Article(id=1304388171224080825, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388135723496407, articleNumber=null, orderNo=null, doi=10.7501/j.issn.0253-2670.2026.13.023, pmid=null, cstr=null, oa=null, hot=0, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1767283200000, receivedDateStr=2026-01-02, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1788919974380, onlineDateStr=2026-09-09, pubDate=null, pubDateStr=null, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1788919974380, onlineIssueDateStr=2026-09-09, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1788919974380, creator=13701087609, updateTime=1788919974380, updator=13701087609, issue=Issue{id=1304388135723496407, tenantId=1146029695717560320, journalId=1302319053441957962, year='2026', volume='57', issue='13', pageStart='4949', pageEnd='5352', issueExtLink='null', onlineDate='null', pubDate='1783785600000', pubDateStr='2026-07-12', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1788919965916, creator='13701087609', updateTime=1788923489765, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1304402915871977875, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388135723496407, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1304402915871977876, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388135723496407, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=5215, endPage=5227, ext={EN=ArticleExt(id=1304388171609956795, articleId=1304388171224080825, tenantId=1146029695717560320, journalId=1302319053441957962, language=EN, title=Identification of CAS gene family in Panax ginseng and functional analysis of PgCAS04-5, columnId=null, journalTitle=Chinese Traditional and Herbal Drugs, columnName=null, runingTitle=null, highlight=null, articleAbstract=Objective To identify members of the cycloartenol synthase (CAS) gene family (PgCAS) using the Panax ginseng transcriptome database, and analyze the expression patterns of PgCAS04-5 in overexpressed and RNAi-positive hair root materials), as well as explore the function of this gene in ginsenoside biosynthesis. Methods Bioinformatics approaches were adopted to characterize the chromosomal localization, conserved motifs, phylogenetic relationships and expression profiles, and the correlations with the gene expression of key saponin synthesis enzymes and the content of monomer saponin of PgCASs. The protein structure and subcellular localization of PgCAS04-5 were predicted. Meanwhile, the expression levels of key enzyme genes and variations in ginsenoside contents were detected in overexpression and RNAi-positive hairy root lines. Results A total of 20 PgCAS transcripts were identified from the P. ginseng transcriptome database. Chromosomal mapping showed that PgCAS genes were unevenly distributed on five chromosomes of P. ginseng. Phylogenetic tree analysis divided the PgCAS family into three subfamilies, and most PgCAS members belonged to subfamily Ⅱ and subfamily Ⅲ. The PgCAS genes displayed obvious spatiotemporal specificity in expression across different cultivars, growth years and tissues. Additionally, the PgCAS family formed an interaction network with key genes involved in ginsenoside biosynthesis, and their