持续反应，直至与半胱氨酸反应生成组氨酸三甲基内盐基半胱氨酸亚砜。Egt2酶与细菌途径中的EgtE酶相似，负责催化生成EGT的最终步骤[3,16]。
在无氧合成途径中，EGT的生物合成涉及2步核心反应，分别为甲基转移和硫转移。在研究较广泛的EanA/EanB酶反应中，甲基转移酶EanA先将SAM的3个甲基转移到组氨酸上生成组氨酸三甲基内盐，与细菌途径EgtD催化反应类似；随后硫转移酶EanB将来自多硫化物的硫原子直接转移至Hercynine的咪唑环上，最终形成EGT。在MES反应中，使用MES代替EanB进行硫转移反应，硫源为半胱氨酸[15,17]。
尽管无氧途径在反应机制上更为简洁，但相比有氧途径，无氧途径的催化效率较低，已鉴定的关键酶EanA和EanB的催化速率（分别为kcat=3.8 min–1和0.5 min–1）远低于有氧途径中EgtD和EgtB的催化速率（分别为kcat=35 min−1和72 min−1）[18]。这一差异表明，在EGT的生物合成过程中，有氧途径因其较高的催化速率而更为高效。而在有氧途径中，相较于细菌途径，真菌途径在步骤上更为简化，涉及的酶种类也更少，简化的酶促反应流程不仅减少了中间步骤，降低了副反应和中间产物损失的风险，而且不需谷氨酸参与，消除了EGT与谷胱甘肽生物合成途径之间的底物竞争，进一步提升了EGT的生物合成效率[3]。
2.2 EGT的工业生产方法
常规生产EGT的方法包括化学合成法和提取法，其中化学合成涉及一系列精确的化学反应来生成EGT。1950年，Health等[19]开发了一种合成EGT的方法，其中包括制备2−硫代组氨酸、引入保护基团、甲基化以及最终的水解和纯化。然而，该方法存在产物易发生消旋化的缺陷。1996年，Xu和Yadan [20]开发了一种新方法，通过有效控制反应条件来避免消旋化，关键步骤是直接转化生成咪唑−2−硫酮。尽管如此，该方法仍无法完全避免消旋化，且过程繁琐，给工业化带来了重大挑战。2023年，中国科学院上海有机化学研究所取得突破，研发了大规模化学合成组氨酸甜菜碱的方法，并利用工程化的裂殖酵母EGT合酶SpEgt1−tr M10和SpEGT2M3实现了EGT的高效合成[21]。目前，由南京艾希帝生物科技有限公司开发的三苯胺基保护法是化学合成法中收率最高的（图3）[22]。尽管化学合成方法在不断进步，但已开发的方法仍面临诸多挑战，例如产物消旋化、多步反应复杂、纯化困难以及原材料成本高昂，所有这些因素都导致了最终产品的价格居高不下[23−25]（表1[25−27]）。
提取法虽应用广泛，但需以蘑菇等担子菌为原料。由于原料中EGT含量低、杂质复杂，该法需消耗大量生物质，且纯化难度高。此外，提取过程涉及发酵、提取和多步纯化等多个繁琐步骤，这限制了提取效率，也使得产量难以提高[28]。
因此，虽然化学合成法和提取法为EGT的制备提供了可用途径，但在成本效益、原料供应和产品纯度方面仍存在局限，致使EGT长期价格偏高，每克售价一度高达约1900$[3,26]。相比之下，利用生物合成发酵获取EGT更为安全、高效，近年来国内外通过改造微生物进行发酵获取EGT的研究取得实质性进展并获得专利，展现出广阔的应用潜力（表2[16,29−37]）。
但目前工业生产中利用微生物发酵生产EGT也存在一些瓶颈。首先，使用生物发酵生产EGT存在一定污染。据报道，每生成1 t活性干酵母，便会伴生约150 t的废水[38]，这一环境问题是制约生物发酵法发展的一大痛点。其次，目前行业缺少对于EGT品质的标准化评判，例如对其活性的评估等。统一标准的缺失，也是EGT生产领域中的一大制约。
3 裂殖酵母作为宿主生物的优势
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3.1 裂殖酵母的内源EGT合成能力
裂殖酵母基因组中已鉴定出EGT合成关键基因Egt1和Egt2，其野生型菌株胞内可检测到内源性EGT，表明该酵母具备天然合成EGT的能力[39]，相比绝大多数模式生物，裂殖酵母本身就具备合成EGT的能力，很有可能通过代谢工程改造以进一步提升其EGT的合成和积累能力（图3）。同时，裂殖酵母作为一种好氧真核生物，其EGT合成途径属于有氧真菌途径，相较其他生物合成途径更为简化、高效。
