[1] 中国科学院中国植物志编辑委员会. 中国植物志: 第7卷[M]. 北京: 科学出版社, 1978: 99.

Editorial Committee of Flora of China, Chinese Academy of Science. Flora of China: Vol. 7. [M]. Beijing: Science Press, 1978: 99.
[2] 张宇航, 邱智东, 傅超美, 等. 刺五加化学成分药动学特性及其体内代谢过程研究进展[J]. 食品工业科技, 2020, 41(16): 334−339, 345.

ZHANG Yuhang, QIU Zhidong, FU Chaomei, et al. Research progress on the pharmacokinetics of Acanthopanax senticosus harms chemical constituents and its metabolism in vivo[J]. Science and Technology of Food Industry, 2020, 41(16): 334−339, 345. DOI: 10.13386/j.issn1002-0306.2020.16.053.
[3] 王知斌, 郭江涛, 姜海, 等. 刺五加叶的化学成分和药理作用研究[J]. 中医药信息, 2013, 30(3): 29−32.

WANG Zhibin, GUO Jiangtao, JIANG Hai, et al. Study on chemical constituents and pharmacological effects of Acanthopanax senticosus leaves[J]. Information on Traditional Chinese Medicine, 2013, 30(3): 29−32. DOI: 10.3969/j.issn.1002-2406.2013.03.012.
[4] 杨晓丹, 井月娥, 卢芳. 刺五加的化学成分研究进展[J]. 中华中医药学刊, 2015, 33(2): 316−318.

YANG Xiaodan, JING Yue’ e, LU Fang. Research Progress on chemical components of Acanthopanax senticosus[J]. Chinese Archives of Traditional Chinese Medicine, 2015, 33(2): 316−318. DOI: 10.13193/j.issn.1673-7717.2015.02.019.
[5] SHAO Chunjie, KASAI R, OHTANI K, et al. Saponins from leaves of Kalopanax septemlobus (Thunb. ) Koidz. : structures of Kalopanax-saponins La, Lb and Lc[J]. Chemical and Pharmaceutical Bulletin, 1989, 37(12): 3251−3254. DOI: 10.1248/cpb.37.3251.
[6] SHAO Chunjie, KASAI R, XU Jingda, et al. Saponins from leaves of Acanthopanax senticosus HARMS., Ciwujia: structures of Ciwujianosides B, C1, C2, C3, C4, D1, D2 and E [J]. Chemical & Pharmaceutical Bulletin, 1988, 36(2): 601-608. DOI:10.1248/cpb.36.601.
[7] LI Zhifeng, WU Zhaohua, CHEN Gang, et al. Two new compounds from Acanthopanax senticosus Harms.[J]. Journal of Asian Natural Products Research, 2009, 11(8): 716−719. DOI: 10.1080/10286020903020578.
[8] 朱立刚, 李志峰. 刺五加的化学成分研究[J]. 黑龙江中医药, 2013, 42(5): 65−66.

ZHU Ligang, LI Zhifeng. Studies on chemical constituents of Acanthopanax senticosus[J]. Heilongjiang Journal of Traditional Chinese Medicine, 2013, 42(5): 65−66.
[9] THIMMAPPA R, GEISLER K, LOUVEAU T, et al. Triterpene biosynthesis in plants[J]. Annual Review of Plant Biology, 2014, 65: 225−257. DOI: 10.1146/annurev-arplant-050312-120229.
[10] RAHIMI S, KIM J, MIJAKOVIC I, et al. Triterpenoid-biosynthetic UDP-glycosyltransferases from plants[J]. Biotechnology Advances, 2019, 37(7): 107394. DOI: 10.1016/j.biotechadv.2019.04.016.
[11] 徐圆圆, 陈仲, 贾黎明, 等. 植物三萜皂苷生物合成途径及调控机制研究进展[J]. 中国科学(生命科学), 2021, 51(5): 525−555.

XU Yuanyuan, CHEN Zhong, JIA Liming, et al. Advances in understanding of the biosynthetic pathway and regulatory mechanism of triterpenoid saponins in plants[J]. Science in China (Series C), 2021, 51(5): 525−555. DOI: 10.1360/SSV-2020-0230.
[12] JIA Ailing, ZHANG Yuhang, GAO Han, et al. A review of Acanthopanax senticosus (Rupr and Maxim.) Harms.: from ethnopharmacological use to modern application[J]. Journal of Ethnopharmacology, 2021, 268: 113586. DOI: 10.1016/j.jep.2020.113586.
[13] 李敏. 刺五加三萜皂苷合成途径关键基因挖掘及功能验证[D]. 哈尔滨: 东北林业大学, 2023.