content was significantly correlated with the accumulation of monomeric ginsenosides. Subcellular localization prediction demonstrated that PgCAS04-5 was localized to chloroplasts. Moreover, the overexpression of this gene was inversely regulated with the expression of four genes including PgCAS04-5, PgDS-1, PgSE2-4 and PgCYP137 in RNAi positive materials, with significant differences. Conclusion A total of 20 members of the PgCAS gene family were identified in this study, and functional differentiation exists among PgCAS family members. PgCAS04-5 positively regulates the biosynthesis of ginsenosides Rg2and Rf, and negatively modulates the synthesis of ginsenosides Rb2and Rd. The results provide a valuable reference for further research on the PgCAS gene fami, authors=MA Ying, LI Chengwei, JIN Ge, YU Shuang, WANG Aimin, YU Jinghui, ZHAO Mingzhu, WANG Kangyu, WANG Yi, ZHANG Meiping, authorsList=MA Ying, LI Chengwei, JIN Ge, YU Shuang, WANG Aimin, YU Jinghui, ZHAO Mingzhu, WANG Kangyu, WANG Yi, ZHANG Meiping, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, 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=1304388171517682106, articleId=1304388171224080825, tenantId=1146029695717560320, journalId=1302319053441957962, language=CN, title=人参CAS基因家族的鉴定及PgCAS04-5基因的功能分析, columnId=1304140203967410414, journalTitle=中草药, columnName=药材与资源, runingTitle=null, highlight=null, articleAbstract=目的 基于人参转录组数据库鉴定环阿屯醇合酶基因(cycloartenol synthase,CAS)家族成员(PgCAS),并分析PgCAS04-5 在过表达及RNAi阳性发根材料中基因的表达模式及人参皂苷生物合成中的功能。方法 利用生物信息学分析了PgCASs的染色体定位、保守基序、进化关系、表达模式、与皂苷合成关键酶基因表达及单体皂含量的相关性;预测PgCAS04-5的蛋白结构与亚细胞定位,同时检测PgCAS04-5在过表达及RNAi阳性发根中关键酶基因的表达情况及皂苷含量变化。结果 在人参转录组数据库中鉴定出20条PgCAS转录本,染色体定位结果表明PgCASs不均匀地分布在人参5条染色体上,构建进化树将PgCASs分为3个亚型,PgCASs主要分布在第Ⅱ和第Ⅲ个亚型中。PgCASs在不同品种、生长年限及组织中的表达模式存在时空特异性,且该家族与皂苷合成关键酶基因形成互作网络并与单体皂苷含量显著相关。PgCAS04-5的亚细胞定位为叶绿体,同时该基因的过表达与RNAi阳性材料中PgCAS04-5、PgDS-1、PgSE2-4、PgCYP137 这4个基因的表达呈反向调控,差异显著。结论 鉴定出20个PgCAS家族成员,PgCAS家族成员功能存在分化,PgCAS04-5基因正向调控人参皂苷Rg₂和人参皂苷Rf的合成,同时负向调控人参皂苷Rb₂和人参皂苷Rd的生物合成,为进一步研究PgCAS基因家族提供参考。, authors=马莹1,2, 李宸玮1,2, 金戈1,2, 于双1,2, 王爱民1,2, 于靖辉1,2, 赵明珠1,2, 王康宇1,2, 王义1,2, 张美萍1,2, authorsList=马莹, 李宸玮, 金戈, 于双, 王爱民, 于靖辉, 赵明珠, 王康宇, 王义, 张美萍, authorCompany=1 吉林农业大学生命科学学院, 吉林 长春 130118; 2 吉林农业大学, 吉林省人参基因资源开发与利用工程研究中心, 吉林 长春 130118, correspAuthors=王义, authorNote=马莹: 马莹,女,硕士研究生,研究方向为生物工程。E-mail:1076900427@qq.com, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=56jfVfN0/cSj3GhkGeolHA==, pdfFileSize=2128007, 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=人参基因育种技术的建立及优良农家品种选育 (20250205032GH))}, authors=null, keywords=[Keyword(id=1304401956886307394, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=CN, orderNo=1, keyword=人参), Keyword(id=1304401956965999171, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=CN, orderNo=2, keyword=环阿屯醇合酶), Keyword(id=1304401957045690948, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=CN, orderNo=3, keyword=基因家族), Keyword(id=1304401957125382725, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=CN, orderNo=4, keyword=生物信息学分析), Keyword(id=1304401957200880198, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=CN, orderNo=5, keyword=功能分析), Keyword(id=1304401957364458055, tenantId=1146029695717560320, journalId=1302319053441957962, articleId=1304388171224080825, language=EN, orderNo=1, keyword=Panax ginseng C. 