Article(id=1320880573862753109, tenantId=1146029695717560320, journalId=1260987677001138203, issueId=1320880262012056083, articleNumber=null, orderNo=null, doi=10.13386/j.issn1002-0306.2025060101, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1749571200000, receivedDateStr=2025-06-11, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1792852069583, onlineDateStr=2026-10-24, pubDate=1789401600000, pubDateStr=2026-09-15, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1792852069583, onlineIssueDateStr=2026-10-24, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1792852069583, creator=13701087609, updateTime=1792852069583, updator=13701087609, issue=Issue{id=1320880262012056083, tenantId=1146029695717560320, journalId=1260987677001138203, year='2026', volume='47', issue='18', pageStart='1', pageEnd='613', issueExtLink='null', onlineDate='null', pubDate='1789401600000', pubDateStr='2026-09-15', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1792851995232, creator='13701087609', updateTime=1792921655980, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1321172440479290195, tenantId=1146029695717560320, journalId=1260987677001138203, issueId=1320880262012056083, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1321172440479290196, tenantId=1146029695717560320, journalId=1260987677001138203, issueId=1320880262012056083, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=284, endPage=294, ext={EN=ArticleExt(id=1320880574059885398, articleId=1320880573862753109, tenantId=1146029695717560320, journalId=1260987677001138203, language=EN, title=Revealing Summer Leaf Purpling Phenotype of Camellia sinensis cv. 'Longrui 1' with Untargeted Metabolomics, columnId=1320880262846722580, journalTitle=Science and Technology of Food Industry, columnName=Modern Analysis and Intelligent Detection, runingTitle=null, highlight=null, articleAbstract=

To elucidate the molecular mechanism underlying the seasonal variation in leaf color of the unique tea plan germplasm 'Longrui 1' (Camellia sinensis cv. 'Longrui 1', Lr1), the dynamic patterns and underlying mechanisms of the seasonal purple phenotype in 'Longrui 1', particularly regarding leaf coloration, quality components, and metabolic profiles, were systematically investigated. A combined approach of biochemical assays, untargeted and targeted metabolomics was employed, using the constitutively green cultivar 'Zhongcha 108' and the constitutively purple cultivar 'Zijuan' as controls. The results indicated that the young shoots of Lr1 transitioned from purple-green in spring to purple in summer, a process accompanied by a significant decrease (P<0.05) in the content of free amino acids and a significant increase (P<0.05) in total polyphenols and catechins. Untargeted metabolomics identified 116 common differentially expressed metabolites (DEMs) between the purple and green phenotypes, with flavonoids being the most predominant class (31 metabolites). Notably, Lr1 exhibited a unique metabolic reprogramming during its seasonal transition, characterized by the specific activation of pathways such as monoterpenoid biosynthesis. Further targeted quantitative approach revealed that the seasonal purpling of Lr1 was driven by an explosive accumulation of cyanidin and delphinidin-O-galactoside in summer, an accumulation pattern that stood in stark contrast to that of 'Zijuan', whose levels decreased in summer. In conclusion, this study demonstrated that the seasonal purpling of Lr1 was a dynamic response process mediated by the massive de novo synthesis of specific anthocyanins (particularly cyanidin), induced by environmental factors. Its metabolic regulatory model was fundamentally different from that of constitutively purple cultivars. This unique germplasm provides a valuable genetic resource and a theoretical basis for understanding the environmental adaptability of tea plants and for breeding novel specialty cultivars.

, authors=Jing HUANG, Lingling WU, Hongying WU, Yuqin NONG, Yanfei LUO, Mingshi WANG, Jingyang YANG, Xiaomin QIN, Yuanquan CHEN*, authorsList=Jing HUANG, Lingling WU, Hongying WU, Yuqin NONG, Yanfei LUO, Mingshi WANG, Jingyang YANG, Xiaomin QIN, Yuanquan CHEN, authorCompany=null, correspAuthors=Yuanquan CHEN, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright © 2026 Science and Technology of Food Industry. All rights reserved., 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=1320880575452394339, articleId=1320880573862753109, tenantId=1146029695717560320, journalId=1260987677001138203, language=CN, title=基于非靶向代谢组学解析茶树新品种‘龙蕊1号’的夏季紫化表型, columnId=1320880262997717526, journalTitle=食品工业科技, columnName=现代分析与智能检测, runingTitle=null, highlight=null, articleAbstract=