LI Min. Mining and Functional Verification of Key Genes in the Synthesis Pathway of Eleutherococcus senticosus Triterpene Saponins[D]. Harbin: Northeast Forestry University, 2023.
[14] BAI Yuechen, FERNÁNDEZ-CALVO P, RITTER A, et al. Modulation of Arabidopsis root growth by specialized triterpenes[J]. The New Phytologist, 2021, 230(1): 228−243. DOI: 10.1111/nph.17144.
[15] HUANG Linzhang, ZHAO Hongfang, HUANG Baokang, et al. Acanthopanax senticosus: review of botany, chemistry and pharmacology[J]. Die Pharmazie, 2011, 66(2): 83−97.
[16] LUO Yun, LU Shan, GAO Ye, et al. Araloside C attenuates atherosclerosis by modulating macrophage polarization via Sirt1-mediated autophagy[J]. Aging, 2020, 12(2): 1704−1724. DOI: 10.18632/aging.102708.
[17] WANG Min, WANG Ruiying, ZHOU Jiahui, et al. Calenduloside E ameliorates myocardial ischemia-reperfusion injury through regulation of AMPK and mitochondrial OPA1[J]. Oxidative Medicine and Cellular Longevity, 2020, 2020: 2415269. DOI: 10.1155/2020/2415269.
[18] LI Fang, LI Wei, FU Hongwei, et al. Pancreatic lipase-inhibiting triterpenoid saponins from fruits of Acanthopanax senticosus[J]. Chemical and Pharmaceutical Bulletin, 2007, 55(7): 1087−1089. DOI: 10.1248/cpb.55.1087.
[19] GE Yuewei, TOHDA C, ZHU Shu, et al. Effects of oleanane-type triterpene saponins from the leaves of Eleutherococcus senticosus in an axonal outgrowth assay[J]. Journal of Natural Products, 2016, 79(7): 1834−1841. DOI: 10.1021/acs.jnatprod.6b00329.
[20] ZHAO Yujia, LI Chun. Biosynthesis of plant triterpenoid saponins in microbial cell factories[J]. Journal of Agricultural and Food Chemistry, 2018, 66(46): 12155−12165. DOI: 10.1021/acs.jafc.8b04657.
[21] VRANOVÁ E, COMAN D, GRUISSEM W. Network analysis of the MVA and MEP pathways for isoprenoid synthesis[J]. Annual Review of Plant Biology, 2013, 64: 665−700. DOI: 10.1146/annurev-arplant-050312-120116.
[22] 王琴波, 王雨晴, 杨谨, 等. 20种中药基原植物鲨烯合酶基因的生物信息学分析[J]. 中草药, 2021, 52(3): 838−844.

WANG Qinbo, WANG Yuqing, YANG Jin, et al. Bioinformatics analysis of squalene synthase gene from 20 Chinese herbal plants[J]. Chinese Traditional and Herbal Drugs, 2021, 52(3): 838−844. DOI: 10.7501/j.issn.0253-2670.2021.03.027.
[23] 李非非, 杨果, 吴鹏, 等. 刺五加HMGR基因的表达及其对皂苷合成的影响[J]. 基因组学与应用生物学, 2015, 34(3): 560−564.

LI Feifei, YANG Guo, WU Peng, et al. Expression of 3-hydroxy-3-methylglutaryl coenzyme a gene in Eleutherococcus senticosus and its influence on saponins synthesis[J]. Genomics and Applied Biology, 2015, 34(3): 560−564. DOI: 10.13417/j.gab.034.000560.
[24] ABBASSI S J, VISHWAKARMA R K, PATEL P, et al. Bacopa monniera recombinant mevalonate diphosphate decarboxylase: biochemical characterization[J]. International Journal of Biological Macromolecules, 2015, 79: 661−668. DOI:10.1016/j.ijbiomac.2015.05.041.
[25] 李志栋, 杨果, 尤鹏升, 等. 刺五加MDD基因启动子的克隆与生物信息学分析[J]. 中草药, 2016, 47(21): 3871−3875.