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Current evaluation of the millennium phytomedicine- ginseng (I): Etymology, pharmacognosy, phytochemistry, market and regulations [J]. Curr Med Chem, 2009, 16(19): 2475-2484. 赵春丽, 李成国, 夏昉, 等. 中国人参产业发展现状及对策 [J]. 应用化学, 2025, 42(9): 1282-1288. Zhang H, Abid S, Ahn J C, et al. Characteristics of Panax ginseng cultivars in Korea and China [J]. Molecules, 2020, 25(11): 2635. Du Y L, Fu X Z, Chu Y Y, et al. Biosynthesis and the roles of plant sterols in development and stress responses [J]. Int J Mol Sci, 2022, 23(4): 2332. 尹艳, 关红雨, 张夏楠. 甾体皂苷生物合成相关酶及基因研究进展 [J]. 天然产物研究与开发, 2016, 28(8): 1332-1336. Hou M Q, Wang R F, Zhao S J, et al. Ginsenosides in Panax genus and their biosynthesis [J]. Acta Pharm Sin B, 2021, 11: 1813-1834. Guo C F, Xu S, Guo X Y. Genome-wide analysis of oxidosqualene cyclase genes in Artemisia annua: Evolution, expression, and potential roles in triterpenoid biosynthesis [J]. Curr Issues Mol Biol, 2025, 47(7): 545. Gas-Pascual E, Berna A, Bach T J, et al. Plant oxidosqualene metabolism: Cycloartenol synthase-dependent sterol biosynthesis in Nicotiana benthamiana [J]. PLoS One, 2014, 9(10): e109156. Parveen I, Wang M, Lee J, et al. Identification and functional characterization of oxidosqualene cyclases from medicinal plant Hoodia gordonii [J]. Plants, 2024, 13(2): 231. Hakim S E, Liu S Y, Herzog R, et al. Expansion of the stereochemical space of triterpenes by mining noncanonical oxidosqualene cyclases across the diversity of green plants [J]. J Am Chem Soc, 2025, 147(12): 10320-10330. Wang K Y, Jiang S C, Sun C Y, et al. The spatial and temporal transcriptomic landscapes of ginseng, Panax ginseng C. A. Meyer [J]. Sci Rep, 2016, 5: 18283. Liu Z H, Suarez Duran H G, Harnvanichvech Y, et al. Drivers of metabolic diversification: How dynamic genomic neighbourhoods generate new biosynthetic pathways in the Brassicaceae [J]. New Phytol, 2020, 227(4): 1109-1123. 焉雅涛. 吉林人参OSC基因家族系统演化、功能分化及表达活动分析 [D]. 长春: 吉林农业大学, 2015. Kushiro T, Shibuya M, Ebizuka Y. β-Amyrin synthase: Cloning of oxidosqualene cyclase that catalyzes the formation of the most popular triterpene among higher plants [J]. Eur J Biochem, 1998, 256(1): 238-244. Corey E J, Matsuda S P, Bartel B. Isolation of an Arabidopsis thaliana gene encoding cycloartenol synthase by functional expression in a yeast mutant lacking lanosterol synthase by the use of a chromatographic screen [J]. Proc Natl Acad Sci USA, 1993, 90(24): 11628-11632. Zhang H, Shibuya M, Yokota S, et al. Oxidosqualene cyclases from cell suspension cultures of Betula platyphylla var. japonica: Molecular evolution of oxidosqualene cyclases in higher plants [J]. Biol Pharm Bull, 2003, 26(5): 642-650. Hayashi H, Hiraoka N, Ikeshiro Y, et al. Molecular cloning and characterization of a cDNA for Glycyrrhiza glabra cycloartenol synthase [J]. Biol Pharm Bull, 2000, 23(2): 231-234. Guhling O, Hobl B, Yeats T, et al. Cloning and characterization of a lupeol synthase involved in the synthesis of epicuticular wax crystals on stem and hypocotyl surfaces of Ricinus communis [J]. Arch Biochem Biophys, 2006, 448(1/2): 60-72. 