高粱蜀黍苷生物合成与代谢调控机制及无/低蜀黍苷种质创新策略

doi:10.3864/j.issn.0578-1752.2026.15.003

中国农业科学 ›› 2026, Vol. 59 ›› Issue (15): 3267-3282.doi: 10.3864/j.issn.0578-1752.2026.15.003

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

高粱蜀黍苷生物合成与代谢调控机制及无/低蜀黍苷种质创新策略

吕孝松¹(), 赖上坤², 朱超³, 刘薇¹, 夏前浩², 姜雷³, 李桂英⁴, 张执金¹, 朱莉¹()

¹ 中国农业科学院生物技术研究所，北京 100081
² 江苏省农业科学院宿迁农科所，江苏宿迁 223800
³ 江苏洋河酒厂股份有限公司，江苏宿迁 223800
⁴ 中国农业科学院作物科学研究所，北京 100081

收稿日期:2025-12-30 接受日期:2026-02-21 出版日期:2026-08-01 发布日期:2026-08-03
通信作者:
朱莉，E-mail：zhuli01@caas.cn
联系方式: 吕孝松，E-mail：lvxiaosong2663@163.com。
基金资助:
江苏省科技计划(BE2023345); 国家自然科学基金(32272054); 国家重点研发计划(2023YFD1200705)

Regulatory Mechanisms of the Biosynthesis and Metabolism of Dhurrin and the Innovation Strategies for Sorghum Germplasm with Non- or Low- Dhurrin Content

LÜ XiaoSong¹(), LAI ShangKun², ZHU Chao³, LIU Wei¹, XIA QianHao², JIANG Lei³, LI GuiYing⁴, ZHANG ZhiJin¹, ZHU Li¹()

¹ Biotechnology Research Institute, Chinese Academy of Agricultural Sciences, Beijing 100081
² Suqian Institute of Agriculatural Sciences, Jiangsu Academy of Agricultural Sciences, Suqian 223800, Jiangsu
³ Jiangsu Yanghe Distillery Co., Ltd., Suqian 223800, Jiangsu
⁴ Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081

Received:2025-12-30 Accepted:2026-02-21 Published:2026-08-01 Online:2026-08-03

摘要/Abstract

摘要：

高粱作为全球第五大禾谷类作物，在保障粮食安全、支撑饲料生产及推动酿造工业发展方面具有不可替代的战略价值。高粱特有的产氰糖苷——蜀黍苷（dhurrin），在植物防御与产业应用中呈现双重特性：一方面，作为关键的抗逆代谢产物，蜀黍苷在抵御干旱、盐碱和病虫害等逆境胁迫中发挥重要作用，增强植株的环境适应能力；另一方面，在组织受损或加工过程中，蜀黍苷可被β-葡萄糖苷酶水解，释放出有毒的氰化氢（HCN），不仅对畜禽造成中毒风险，还可能影响酿造产品的风味稳定性与品质。本文系统梳理了蜀黍苷在高粱体内的时空分布规律，深入解析其生物合成、活化、解毒和再循环等环节构成的复杂代谢网络及其调控机制。研究表明，蜀黍苷的生物合成主要依赖细胞色素P450酶家族（CYP79A1和CYP71E1）的级联催化，由UGT85B1等糖基转移酶催化糖基化反应，并通过多酶复合物的协同反应完成；其活化过程与植物组织损伤程度密切相关，β-葡萄糖苷酶的水解作用是关键步骤。在饲用领域，未充分处理的高粱秸秆或青贮饲料中残留的蜀黍苷可能引发动物氢氰酸中毒，限制其饲用安全性；在酿造过程中，蜀黍苷降解产物氰化物可进一步反应生成致癌物氨基甲酸乙酯（ethyl carbamate，EC），对白酒的食品安全构成潜在威胁。针对上述风险，本文归纳了包括物理预处理、酶抑制剂添加，以及微生物发酵降解在内的多种减毒策略。更重要的是，基于对蜀黍苷代谢通路的深入认知，本文进一步提出通过分子设计育种创制无或低蜀黍苷高粱新种质的可行性路径，利用CRISPR/Cas9等基因编辑技术定向敲除关键合成基因，旨在从源头上降低风险。本研究为高粱蜀黍苷的代谢定向调控、安全加工与高效利用提供了系统的理论支持与实践指导，对促进高粱全产业链的绿色可持续发展具有重要意义。

关键词: 高粱, 蜀黍苷, 代谢途径, 调控因素, 育种策略

Abstract:

Sorghum bicolor (L.) Moench, as the fifth most important cereal crop globally, plays an irreplaceable strategic role in ensuring food security, supporting feed production, and promoting the development of the brewing industry. Dhurrin, as cyanogenic glycoside unique to sorghum, exhibits a dual role in plant defense and industrial applications. On one hand, as a key stress-responsive metabolite, dhurrin enhances plant resilience against adverse stresses such as drought, salinity, and pests and diseases. On the other hand, upon tissue damage or during processing, dhurrin can be hydrolyzed by β-glucosidase to release toxic hydrogen cyanide (HCN), posing potential risks of livestock health and brewing quality. This paper systematically reviews the spatiotemporal distribution characteristics of dhurrin, elucidates its metabolic network (encompassing biosynthesis, activation, detoxification, and recycling) and regulatory mechanisms. Research indicates that the biosynthesis of dhurrin mainly relies on the sequential catalysis of cytochrome P450 enzyme family (CYP79A1 and CYP71E1), with the glycosylation reaction catalyzed by glycosyltransferases such as UGT85B1, and is completed through the cooperative reaction of multi-enzyme complexes; its activation process is closely related to the degree of plant tissue damage, and the hydrolysis action of β-glucosidase is a key step. In the field of animal feed utilization, residual dhurrin in insufficiently processed sorghum stalks or silage may induce HCN poisoning in animals, limiting its safety as feedstock. During the brewing process, cyanide released from dhurrin degradation can further react to form the carcinogen ethyl carbamate (EC), which poses a potential threat to the food safety of Baijiu (Chinese liquor). In response to the aforementioned risks, this paper summarizes multiple detoxification strategies, including physical pretreatment, enzyme inhibitors, and microbial degradation. More importantly, based on a deep understanding of the dhurrin metabolic pathway, this paper further proposes a feasible approach for developing non- or low-dhurrin sorghum germplasm through molecular breeding. By utilizing gene-editing technologies such as CRISPR/Cas9 to precisely knock out key biosynthetic genes, thereby reducing risks at the source. This research provides comprehensive theoretical support and practical guidance for the targeted regulation of metabolism, safe processing, and efficient utilization of dhurrin in sorghum, which is of great significance for promoting the green and sustainable development of the entire sorghum industry chain.

Key words: Sorghum bicolor, dhurrin, metabolic pathway, regulatory factors, breeding strategy

吕孝松, 赖上坤, 朱超, 刘薇, 夏前浩, 姜雷, 李桂英, 张执金, 朱莉. 高粱蜀黍苷生物合成与代谢调控机制及无/低蜀黍苷种质创新策略[J]. 中国农业科学, 2026, 59(15): 3267-3282.

LÜ XiaoSong, LAI ShangKun, ZHU Chao, LIU Wei, XIA QianHao, JIANG Lei, LI GuiYing, ZHANG ZhiJin, ZHU Li. Regulatory Mechanisms of the Biosynthesis and Metabolism of Dhurrin and the Innovation Strategies for Sorghum Germplasm with Non- or Low- Dhurrin Content[J]. Scientia Agricultura Sinica, 2026, 59(15): 3267-3282.

图/表 4

表1

图1

图2

表2

参与高粱蜀黍苷生物合成与代谢过程的关键基因及其功能"