LI Zhidong, YANG Guo, YOU Pengsheng, et al. Cloning and bioinformatic analysis of mevalonate diphosphate decarboxylase gene promoter in Eleutherococcus senticosus[J]. Chinese Traditional and Herbal Drugs, 2016, 47(21): 3871−3875. DOI: 10.7501/j.issn.0253-2670.2016.21.020.
[26] 邢朝斌, 龙月红, 何闪, 等. 刺五加甲羟戊酸焦磷酸脱羧酶基因的克隆与表达分析[J]. 西北植物学报, 2012, 32(10): 1950−1956.

XING Zhaobin, LONG Yuehong, HE Shan, et al. Cloning and expression analysis of mevalonate diphosphate decarboxylasa gene in Eleutherococcus senticosus[J]. Acta Botanica Boreali-Occidentalia Sinica, 2012, 32(10): 1950−1956. DOI: 10.3969/j.issn.1000-4025.2012.10.003.
[27] 周秘, 柴丽花, 修乐山, 等. 刺五加法呢基焦磷酸合酶基因的表达及其与皂苷含量的相关性分析[J]. 河南农业科学, 2013, 42(12): 106−109.

ZHOU Mi, CHAI Lihua, XIU Leshan, et al. Expression level of Eleutherococcus senticosus farnesyl diphosphate dynthase gene and its correlation with saponin content[J]. Journal of Henan Agricultural Sciences, 2013, 42(12): 106−109. DOI: 10.3969/j.issn.1004-3268.2013.12.025.
[28] 邢朝斌, 龙月红, 李明, 等. 刺五加亚精胺合成酶基因的克隆及内生真菌对其表达的影响[J]. 中草药, 2012, 43(11): 2260−2265.

XING Zhaobin, LONG Yuehong, LI Ming, et al. Cloning of spermidine synthase gene in Eleutherococcus senticosus and effect of endophytic fungus on its expression[J]. Chinese Traditional and Herbal Drugs, 2012, 43(11): 2260−2265.
[29] 李敏, 王亢宗, 李艳林, 等. 刺五加β-香树脂醇合酶基因的挖掘及功能验证[J]. 世界科学技术-中医药现代化, 2022, 24(5): 1802−1810.

LI Min, WANG Kangzong, LI Yanlin, et al. Excavated gene and functional verification of β-amyrin synthase in Eleutherococcus senticosus[J]. World Science and Technology-Modernization of Traditional Chinese Medicine, 2022, 24(5): 1802−1810. DOI: 10.11842/wst.20211122004.
[30] 邢朝斌, 吴鹏, 龙月红, 等. 刺五加与其愈伤组织中皂苷合成关键酶基因表达的差异分析[J]. 中药材, 2012, 35(8): 1210−1213.

XING Zhaobin, WU Peng, LONG Yuehong, et al. Analyze on the expression differences of key genes in saponins biosynthesize in Eleutherococcus senticosus and its callus[J]. Journal of Chinese Medicinal Materials, 2012, 35(8): 1210−1213. DOI: 10.13863/j.issn1001-4454.2012.08.014.
[31] 龙月红, 李非非, 杨果, 等. 刺五加β-香树酯醇合成酶基因的克隆及其表达与皂苷量的相关性[J]. 中草药, 2015, 46(9): 1354−1359.

LONG Yuehong, LI Feifei, YANG Guo, et al. Gene cloning and expression level of bAS and their correlation with content of saponins in Eleutherococcus senticosus[J]. Chinese Traditional and Herbal Drugs, 2015, 46(9): 1354−1359. DOI: 10.7501/j.issn.0253-2670.2015.09.018.
[32] 刘镛. 植物三萜合成通路关键酶基因的适应性进化研究[D]. 南宁: 广西医科大学, 2013.

LIU Yong. Study on Adaptive Evolution of Key Enzyme Genes in Plant Triterpenoid Synthesis Nathway[D]. Nanning: Guangxi Medical University, 2013.
[33] 龙月红, 李非非, 杨果, 等. 刺五加鲨烯环氧酶基因家族2成员表达特性的分析[J]. 中国中药杂志, 2015, 40(1): 59−62.

