生物技术通报 ›› 2023, Vol. 39 ›› Issue (5): 44-53.doi: 10.13560/j.cnki.biotech.bull.1985.2022-1249
周定定1,2(), 李辉虎3, 汤兴涌4, 余发新3, 孔丹宇2, 刘毅2()
收稿日期:
2022-10-11
出版日期:
2023-05-26
发布日期:
2023-06-08
通讯作者:
刘毅,男,博士,副研究员,研究方向:药用与功能植物学;E-mail: liuy@lsbg.cn作者简介:
周定定,女,硕士研究生,研究方向:药用与功能植物学;E-mail: 1223918603@qq.com李辉虎同为本文第一作者
基金资助:
ZHOU Ding-ding1,2(), LI Hui-hu3, TANG Xing-yong4, YU Fa-xin3, KONG Dan-yu2, LIU Yi2()
Received:
2022-10-11
Published:
2023-05-26
Online:
2023-06-08
摘要:
甘草是一种被多个国家药典收录的常用中草药,由豆科甘草属特定物种的根和匍匐茎炮制而成。甘草含有丰富的次生代谢物和多种活性成分,具有抗肿瘤、抗菌、抗病毒、抗炎和增强免疫等多方疗效。甘草酸和甘草苷是甘草最重要的活性物质。近年来甘草酸和甘草苷的生物合成和调控的研究进展突出。为此,本文综述了甘草中甘草酸和甘草苷的生物合成途径,以及非生物胁迫、生物因素对合成通路调控的研究进展;也对甘草酸和甘草苷的生物合成及调控的未来研究提出展望,以期对该领域的未来发展带来一定帮助。
周定定, 李辉虎, 汤兴涌, 余发新, 孔丹宇, 刘毅. 甘草酸和甘草苷生物合成与调控的研究进展[J]. 生物技术通报, 2023, 39(5): 44-53.
ZHOU Ding-ding, LI Hui-hu, TANG Xing-yong, YU Fa-xin, KONG Dan-yu, LIU Yi. Research Progress in the Biosynthesis and Regulation of Glycyrrhizic Acid and Liquiritin[J]. Biotechnology Bulletin, 2023, 39(5): 44-53.
图2 甘草酸的生物合成途径 IPP:异戊烯基焦磷酸,isopentenyl pyrophosphate;GPP:牻牛儿基焦磷酸,geranyl pyrophosphate;FPP:法尼基焦磷酸,farnesyl pyrophosphate;角鲨烯,Squalene;2,3-环氧角鲨烯,2,3-oxidosqualene; β-香树素,β-amyrin;11-氧代-β-香树脂醇,11-oxo-β-amyrin;甘草次酸,Glycyrrhetinic acid;3-O-单葡萄糖醛酸基甘草次酸,Glycyrrhetinic acid-3-O-monoglucoronide;甘草酸,Glycyrrhizin ;GPS:牻牛儿基焦磷酸合酶,geranylgeranyl diphosphate synthase;FPS:法尼基焦磷酸合酶,Farnesyl pyrophosphate synthase;SQS:角鲨烯合酶,Squalene synthase;SQE:角鲨烯环氧化酶,squalene epoxidase;β-AS:β-香树素合酶,β-amyrin synthase
Fig. 2 Glycyrrhizic acid biosynthetic pathway in licorice
图3 甘草苷的生物合成途径 Phenylalanine:苯丙氨酸;Cinnamic acid:肉桂酸;Coumaric acid:香豆酸;Coumaroyl-CoA:香豆酰辅酶A;Malonyl-CoA:丙二酰辅酶A;Naringenin chalcone:柚皮素查尔酮; Isoliquiritigenin:异甘草素;Liquiritigenin:甘草素;Liquiritin:甘草苷;PAL:苯丙氨酸解氨酶,phenylalanine ammonla-lyase;C4H:肉桂酸4-羟基化酶,cinnamate-4-hydroxylase;4CL:4-香豆酸辅酶A连接酶,4-coumarate-CoA ligase;CHS:查尔酮合酶,chalcone synthase;CHR:查尔酮还原酶,chalcone reductase
Fig. 3 Liquiritin biosynthetic pathway in licorice
类型Category | 影响因素Affecting factors | 对甘草酸含量影响Effect on glycyrrhizic acid content | 参考文献References |
---|---|---|---|
非生物胁迫 Abiotic stress | 低温胁迫 Low temperature stress 光照 Light | 低温处理提高甘草酸含量一倍 | [ [ |
红光,低强度和高强度UV-B处理都提高甘草酸苷 | |||
低光强增加甘草酸含量,100 μmol m-2 s-1的光照下甘草酸浓度最高 | [ | ||
干旱胁迫 Drought stress | 适度干旱提高甘草酸含量和总量,严重干旱提高甘草酸含量,降低甘草酸总量 | [ | |
10% PEG6000处理提高改良霍格兰营养液培养的甘草的甘草酸含量,10%和20% PEG6000处理降低1/2MS营养液培养的甘草的甘草酸含量 | [ | ||
盐胁迫 Salt stress | 盐胁迫(NaCl)诱导SQS、β-AS和CYP88D6的表达,提高甘草酸含量 | [ | |
复合盐胁迫(NaCl、CaCl2和MgSO4)抑制甘草酸积累 | [ | ||
营养胁迫 Nutritional stress | 高磷促进甘草酸积累 | [ | |