邢朝斌, 龙月红, 吴鹏, 等. 刺五加环阿屯醇合酶基因的克隆及其表达分析 [J]. 中草药, 2012, 43(7): 1387-1392. 郑蔚, 代佳妮, 严和琴, 等. 越南油茶HMGCR基因的克隆及表达分析 [J]. 分子植物育种, 2022, 20(13): 4296-4304. 刘梦迪, 李长福, 章焰生. 葫芦巴环阿屯醇合酶基因的分离及其对薯蓣皂素合成的影响 [J]. 植物科学学报, 2019, 37(1): 87-92. Zhao H, Tang Q, Mo C, et al. Cloning and characterization of squalene synthase and cycloartenol synthase from Siraitia grosvenorii [J]. Acta Pharmaceutica Sinia B, 2017, 7(2): 215-222. Duan Y Y, Du W Y, Song Z J, et al. Functional characterization of a cycloartenol synthase and four glycosyltransferases in the biosynthesis of cycloastragenol-type astragalosides from Astragalus membranaceus [J]. Acta Pharm Sin B, 2023, 13(1): 271-283. Lin X Y, Kaul S, Rounsley S, et al. Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana [J]. Nature, 1999, 402(6763): 761-768. Atsumi G, Kagaya U, Tabayashi N, et al. Analysis of the mechanisms regulating the expression of isoprenoid biosynthesis genes in hydroponically-grown Nicotiana benthamiana plants using virus-induced gene silencing [J]. Sci Rep, 2018, 8: 14804. Gas-Pascual E, Berna A, Bach T J, et al. Plant oxidosqualene metabolism: Cycloartenol synthase-dependent sterol biosynthesis in Nicotiana benthamiana [J]. PLoS One, 2014, 9(10): e109156. Schnable P S, Ware D, Fulton R S, et al. The B73 maize genome: Complexity, diversity, and dynamics [J]. Science, 2009, 326(5956): 1112-1115. 于靖辉. 人参内生菌广泛靶向代谢组学分析及其对人参不定根的影响 [D]. 长春: 吉林农业大学, 2023 Yu J, Wang D, Zhao T, et al. Potential participants and regulatory factors in ginsenoside biosynthesis of Panax ginseng C.A. Meyer: The role of endophytic fungus PBF-08[J]. Plant Science, 2025, 358: 112553. Jiang Z Q, Tu L C, Yang W F, et al. The chromosome-level reference genome assembly for Panax notoginseng and insights into ginsenoside biosynthesis [J]. Plant Commun, 2021, 2(1): 100113. Van de Peer Y, Mizrachi E, Marchal K. The evolutionary significance of polyploidy [J]. Nat Rev Genet, 2017, 18(7): 411-424. Heidari P, Puresmaeli F, Mora-Poblete F. Genome-wide identification and molecular evolution of the magnesium transporter (MGT) gene family in Citrullus lanatus and Cucumissativus [J]. Agronomy, 2022, 12(10): 2253. Faraji S, Heidari P, Amouei H, et al. Investigation and computational analysis of the sulfotransferase (SOT) gene family in potato (Solanum tuberosum): Insights into sulfur adjustment for proper development and stimuli responses [J]. Plants, 2021, 10(12): 2597. Almeida de Jesus D, Batista D M, Monteiro E F, et al. Structural changes and adaptative evolutionary constraints in flowering locus t and TERMINAL FLOWER1-like genes of flowering plants [J]. Front Genet, 2022, 13: 954015. 赵昕, 高雅, 胡英考, 等. 水稻CAS基因的克隆及分析 [J]. 安徽农业科学, 2009, 37(28): 13488-13489. Milla P, Viola F, Bosso S O, et al. Subcellular localization of oxidosqualene cyclases from Arabidopsis thaliana, Trypanosoma cruzi, and Pneumocystis carinii expressed in yeast [J]. Lipids, 2002, 37(12): 1171-1176. 许勇, 李静, 蔡标, 等. 贝母中环阿屯醇合成酶的生物信息学分析 [J]. 生物技术通讯, 2018, 29(4): 481-485. Rodriguez-Concepcion M, Lim S, Ha S H. Family alliances feeding the carotenoid pathway in tomato [J]. J Exp Bot, 2025, 76(21): 6327-6334. 李月, 庞亚如, 成旭, 等. 酿酒酵母中胆固醇生物合成与优化的研究进展 [J]. 微生物学通报, 2022, 49(11): 4869-4885.)