LONG Yuehong, LI Feifei, YANG Guo, et al. Expression profiles analysis of two member of squaleneepoxidase gene family from Eleutherococcus senticosus[J]. China Journal of Chinese Materia Medica, 2015, 40(1): 59−62. DOI: 10.4268/cjcmm20150111.
[34] 龙月红, 尤鹏升, 国红玉, 等. 刺五加SS和SE拷贝数与皂苷含量的相关性分析[J]. 分子植物育种, 2017, 15(2): 725−729.

LONG Yuehong, YOU Pengsheng, GUO Hongyu, et al. The correlation analysis on SS and SE of copy number and saponins content in Eleutherococcus senticosus[J]. Molecular Plant Breeding, 2017, 15(2): 725−729. DOI: 10.13271/j.mpb.015.000725.
[35] 吴鹏, 修乐山, 李非非, 等. 刺五加P450基因原核表达体系的建立与表达条件的优化[J]. 中国中药杂志, 2015, 40(7): 1274−1277.

WU Peng, XIU Leshan, LI Feifei, et al. Establishment of prokaryotic expression and optimization of expression conditions of Eleutherococcus senticosus P450 gene[J]. China Journal of Chinese Materia Medica, 2015, 40(7): 1274−1277. DOI: 10.4268/cjcmm20150710.
[36] 邢朝斌, 吴鹏, 李非非, 等. 刺五加细胞色素P450基因的克隆与表达分析[J]. 生物技术通报, 2014, 30(1): 112−115.

XING Zhaobin, WU Peng, LI Feifei, et al. Cloning and expression analysis of cytochrome P450 gene in Eleutherococcus senticosus[J]. Biotechnology Bulletin, 2014, 30(1): 112−115. DOI: 10.13560/j.cnki.biotech.bull.1985.2014.01.024.
[37] 吴鹏, 谷俊涛, 修乐山, 等. 刺五加P450基因时空表达差异及与皂苷含量的相关性分析[J]. 河北农业大学学报, 2014, 37(3): 29−33.

WU Peng, GU Juntao, XIU Leshan, et al. Differential expression of Eleutherococcus senticosus P450 gene in time and space and the correlation analysis between expression level of E. senticosus P450 gene and saponins content[J]. Journal of Agricultural University of Hebei, 2014, 37(3): 29−33. DOI: 10.13320/j.cnki.jauh.2014.0059.
[38] 宋鑫, 崔明晖, 张朵朵, 等. 刺五加P450基因的筛选及其表达对皂苷含量的影响[J]. 中草药, 2021, 52(16): 5012−5020.

SONG Xin, CUI Minghui, ZHANG Duoduo, et al. Identification of P450 gene of Eleutherococcus senticosus and its effect on saponin content[J]. Chinese Traditional and Herbal Drugs, 2021, 52(16): 5012−5020. DOI: 10.7501/j.issn.0253-2670.2021.16.026.
[39] 朱灵英, 郭娟, 张爱丽, 等. 参与植物三萜生物合成的细胞色素P450酶研究进展[J]. 中草药, 2019, 50(22): 5597−5610.

ZHU Lingying, GUO Juan, ZHANG Aili, et al. Research progress on CYP450 involved in medicinal plant triterpenoid biosynthesis[J]. Chinese Traditional and Herbal Drugs, 2019, 50(22): 5597−5610. DOI: 10.7501/j.issn.0253-2670.2019.22.031.
[40] 修乐山, 柴丽花, 周秘, 等. 刺五加GAPDH基因全长cDNA的克隆与生物信息学分析[J]. 广东农业科学, 2013, 40(16): 124−127.

XIU Leshan, CHAI Lihua, ZHOU Mi, et al. Cloning and bioinformatics analysis of cDNA of GAPDH gene from Eleutherococcus senticosus[J]. Guangdong Agricultural Sciences, 2013, 40(16): 124−127. DOI: 10.3969/j.issn.1004-874X.2013.16.040.
[41] 修乐山, 李非非, 周秘, 等. 刺五加糖基转移酶基因的表达及其对皂苷含量的影响[J]. 基因组学与应用生物学, 2014, 33(1): 128−132.

XIU Leshan, LI Feifei, ZHOU Mi, et al. Expression of glycosyltransferase gene in Eleutherococcus senticosus and its influence on saponins content[J]. Genomics and Applied Biology, 2014, 33(1): 128−132. DOI: 10.13417/j.gab.033.000128.
[42] 邢朝斌, 修乐山, 吴鹏, 等. 刺五加葡萄糖基转移酶基因的克隆与表达分析[J]. 西北农业学报, 2014, 23(2): 126−130.