为阐明特异茶树种质资源‘龙蕊1号’芽叶颜色随季节变化的分子机制,本研究以稳定绿色品种‘中茶108’和紫色品种‘紫娟’为对照,综合运用生化测定、非靶向与靶向代谢组学技术,系统解析了‘龙蕊1号’季节性紫化表型在叶片色泽、品质成分与代谢谱等方面的动态规律与形成基础。结果表明,‘龙蕊1号’芽叶由春季的紫绿色转变为夏季的紫色,该过程伴随着游离氨基酸含量显著(P<0.05)下降及总多酚和儿茶素含量显著(P<0.05)上升。非靶向代谢组学在紫色与绿色表型间共筛选出116个共有差异代谢物,其中类黄酮占比最高(31种)。‘龙蕊1号’在季节性转变中表现出独特的代谢重编程,其单萜类生物合成等通路被特异性激活,进一步靶向分析的结果显示,‘龙蕊1号’的季节性紫化是由矢车菊素和飞燕草素-O-半乳糖苷在夏季的爆发式增长所驱动的,此积累模式与‘紫娟’(夏季含量下降)形成鲜明对比。综上,‘龙蕊1号’的季节性紫化是一种由环境因子诱导、特定花青苷(尤其是矢车菊素)大量从头合成所介导的动态响应过程,其代谢调控模式与组成型紫色品种存在本质区别。该特异种质为了解茶树环境适应性及培育特色新品种提供了宝贵的遗传资源与理论依据。

, authors=黄静, 吴玲玲, 巫虹颖, 农玉琴, 骆妍妃, 王明释, 阳景阳, 覃潇敏, 陈远权*, authorsList=黄静, 吴玲玲, 巫虹颖, 农玉琴, 骆妍妃, 王明释, 阳景阳, 覃潇敏, 陈远权, authorCompany=null, correspAuthors=陈远权, authorNote=

黄静(1998−),女,硕士,研究方向:茶树育种与茶叶品质,E-mail:

, correspAuthorsNote=
陈远权(1982−),男,硕士,高级农艺师,研究方向:茶树育种栽培与茶叶加工,E-mail:。
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Journal of Agricultural and Food Chemistry, 2022, 70(6): 1830−1839., articleTitle=null, refAbstract=null)], funds=null, companyList=[AuthorCompany(id=1320880575670498148, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, xref=null, ext=[AuthorCompanyExt(id=1320880575678886758, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, companyId=1320880575670498148, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Guangxi South Subtropical Agricultural Sciences Research Institute, Chongzuo 532415, China), AuthorCompanyExt(id=1320880575687275367, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, companyId=1320880575670498148, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=广西南亚热带农业科学研究所,广西崇左 532415)])], figs=[ArticleFig(id=1320880580728828834, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.1, caption=Bud and leaf phenotypes (A), free amino acids (B), total polyphenols (C), and catechin (D) contents of 'Zhongcha 108' 'Zijuan', and 'Longrui No.1' in spring and summer seasons, figureFileSmall=6W0bFXh/P3UYh8wn+cITfw==, figureFileBig=tUn5/WWdNrQOLUnASVS5lw==, tableContent=null), ArticleFig(id=1320880580808520611, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图1, caption=‘中茶108’‘紫娟’与‘龙蕊1号’在春、夏两季的芽叶表型(A)、游离氨基酸(B)、总多酚(C)及儿茶素(D)含量

注:B~D中,小写字母表示具有显著性差异(P<0.05)。

, figureFileSmall=6W0bFXh/P3UYh8wn+cITfw==, figureFileBig=tUn5/WWdNrQOLUnASVS5lw==, tableContent=null), ArticleFig(id=1320880582540768168, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.2, caption=Correlation matrix of metabolic profiles (A), OPLS-DA scores (B), and model diagnostics (C) for 'Zhongcha 108' 'Zijuan', and 'Longrui No.1' in spring and summer seasons, figureFileSmall=g2hUyamsFb5uP+581uLLZg==, figureFileBig=I514LcAU1VAQWfcLvzjR7A==, tableContent=null), ArticleFig(id=1320880582637237161, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图2, caption=‘中茶108’‘紫娟’与‘龙蕊1号’在春、夏两季代谢谱的相关性矩阵(A)、OPLS-DA得分(B)以及模型诊断(C), figureFileSmall=g2hUyamsFb5uP+581uLLZg==, figureFileBig=I514LcAU1VAQWfcLvzjR7A==, tableContent=null), ArticleFig(id=1320880582721123242, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.3, caption=DEMs among different leaf color samples, figureFileSmall=D+QPW2kqvfhbBJxm5ZYKrw==, figureFileBig=+iCbk4MLJx9TSJ8gycS3Ww==, tableContent=null), ArticleFig(id=1320880582788232107, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图3, caption=不同叶色样本间的差异代谢物