Objective To identify members of the cycloartenol synthase (CAS) gene family (PgCAS) using the Panax ginseng transcriptome database, and analyze the expression patterns of PgCAS04-5 in overexpressed and RNAi-positive hair root materials), as well as explore the function of this gene in ginsenoside biosynthesis. Methods Bioinformatics approaches were adopted to characterize the chromosomal localization, conserved motifs, phylogenetic relationships and expression profiles, and the correlations with the gene expression of key saponin synthesis enzymes and the content of monomer saponin of PgCASs. The protein structure and subcellular localization of PgCAS04-5 were predicted. Meanwhile, the expression levels of key enzyme genes and variations in ginsenoside contents were detected in overexpression and RNAi-positive hairy root lines. Results A total of 20 PgCAS transcripts were identified from the P. ginseng transcriptome database. Chromosomal mapping showed that PgCAS genes were unevenly distributed on five chromosomes of P. ginseng. Phylogenetic tree analysis divided the PgCAS family into three subfamilies, and most PgCAS members belonged to subfamily Ⅱ and subfamily Ⅲ. The PgCAS genes displayed obvious spatiotemporal specificity in expression across different cultivars, growth years and tissues. Additionally, the PgCAS family formed an interaction network with key genes involved in ginsenoside biosynthesis, and their content was significantly correlated with the accumulation of monomeric ginsenosides. Subcellular localization prediction demonstrated that PgCAS04-5 was localized to chloroplasts. Moreover, the overexpression of this gene was inversely regulated with the expression of four genes including PgCAS04-5, PgDS-1, PgSE2-4 and PgCYP137 in RNAi positive materials, with significant differences. Conclusion A total of 20 members of the PgCAS gene family were identified in this study, and functional differentiation exists among PgCAS family members. PgCAS04-5 positively regulates the biosynthesis of ginsenosides Rg2and Rf, and negatively modulates the synthesis of ginsenosides Rb2and Rd. The results provide a valuable reference for further research on the PgCAS gene fami
Key words
Panax ginseng C. A. Mey
/
cycloartenol synthase
/
gene family
/
bioinformatics analysis
/
functional analysis
MA Ying, LI Chengwei, JIN Ge, YU Shuang, WANG Aimin, YU Jinghui, ZHAO Mingzhu, WANG Kangyu, WANG Yi, ZHANG Meiping.
Identification of CAS gene family in Panax ginseng and functional analysis of PgCAS04-5[J].
Chinese Traditional and Herbal Drugs,
2026
, 57
(13)
: 5215
-5227
.