3.2 裂殖酵母是优秀的模式生物
裂殖酵母作为一种易于培养和操作的真核模式生物（图4），以快速的增殖速度和简单的遗传系统而受到科研人员的青睐，基因组较小（全长约14 Mb）且已经全部测序，这为遗传改造提供了便利。同时相对于非模式生物，裂殖酵母的转化效率很高，使用醋酸锂转化法可以产生较多的转化子，适合进行高效的遗传操作，拥有极佳的代谢工程潜力[40]。
4 代谢工程改造裂殖酵母的策略
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目前已知的代谢工程改造裂殖酵母的方法包括启动子改造、培养基碳饥饿和氮饥饿、基因突变等，均能有效提高EGT产量。
4.1 启动子改造
裂殖酵母自身具备EGT合成基因，即Egt1和Egt2，提高EGT产量的思路之一是通过强启动子来增强Egt1或Egt2的表达，从而促进EGT合成。
为了探索裂殖酵母内源EGT合成通路的关键基因，Pluskal等[39]对裂殖酵母中的Egt1和Egt2基因分别进行了敲除，实验结果显示，敲除Egt1后裂殖酵母EGT产量降为0，而敲除Egt2后，EGT的合成虽然减少，但细胞仍然能够合成EGT，表明Egt2可能不是EGT合成的关键基因。后续实验表明，在磷酸吡哆醛（PLP）存在的情况下，裂殖酵母中的组氨酸三甲基内盐基半胱氨酸亚砜能自发地转化为EGT，研究者推测反应可能由其他PLP结合酶催化完成（裂殖酵母基因组包含至少26种不同的PLP结合酶），因此判断裂殖酵母EGT合成的关键基因仅为Egt1，Egt1催化反应被认为是EGT合成的限速步骤，而Egt2催化反应不是。基于这些发现，推测强化Egt1的启动子应当能有效提高EGT产量。
后续的研究中，研究者利用nmt1启动子来增强Egt1基因的表达。nmt1启动子是裂殖酵母的诱导性强启动子，在缺乏硫胺素的条件下具有很高的诱导强度[41]。研究者将Egt1原始启动子替换成nmt1启动子，并在没有硫胺素的培养基中进行培养。通过代谢组学分析，他们证实了在改造后的裂殖酵母中，EGT及其前体物质组氨酸三甲基内盐和组氨酸三甲基内盐基半胱氨酸亚砜得到了积累，并且EGT胞内浓度从0.3 μM提高至1606 μM，相比野生型增加了超过5000倍[39]。这一结果说明通过启动子替换过表达Egt1基因能有效提高EGT产量。
4.2 培养基碳饥饿
Pluskal等[42]的另一项研究探究了裂殖酵母在碳饥饿状态的代谢变化。实验数据显示，当裂殖酵母处于葡萄糖浓度低于2.2 mM的条件时，细胞内SAM的浓度增加了约20倍。SAM是Egt1第一步反应的底物，理论上其浓度上升能够促进组氨酸三甲基内盐的产生，而组氨酸三甲基内盐作为EGT合成的重要中间体，其产量的增加能进一步促进EGT的合成，最终提高EGT的产量。
同时，实验表明当培养基的葡萄糖浓度低于2.2 mM时，裂殖酵母的分裂速度大幅减慢，在1.1 mM葡萄糖浓度下，不分裂的细胞数量显著增加。因此，研究者将1.1 mM的葡萄糖浓度定义为“碳饥饿”状态，这是一个关键的阈值，低于此浓度，裂殖酵母的细胞生长和分裂受到抑制。于是Pluskal等[42]采用1.1 mM的葡萄糖浓度进行实验，这一浓度能够在不显著抑制细胞生长的前提下模拟碳源限制的环境。实验数据表明，在这种碳饥饿条件下，EGT的产量得到了显著提升，EGT胞内浓度从0.3 μM提高至41.6 μM。
4.3 培养基氮饥饿
Pluskal等[42]还对裂殖酵母进行了氮饥饿处理（在培养基中去除氮源，即不添加NH4Cl），在处理60 min后，裂殖酵母细胞内的组氨酸三甲基内盐基半胱氨酸亚砜含量增加了超过10倍，理论上可以为EGT的合成提供更多的前体物质。这一发现表明，氮饥饿条件可能通过影响裂殖酵母的氮代谢，进而加速EGT合成途径中的关键反应，导致EGT合成效率的提升。Pluskal等[39]还检测了裂殖酵母在氮饥饿培养条件下的EGT产量，实验数据显示EGT胞内浓度从0.3 μM提高至157.4 μM，表明氮饥饿能够有效提高裂殖酵母EGT产量。
4.4 发展方向：借鉴酿酒酵母的工程化思路
4.4.1 EGT合成途径改造