基因ID Gene ID	酶名称 Enzyme name	主要功能 Main function	参考文献 References
蜀黍苷生物合成 Dhurrin biosynthesis
Sobic.001G012300	细胞色素酶P450 Cytochrome enzyme P450 SbCYP79A1	专一性催化酪氨酸生成对羟基苯乙醛肟，是蜀黍苷生物合成的限速步骤 Specifically catalyze the conversion of tyrosine into p-hydroxyphenylacetaldoxime, and is a key rate-limiting step in the biosynthesis of dhurrin	[17,21,37]
Sobic.001G012200	细胞色素酶P450 Cytochrome enzyme P450 SbCYP71E1	负责催化对羟基苯乙醛肟转化为对羟基乙醛腈，是合成蜀黍苷前体的关键步骤 Response for the conversion of p-hydroxyphenylacetaldoxime to p-hydroxymandelonitrile in dhurrin biosynthesis, which is a crucial step in the synthesis of the precursor of dhurrin	[17,37]
Sobic.001G012400	UDP-葡萄糖基转移酶 UDP-glucosyltransferase SbUGT85B1	参与多酶复合体的形成，其催化的糖基化反应是蜀黍苷生物合成的最后一个步骤 Participate in the formation of the multi-enzyme complex, and its glycosylation reaction is the final step in dhurrin biosynthesis	[17,38,40]
蜀黍苷生物活化Dhurrin bioactivation
Sobic.008G079800 Sobic.008G080400 Sobic.008G080100 Sobic.008G080600	β-葡萄糖苷酶 β -glucosidase DHR1 DHR2 DHR-like 3 DHR-like 4	在植物组织受损时负责催化蜀黍苷水解生成氰醇中间体，是释放HCN的关键步骤 When the tissue is damaged, DHR is responsible for the hydrolysis of dhurrin to generate the cyanohydrin intermediate, which is a key step in releasing HCN	[17,44-45]
Sobic.004G335500	α‑羟腈裂解酶 α -hydroxynitrile lyase HNL	负责将氰醇裂解，释放HCN与对羟基苯甲醛 Responsible for the cleavage of cyanohydrin, releasing HCN and p-hydroxybenzaldehyde (pHB)	[17,44,46]
蜀黍苷脱毒Dhurrin detoxification
Sobic.006G016900	β-氰丙氨酸合成酶 β -cyanoalanine synthase β- CAS C1	负责催化半胱氨酸与剧毒HCN结合生成低毒的β-氰基-L-丙氨酸，是氰化物解毒的核心基因 Responsible for catalyzing the combination of cysteine with the highly toxic HCN to generate the less toxic β-cyanomethyl-L-alanine (BCA), and is the core gene for dhurrin detoxification	[17,47-49,61]
Sobic.004G225200 Sobic.004G225100	硝化酶 Nitrifying enzyme SbNIT4A SbNIT4B1/4B2	负责将HCN中的氮整合生成天冬酰胺、天冬氨酸和氨，以防氰化物中毒 Responsible for integrating the nitrogen in HCN to form asparagine, aspartic acid and ammonia, in order to prevent cyanide poisoning	[17,50]
蜀黍苷再循环Dhurrin recycling
Sobic.002G421200 Sobic.009G033200	谷胱甘肽S转移酶 Glutathione S-transferase SbGSTL1 SbGSTL2	无组织损伤条件下，负责将蜀黍苷转化为对羟基苯基乙腈，并与硝化酶协同作用将蜀黍苷衍生物定向转化为对羟基苯基乙酸（pHPAAc）和游离氨（NH3），且不释放氰化氢 Under non-destructive conditions of the tissue, responsible for converting dhurrin into p- hydroxyphenylacetonitrile, and cooperate with NIT4A/B2 to specifically convert dhurrin derivatives into p-hydroxyphenylacetic acid (pHPAAc) and NH3, without releasing HCN	[17,51]
蜀黍苷转运 Dhurrin transport
Sobic.001G012600	MATE家族液泡转运蛋白 MATE family vacuole transport proteins SbMATE2	SbMATE2具有底物选择性，仅转运非内源蜀黍氰苷，但不能转运花青素-3-O-葡萄糖苷或硫代葡萄糖苷吲哚-3-甲基硫代葡萄糖苷，主要介导蜀黍苷的液泡区室化储存 SbMATE2 has substrate selectivity and only transports non-endogenous dhurrin, but not the anthocyanin cyanidin 3-O-glucoside or the glucosinolate indol-3-yl-methyl glucosinolate. It is mainly mediates the compartmentalized storage of dhurrin in the vacuole.	[17,53]
Sobic.001G133900	NPF家族转运蛋白 NPF family transporter proteins SbCGTR1	与木薯氰苷转运蛋白显著同源，具有长程或跨细胞器转运蜀黍苷的潜力，以避免自毒 SbCGTR1 is highly homologous to the cassava cyanogenic glycoside transporter and has the potential for long- distance or inter-organelle transport of dhurrin to avoid autotoxicity	[17,54]
蜀黍苷生物合成调控因子 Transcription factors regulating the dhurrin biosynthesis
Sobic.010G173400	GATA转录因子 GATA transcription factor SbGATA22	属于具LLM结构域的GATA转录因子，可能是蜀黍苷生物合成负调控因子 Belongs to B-GATA transcription factor with LLM domain and may act as a negative regulator of dhurrin biosynthesis in sorghum	[17,66]

表2

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分类维度 Classification dimension	关键特征 Key characteristics	主要影响因素 Primary influencing factors	参考文献 References
品种差异 Varietal differences	高氰品种：30.26—136.70 mg·kg^-1；低氰品种：<1.00 mg·kg^-1；非糯性>糯性；野生种：低氰或无氰；驯化种：高氰 High dhurrin varieties: 30.26-136.70 mg·kg^-1; Low dhurrin varieties: <1.00 mg·kg^-1; Non-glutinous sorghum > glutinous sorghum; Wild species: low dhurrin or dhurrin-free; Domesticated species: high dhurrin	基因型（CYP79A1-CYP71E1- UGT85B1） Geneotype (CYP79A1-CYP71E1-UGT85B1)	[7-12]
组织分布 Tissue distribution	幼苗叶片>茎秆>籽粒（成熟期痕量）；新生组织（叶腋）>主茎 Young leaves>Stem>Mature seeds (trace levels); New tissues (leaf axils)>Main stem	发育阶段与防御需求 Developmental stage and defense demand	[16-20]
环境调控 Environmental regulation	干旱、施氮、机械损伤等胁迫显著提升含量；西南产区含量普遍低于东北/华中产区 Drought, nitrogen application, mechanical damage significantly increase content; Content in Southwest region generally lower than that in Northeast/Central China	ABA/JA信号通路、环境条件 ABA/JA signaling pathways, environmental conditions	[7,25-31]
动态变化 Dynamic changes	幼苗期含量最高，随生长下降；成熟秸秆高于籽粒；再生苗含量回升；受昼夜节律调控 Highest content in seedling stage, decreases with growth; Mature stems higher than seeds; Regenerated seedlings show increased content; may be regulated by the circadian rhythm	生长阶段与再生防御需求 Growth stage and regeneration defense needs	[17,32-36]

高粱蜀黍苷生物合成与代谢调控机制及无/低蜀黍苷种质创新策略

Regulatory Mechanisms of the Biosynthesis and Metabolism of Dhurrin and the Innovation Strategies for Sorghum Germplasm with Non- or Low- Dhurrin Content

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