DOI: 10.7501/j.issn.0253-2670.2026.13.023
基金
收起
人参基因育种技术的建立及优良农家品种选育 (20250205032GH)
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Jia L, Zhao Y Q. Current evaluation of the millennium phytomedicine- ginseng (I): Etymology, pharmacognosy, phytochemistry, market and regulations [J]. Curr Med Chem, 2009, 16(19): 2475-2484. 赵春丽, 李成国, 夏昉, 等. 中国人参产业发展现状及对策 [J]. 应用化学, 2025, 42(9): 1282-1288. Zhang H, Abid S, Ahn J C, et al. Characteristics of Panax ginseng cultivars in Korea and China [J]. Molecules, 2020, 25(11): 2635. Du Y L, Fu X Z, Chu Y Y, et al. Biosynthesis and the roles of plant sterols in development and stress responses [J]. Int J Mol Sci, 2022, 23(4): 2332. 尹艳, 关红雨, 张夏楠. 甾体皂苷生物合成相关酶及基因研究进展 [J]. 天然产物研究与开发, 2016, 28(8): 1332-1336. Hou M Q, Wang R F, Zhao S J, et al. Ginsenosides in Panax genus and their biosynthesis [J]. Acta Pharm Sin B, 2021, 11: 1813-1834. Guo C F, Xu S, Guo X Y. Genome-wide analysis of oxidosqualene cyclase genes in Artemisia annua: Evolution, expression, and potential roles in triterpenoid biosynthesis [J]. Curr Issues Mol Biol, 2025, 47(7): 545. Gas-Pascual E, Berna A, Bach T J, et al. Plant oxidosqualene metabolism: Cycloartenol synthase-dependent sterol biosynthesis in Nicotiana benthamiana [J]. PLoS One, 2014, 9(10): e109156. Parveen I, Wang M, Lee J, et al. Identification and functional characterization of oxidosqualene cyclases from medicinal plant Hoodia gordonii [J]. Plants, 2024, 13(2): 231. Hakim S E, Liu S Y, Herzog R, et al. Expansion of the stereochemical space of triterpenes by mining noncanonical oxidosqualene cyclases across the diversity of green plants [J]. J Am Chem Soc, 2025, 147(12): 10320-10330. Wang K Y, Jiang S C, Sun C Y, et al. The spatial and temporal transcriptomic landscapes of ginseng, Panax ginseng C. A. Meyer [J]. Sci Rep, 2016, 5: 18283. Liu Z H, Suarez Duran H G, Harnvanichvech Y, et al. Drivers of metabolic diversification: How dynamic genomic neighbourhoods generate new biosynthetic pathways in the Brassicaceae [J]. New Phytol, 2020, 227(4): 1109-1123. 焉雅涛. 吉林人参OSC基因家族系统演化、功能分化及表达活动分析 [D]. 长春: 吉林农业大学, 2015. Kushiro T, Shibuya M, Ebizuka Y. β-Amyrin synthase: Cloning of oxidosqualene cyclase that catalyzes the formation of the most popular triterpene among higher plants [J]. Eur J Biochem, 1998, 256(1): 238-244. Corey E J, Matsuda S P, Bartel B. Isolation of an Arabidopsis thaliana gene encoding cycloartenol synthase by functional expression in a yeast mutant lacking lanosterol synthase by the use of a chromatographic screen [J]. Proc Natl Acad Sci USA, 1993, 90(24): 11628-11632. Zhang H, Shibuya M, Yokota S, et al. Oxidosqualene cyclases from cell suspension cultures of Betula platyphylla var. japonica: Molecular evolution of oxidosqualene cyclases in higher plants [J]. Biol Pharm Bull, 2003, 26(5): 642-650. Hayashi H, Hiraoka N, Ikeshiro Y, et al. Molecular cloning and characterization of a cDNA for Glycyrrhiza glabra cycloartenol synthase [J]. Biol Pharm Bull, 2000, 23(2): 231-234. Guhling O, Hobl B, Yeats T, et al. Cloning and characterization of a lupeol synthase involved in the synthesis of epicuticular wax crystals on stem and hypocotyl surfaces of Ricinus communis [J]. Arch Biochem Biophys, 2006, 448(1/2): 60-72. 