XING Zhaobin, XIU Leshan, WU Peng, et al. Cloning and expression analysis of glucosyltransferase gene in Eleutherococcus senticosus[J]. Acta Agriculturae Boreali-Occidentalis Sinica, 2014, 23(2): 126−130. DOI: 10.7606/j.issn.1004-1384.2014.02.022.
[43] YAO Lu, LU Jun, WANG Juan, et al. Advances in biosynthesis of triterpenoid saponins in medicinal plants[J]. Chinese Journal of Natural Medicines, 2020, 18(6): 417−424. DOI: 10.1016/S1875-5364(20)30049-2.
[44] DENG Bing, HUANG Zhuangjia, GE Feng, et al. An AP2/ERF family transcription factor PnERF1 raised the biosynthesis of saponins in Panax notoginseng[J]. Journal of Plant Growth Regulation, 2017, 36(3): 691−701. DOI: 10.1007/s00344-017-9672-z.
[45] 龙月红, 宋菊, 国红玉, 等. 刺五加转录因子基因的挖掘[J]. 北方园艺, 2017(20): 153−156.

LONG Yuehong, SONG Ju, GUO Hongyu, et al. Exploring transcription factors in Eleutherococcus senticosus[J]. Northern Horticulture, 2017(20): 153−156. DOI: 10.11937/bfyy.20170195.
[46] 董靖, 赵雪蕾, 王硕, 等. 刺五加NAC转录因子基因的适应性进化分析[J]. 林业科技通讯, 2024(5): 33−38.

DONG Jing, ZHAO Xuelei, WANG Shuo, et al. Adaptive evolutionary analysis of the NAC transcription factor genes of Eleutherococcus senticosus[J]. Forest Science and Technology, 2024(5): 33−38. DOI: 10.13456/j.cnki.lykt.2023.09.01.0003.
[47] 董靖. 刺五加NAC的鉴定及其在DNA甲基化调控皂苷合成中的作用[D]. 唐山: 华北理工大学, 2024.

DONG Jing. Identification of Eleutherococcus senticosus NAC and Its Role in DNA Methylation Regulating Saponin Synthesis[D]. Tangshan: North China University of Science and Technology, 2024.
[48] LI Chang, MA Jiacheng, WU Peng, et al. Identification and adaptive evolution analysis of FAR1/FHY3 in Eleutherococcus senticosus and prediction of target genes regulating saponin synthesis[J]. Medicinal Plant Biology, 2025, 4: e16. DOI: 10.48130/mpb-0025-0013.
[49] 赵雪蕾. 刺五加DNA甲基化相关酶基因家族的鉴定及其功能分析[D]. 唐山: 华北理工大学, 2024.

ZHAO Xuelei. Identification and Functional Analysis of DNA Methylation-Related Enzyme Gene Family in Eleutherococcus senticosus[D]. Tangshan: North China University of Science and Technology, 2024.
[50] 王硕. 干旱胁迫调控刺五加DNA甲基化和皂苷合成的机制研究[D]. 唐山: 华北理工大学, 2024.

WANG Shuo. Study on the Mechanism of Drought Stress Regulating DNA Methylation and Saponin Synthesis of Eleutherococcus senticosus[D]. Tangshan: North China University of Science and Technology, 2024.
[51] 崔明晖. 刺五加MDD基因启动子DNA甲基化对皂苷合成的影响机制[D]. 唐山: 华北理工大学, 2022.

CUI Minghui. Mechanism of DNA Methylation of MDD Gene Promoter on Saponin Synthesis in Eleutherococcus senticosus[D]. Tangshan: North China University of Science and Technology, 2022.
[52] CHUNG S Y, SEKI H, FUJISAWA Y, et al. A cellulose synthase-derived enzyme catalyses 3-O-glucuronosylation in saponin biosynthesis[J]. Nature Communications, 2020, 11: 5664. DOI: 10.1038/s41467-020-19399-0.
[53] YAN Xing, FAN Yun, WEI Wei, et al. Production of bioactive ginsenoside compound K in metabolically engineered yeast[J]. Cell Research, 2014, 24(6): 770−773. DOI: 10.1038/cr.2014.28.