Egt1与Egt2是真菌有氧合成麦角硫因过程中的2个核心基因。在酿酒酵母的研究中，van der Hoek 等[36]通过筛选不同直系同源基因组合，发现来自粗糙脉孢菌（Neurospora crassa）的Egt1与麦角菌的Egt2组合能够获得最高产量，其中粗糙脉孢菌来源的Egt1尤为关键。在裂殖酵母中亦可借鉴该策略，尝试引入外源EGT合成途径。
此外，基因表达水平直接影响相应蛋白质的合成，进而影响麦角硫因的产量，因此提高EGT生物合成相关基因的表达水平是优化生产菌株的有效策略。基于这一思路，van der Hoek等[36]在酵母基因组中额外整合了1拷贝粗糙脉孢菌来源的Egt1与麦角菌来源的Egt2，使EGT产量较单拷贝策略显著提高了80%，在培养84 h后达到（598 ±18） mg·L−1。类似地，在裂殖酵母中通过增强内源EGT合成基因的表达，同样具备进一步提高产量的潜力。
4.4.2 底盘菌株改造
代谢前体的供应提升也是产量提升的重要方式。鉴于组氨酸是EGT生物合成中的关键前体，提高其产量已成为增加EGT产率的有效策略。为此，van der Hoek等[37]开发了一种创新的筛选方法，用于鉴定高产组氨酸的酿酒酵母菌株。研究人员将培养基中的组氨酸替换为其毒性类似物β−（1,2,4−三唑−3−基）−DL−丙氨酸。若菌株在此条件下生长正常，则表明其能够内源合成足够的组氨酸以满足生长需求，从而显示出增强的组氨酸生物合成能力。初步筛选出的菌株随后被分别接种于2种培养基：一种含有30 mM组氨酸，另一种不含组氨酸。鉴于30 mM组氨酸对酿酒酵母具有毒性[43]，将在含组氨酸培养基中生长受阻而在无组氨酸培养基中正常生长的菌株鉴定为具备高产组氨酸潜力的目标菌株。分析结果表明，这些筛选获得的高产组氨酸菌株的麦角硫因产量达到了61 mg·L−1。类似地，有望可以通过相同原理构造组氨酸高产的裂殖酵母菌株以提高EGT产量。
EGT途径中其余的重要代谢前体如SAM等的供应量也十分重要，可以通过对这些代谢途径的关键基因位点进行改造。例如van der Hoek 等[37]系统筛选了27个与EGT前体合成相关的基因，通过基因敲除和过表达实验发现，同时敲除SPE2基因并过表达MET14基因可显著提高EGT产量。硫基作为EGT强抗氧化性能的关键结构，其组成依赖硫元素；MET14基因编码腺苷磷硫酸激酶，在细胞硫利用调控中起核心作用。SPE2基因则编码SAM脱羧酶，是调控细胞内S−腺苷甲硫氨酸（SAM）代谢的关键酶。SAM作为甲基供体，是EGT有氧合成途径中不可或缺的中间体。经160 h培养后，该优化策略使EGT产量达到（2.39±0.08） g·L−1。类似的，在裂殖酵母中也许可以通过敲除或者过表达相关基因进行调整。
5 裂殖酵母生产EGT的挑战与展望
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5.1 挑战
目前有许多对不同宿主生物进行改造以生产EGT的研究。在酵母菌中，酿酒酵母作为研究较为透彻的模式生物，生产应用较广，基因改造技术也较为成熟，是最主流的宿主选择。裂殖酵母相关的改造相对较少，机制研究也远不如酿酒酵母等其他模式生物，使得以裂殖酵母为底盘的代谢改造具备相当大的挑战性。
Pluskal等[39]在对裂殖酵母进行Egt1/Egt2基因敲除的实验中发现，即使敲除了Egt1和Egt2基因，裂殖酵母的生长和形态等各方面并未受到明显影响，这可能表明EGT对裂殖酵母的基本生理活动并无明显作用。而即使在氧化胁迫的条件下，裂殖酵母中Egt1的基因表达水平也没有显著提高，这表明EGT可能不是裂殖酵母应对氧化胁迫的主要策略。可以由此推断在裂殖酵母中生产EGT的挑战性：一方面，EGT的合成似乎不是裂殖酵母生存所必需的，在没有额外的代谢工程干预的情况下，裂殖酵母可能不会将大量资源投入EGT的合成中；另一方面，EGT可能不仅与抗氧化机制相关，其在裂殖酵母中的生理作用尚未明确，参与的调控机制可能比预期的更为复杂，这都增加了通过遗传改造来提高EGT产量的难度。