邢朝斌, 龙月红, 吴鹏, 等. 刺五加环阿屯醇合酶基因的克隆及其表达分析 [J]. 中草药, 2012, 43(7): 1387-1392. 郑蔚, 代佳妮, 严和琴, 等. 越南油茶HMGCR基因的克隆及表达分析 [J]. 分子植物育种, 2022, 20(13): 4296-4304. 刘梦迪, 李长福, 章焰生. 葫芦巴环阿屯醇合酶基因的分离及其对薯蓣皂素合成的影响 [J]. 植物科学学报, 2019, 37(1): 87-92. Zhao H, Tang Q, Mo C, et al. Cloning and characterization of squalene synthase and cycloartenol synthase from Siraitia grosvenorii [J]. Acta Pharmaceutica Sinia B, 2017, 7(2): 215-222. Duan Y Y, Du W Y, Song Z J, et al. Functional characterization of a cycloartenol synthase and four glycosyltransferases in the biosynthesis of cycloastragenol-type astragalosides from Astragalus membranaceus [J]. Acta Pharm Sin B, 2023, 13(1): 271-283. Lin X Y, Kaul S, Rounsley S, et al. Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana [J]. Nature, 1999, 402(6763): 761-768. Atsumi G, Kagaya U, Tabayashi N, et al. Analysis of the mechanisms regulating the expression of isoprenoid biosynthesis genes in hydroponically-grown Nicotiana benthamiana plants using virus-induced gene silencing [J]. Sci Rep, 2018, 8: 14804. Gas-Pascual E, Berna A, Bach T J, et al. Plant oxidosqualene metabolism: Cycloartenol synthase-dependent sterol biosynthesis in Nicotiana benthamiana [J]. PLoS One, 2014, 9(10): e109156. Schnable P S, Ware D, Fulton R S, et al. The B73 maize genome: Complexity, diversity, and dynamics [J]. Science, 2009, 326(5956): 1112-1115. 于靖辉. 人参内生菌广泛靶向代谢组学分析及其对人参不定根的影响 [D]. 长春: 吉林农业大学, 2023 Yu J, Wang D, Zhao T, et al. Potential participants and regulatory factors in ginsenoside biosynthesis of Panax ginseng C.A. Meyer: The role of endophytic fungus PBF-08[J]. Plant Science, 2025, 358: 112553. Jiang Z Q, Tu L C, Yang W F, et al. The chromosome-level reference genome assembly for Panax notoginseng and insights into ginsenoside biosynthesis [J]. Plant Commun, 2021, 2(1): 100113. Van de Peer Y, Mizrachi E, Marchal K. The evolutionary significance of polyploidy [J]. Nat Rev Genet, 2017, 18(7): 411-424. Heidari P, Puresmaeli F, Mora-Poblete F. Genome-wide identification and molecular evolution of the magnesium transporter (MGT) gene family in Citrullus lanatus and Cucumissativus [J]. Agronomy, 2022, 12(10): 2253. Faraji S, Heidari P, Amouei H, et al. Investigation and computational analysis of the sulfotransferase (SOT) gene family in potato (Solanum tuberosum): Insights into sulfur adjustment for proper development and stimuli responses [J]. Plants, 2021, 10(12): 2597. Almeida de Jesus D, Batista D M, Monteiro E F, et al. Structural changes and adaptative evolutionary constraints in flowering locus t and TERMINAL FLOWER1-like genes of flowering plants [J]. Front Genet, 2022, 13: 954015. 赵昕, 高雅, 胡英考, 等. 水稻CAS基因的克隆及分析 [J]. 安徽农业科学, 2009, 37(28): 13488-13489. Milla P, Viola F, Bosso S O, et al. Subcellular localization of oxidosqualene cyclases from Arabidopsis thaliana, Trypanosoma cruzi, and Pneumocystis carinii expressed in yeast [J]. Lipids, 2002, 37(12): 1171-1176. 许勇, 李静, 蔡标, 等. 贝母中环阿屯醇合成酶的生物信息学分析 [J]. 生物技术通讯, 2018, 29(4): 481-485. Rodriguez-Concepcion M, Lim S, Ha S H. Family alliances feeding the carotenoid pathway in tomato [J]. J Exp Bot, 2025, 76(21): 6327-6334. 李月, 庞亚如, 成旭, 等. 酿酒酵母中胆固醇生物合成与优化的研究进展 [J]. 微生物学通报, 2022, 49(11): 4869-4885.