注:A. 不同叶色差异代谢物上下调情况;B. 不同叶色对比差异代谢物韦恩图;C. 共有叶色差异代谢物分类,图中冒号后的数字表示该分类下的差异代谢物种数。

, figureFileSmall=D+QPW2kqvfhbBJxm5ZYKrw==, figureFileBig=+iCbk4MLJx9TSJ8gycS3Ww==, tableContent=null), ArticleFig(id=1320880582859535276, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.4, caption=Volcano plot (A) and KEGG pathway enrichment (B) analysis of DEMs on 'Lr1' 'ZC108' and 'ZJ' across spring and summer seasons, figureFileSmall=/HupP9Xa20OXYpkyMQISnA==, figureFileBig=0z5agHsOWERhSFEHqQ5xFg==, tableContent=null), ArticleFig(id=1320880582922449837, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图4, caption=‘龙蕊1号’‘中茶108’和‘紫娟’的春、夏季节差异代谢物火山图(A)和KEGG通路富集情况(B), figureFileSmall=/HupP9Xa20OXYpkyMQISnA==, figureFileBig=0z5agHsOWERhSFEHqQ5xFg==, tableContent=null), ArticleFig(id=1320880583002141614, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.5, caption=Seasonal metabolic shifts in 'Lr1' between spring and summer, figureFileSmall=uLROr0TuD81qIEkdwh5jNg==, figureFileBig=+IV600x6SlW2xU5wHNrtgQ==, tableContent=null), ArticleFig(id=1320880583069250479, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图5, caption=‘龙蕊1号’春夏季节间主要差异代谢物的变化

注:A. ‘龙蕊1号’春夏季节间差异代谢物的分类;B. FC>1.5或FC<0.67的差异代谢物;C. 夏季上调积累的二氢硫辛酸、长蒴黄麻素、氯莫环素和毛竹酚的含量,*表示两个季节间的显著性检验结果,P<0.05,**表示P<0.01。

, figureFileSmall=uLROr0TuD81qIEkdwh5jNg==, figureFileBig=+IV600x6SlW2xU5wHNrtgQ==, tableContent=null), ArticleFig(id=1320880583157330864, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Fig.6, caption=Seasonal dynamics in 'ZC08' 'ZJ ', and 'Lr1' on the contents of cyanidinn (A), pelargonidin (B), delphinidin-3-O-galactoside (C) and malvidin-3-O-galactoside (D), figureFileSmall=rdD/czR04Da5FxX0WNATBw==, figureFileBig=6SYLOiaEaKOj/39pp0k3IQ==, tableContent=null), ArticleFig(id=1320880583220245425, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=图6, caption=‘中茶108’‘紫娟’与‘龙蕊1号’中矢车菊素(A)、天竺葵色素(B)、飞燕草素-O-半乳糖苷(C)和锦葵色素-O-半乳糖苷(D)含量的季节性差异

注:ND表示未检出,不同小写字母表示数据差异显著(P<0.05)。

, figureFileSmall=rdD/czR04Da5FxX0WNATBw==, figureFileBig=6SYLOiaEaKOj/39pp0k3IQ==, tableContent=null), ArticleFig(id=1320880583274771378, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=EN, label=Table 1, caption=