缺Mn处理降低甘草酸积累 土壤中总钾含量与甘草酸含量呈负相关 | [ [ | ||
生物因素 Biotic factors | 根瘤菌Rhizobacteria Mesorhizobium sp J8 | 接种J8后,甘草酸含量提高到3.2倍 | [ |
丛枝菌根真菌 Arbuscular mycorrhizal fungi Glomus mosseae | 接种Glomus mosseae甘草酸含量增加3-9倍 | [ | |
酵母Yeast Meyerozyma guilliermondii | 接种M. guilliermondii后,甘草酸含量比对照高5.3倍 | [ | |
酵母Yeast Pichia pastoris | 接种Pichia pastoris后甘草酸浓度提高3.89倍 | [ |
表1 不同因素对甘草酸含量影响
Table 1 Factors affecting the content of glycyrrhizic acid
类型Category | 影响因素Affecting factors | 对甘草酸含量影响Effect on glycyrrhizic acid content | 参考文献References |
---|---|---|---|
非生物胁迫 Abiotic stress | 低温胁迫 Low temperature stress 光照 Light | 低温处理提高甘草酸含量一倍 | [ [ |
红光,低强度和高强度UV-B处理都提高甘草酸苷 | |||
低光强增加甘草酸含量,100 μmol m-2 s-1的光照下甘草酸浓度最高 | [ | ||
干旱胁迫 Drought stress | 适度干旱提高甘草酸含量和总量,严重干旱提高甘草酸含量,降低甘草酸总量 | [ | |
10% PEG6000处理提高改良霍格兰营养液培养的甘草的甘草酸含量,10%和20% PEG6000处理降低1/2MS营养液培养的甘草的甘草酸含量 | [ | ||
盐胁迫 Salt stress | 盐胁迫(NaCl)诱导SQS、β-AS和CYP88D6的表达,提高甘草酸含量 | [ | |
复合盐胁迫(NaCl、CaCl2和MgSO4)抑制甘草酸积累 | [ | ||
营养胁迫 Nutritional stress | 高磷促进甘草酸积累 | [ | |
缺Mn处理降低甘草酸积累 土壤中总钾含量与甘草酸含量呈负相关 | [ [ | ||
生物因素 Biotic factors | 根瘤菌Rhizobacteria Mesorhizobium sp J8 | 接种J8后,甘草酸含量提高到3.2倍 | [ |
丛枝菌根真菌 Arbuscular mycorrhizal fungi Glomus mosseae | 接种Glomus mosseae甘草酸含量增加3-9倍 | [ | |
酵母Yeast Meyerozyma guilliermondii | 接种M. guilliermondii后,甘草酸含量比对照高5.3倍 | [ | |
酵母Yeast Pichia pastoris | 接种Pichia pastoris后甘草酸浓度提高3.89倍 | [ |
类型Category | 影响因素Factors | 对甘草苷含量影响Effect on liquiritin content | 参考文献References |
---|---|---|---|
非生物胁迫 Abiotic stress | 光照 Light | 低光强(100 μmol m-2 s-1)增加甘草苷含量 | [ |
干旱胁迫 Drought stress | 适度干旱提高甘草苷含量 | [ | |
盐胁迫 Salt stress | 盐处理上调PAL、C4H、4CL、CHS、GuUGT2和GuUGT3等甘草苷表达,促进甘草苷生成 | [ | |
营养胁迫 Nutritional stress | 高磷提高甘草苷浓度 | [ | |
缺Mn处理下显著降低甘草苷含量 | [ | ||
土壤中总钾含量与甘草苷浓度呈显著负相关 | [ | ||
生物因素 Biotic factor | 丛枝菌根真菌 Arbuscular mycorrhizal fungi Rhizophagus irregularis | 接种R. irregularis后促进CHS的表达,增加甘草苷浓度 | [ |
表2 不同因素对甘草苷含量影响
Table 2 Factors influencing the content of liquiritin
类型Category | 影响因素Factors | 对甘草苷含量影响Effect on liquiritin content | 参考文献References |
---|---|---|---|
非生物胁迫 Abiotic stress | 光照 Light | 低光强(100 μmol m-2 s-1)增加甘草苷含量 | [ |
干旱胁迫 Drought stress | 适度干旱提高甘草苷含量 | [ | |
盐胁迫 Salt stress | 盐处理上调PAL、C4H、4CL、CHS、GuUGT2和GuUGT3等甘草苷表达,促进甘草苷生成 | [ | |
营养胁迫 Nutritional stress | 高磷提高甘草苷浓度 | [ | |
缺Mn处理下显著降低甘草苷含量 | [ | ||
土壤中总钾含量与甘草苷浓度呈显著负相关 | [ | ||
生物因素 Biotic factor | 丛枝菌根真菌 Arbuscular mycorrhizal fungi Rhizophagus irregularis | 接种R. irregularis后促进CHS的表达,增加甘草苷浓度 | [ |
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