同时，一个值得注意的事实是，裂殖酵母自身能够合成EGT，但其生理作用并不明确，这与对其他生产EGT物种的研究结果相符。随着越来越多样化的EGT的生产物种和合成机制的发现，许多研究者推测EGT可能事实上具备与目前已知的截然不同的生理作用。例如高浓度EGT是否抑制酵母细胞生长，相关研究是缺乏的。
5.2 展望
目前构建EGT高产菌株的研究层出不穷，在常用模式生物中，改造思路已经较为成熟。其中，通过导入外源EGT合成基因，尤其是粗糙脉孢菌Egt1是目前提升合成EGT能力最有效的方式[37]。而在使用裂殖酵母构建EGT高产菌株的相关实验中，由于裂殖酵母本身具有内源EGT合成通路，引入外源EGT合成基因的实验鲜有见闻，这为更进一步的改造提供了广泛的探索空间。未来可以通过引入来自粗糙脉孢菌的NcEgt1加强EGT真菌途径的合成，也可以通过引入其他常见的EGT合成途径中的相关酶强化表达EGT的合成路径。
因为工业上使用生物发酵法会产生大量的废水对环境造成污染[38]，目前在酿酒酵母生产EGT领域中，有研究团队致力于研究使用甘油这一可再生的工业生产中常见的副产物作为碳源用于EGT生产中[44]。这一举措可以大大降低工业生产EGT对环境的污染，而裂殖酵母生产EGT的工艺在这方面是一片空白，可以进行大胆的开荒实验。
相比酿酒酵母等其他模式生物，目前裂殖酵母相关的研究仍然较少，对其的理解仍远不够透彻。通过代谢工程改造裂殖酵母的研究，不仅能够发挥裂殖酵母的优势、提高EGT的产量、促进EGT商业化应用普及，而且可能有助于我们进一步明确EGT的生理功能，同时增进对裂殖酵母代谢机制的了解。
6 总结
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裂殖酵母作为优良的模式生物，凭借其内源性EGT合成能力、简化的真菌合成途径及卓越的代谢可塑性，具有广阔的发展前景。然而，裂殖酵母的遗传工具匮乏、EGT生理功能不明确及缺乏标准化检测方法等问题仍需突破。
未来研究应聚焦以下方向：（1） 开发高效遗传操作工具，以精准调控代谢通路；（2） 探索EGT在裂殖酵母中的潜在生理作用，解析其代谢调控网络；（3） 整合绿色生产工艺以减少环境负担；（4） 建立EGT活性与纯度的标准化评价体系。随着代谢工程技术的发展，裂殖酵母有望突破EGT生产瓶颈，成为工业化制备EGT的核心宿主生物，为EGT的商业化应用探索出一条新的路径，同时为真核微生物代谢工程提供新范式。
基金
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  • 国家重点研发计划合成生物学专项(2024YFA0916503)
  • 福建省自然科学基金攻青项目(2025J011005)
  • 厦门大学校长基金本科生项目(20720242001)
文献
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doi: 10.3981/j.issn.1000-7857.2025.05.00157
  • 接收时间：2025-05-26
  • 首发时间：2026-01-14
  • 出版时间：2025-12-28
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  • 收稿日期：2025-05-26
  • 修回日期：2025-11-27
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国家重点研发计划合成生物学专项(2024YFA0916503)
福建省自然科学基金攻青项目(2025J011005)
厦门大学校长基金本科生项目(20720242001)
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    厦门大学生命科学学院，厦门 361102
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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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