Shared DEMs among different leaf color comparisons

, figureFileSmall=null, figureFileBig=null, tableContent=
代谢物中文名称代谢物英文名称二级分类
假荆芥甙Nepitrin黄酮糖苷
木犀草素-7-O-葡萄糖苷Luteolin-7-O-Glucoside黄酮糖苷
6''-(4-羟基肉桂酰基)黄芪甲苷4'-葡萄糖苷6''-(4-Hydroxycinnamoyl) astragalin 4'-glucoside黄酮糖苷
山奈酚-3-O-α-L-阿拉伯糖苷Kaempferol-3-O-alpha-L-arabinoside黄酮糖苷
异槲皮苷Isoquercitrin黄酮糖苷
异甘草苷Isoliquiritin黄酮糖苷
根皮苷Phlorizin黄酮糖苷
山奈酚3-[6'''-对香豆糖基-(1→2)-鼠李糖苷]Kaempferol 3-[6'''-p-coumarylglucosyl-(1→2)-rhamnoside]黄酮糖苷
5,7 -二羟基-6-甲氧基黄酮5-鼠李糖苷5,7-Dihydroxy-6-methoxyflavone 5-rhamnoside黄酮糖苷
儿茶素7-葡萄糖苷Catechin 7-glucoside黄酮糖苷
斯皮诺素CSpinosin C黄酮糖苷
柚皮苷Naringin黄酮糖苷
高圣草酚Homoeriodictyol黄酮
针依瓦菊素Acerosin黄酮
矮牵牛素-3-O-Beta-葡萄糖苷Petunidin-3-O-beta-glucopyranoside花青素
矢车菊素-3-O-鼠李糖苷Cyanidin-3-O-Rhamnoside花青素
矮牵牛素3-(6''-对香豆酰基葡萄糖苷)Petunidin 3-(6''-p-coumaroyl-glucoside)花青素
4β-(2-氨基乙基硫代)表儿茶素3-没食子酸酯4beta-(2-Aminoethylthio) epicatechin 3-gallate黄烷醇
二苯甲酮ATheadibenzotropolone A黄烷醇
3'-羟基雌马酚3'-Hydroxyequol异黄酮
染料木素4'-O-葡萄糖醛酸苷Genistein 4'-O-glucuronide异黄酮
黄檀素Dalbergin新黄酮
鼠李醚Rhamnetin黄酮醇
没食子酸-(4alpha→8)-没食子酸-(4alpha→8)-没食子酸Gallocatechin-(4alpha→8)-gallocatechin-(4alpha→8)-gallocatechin双黄酮和多黄酮
苞叶素Bracteatin其他黄酮类
4-p-香豆酰基-1,5-喹啉内酯4-p-Coumaroyl-1,5-quinolactone肉桂酸及其衍生物
桂叶苷AOsmanthuside A肉桂酸及其衍生物
2-O-对-香豆酰羟基柠檬酸2-O-p-Coumaroylhydroxycitric acid肉桂酸及其衍生物
4R,5R,6S-三羟基-2-羟甲基-2-环己烯-1-酮6-(2-羟基-6-甲基苯甲酸酯)4R,5R,6S-Trihydroxy-2-hydroxymethyl-2-cyclohexen-1-one 6-(2-hydroxy-6-methylbenzoate)苯甲酸及其衍生物
2,3,6-三-O-没食子酰基-Beta-D-葡萄糖硫苷Methyl 2,3,6-tri-O-galloyl-beta-D-glucopyranoside苯甲酸及其衍生物
甲基-6-O-没食子酰基-Beta-D-葡萄糖硫苷Methyl 6-O-galloyl-beta-D-glucopyranoside苯甲酸及其衍生物
足叶草毒素Podofilox木脂素类及其衍生物
杨梅苷IMyricatomentoside I木脂素类及其衍生物
芹菜香豆精苷Apiumoside香豆素类及其衍生物
8-羟基-R-醋硝香豆素8-Hydroxy-R-acenocoumarol香豆素类及其衍生物
2-O-(4-羟基肉桂酰基)-1,6-二-O-没食子酰基-β-D-吡喃葡萄糖2-O-(4-Hydroxycinnamoyl)-1,6-di-O-galloyl-beta-D-glucopyranose鞣质
6 -肉桂酰-1-没食子酰葡萄糖6-Cinnamoyl-1-galloylglucose鞣质
山茶酮二醇Camellenodiol三萜
3-羟基-8'-Apo-ε-胡萝卜素-8'-醛3-Hydroxy-8'-apo-epsilon-caroten-8'-al三萜
大豆甾醇ASoyasapogenol A三萜
甘草次酸CGanolucidic acid C三萜
D-吡喃葡萄糖醛酸Tragopogonsaponin A三萜皂苷
20,24 -环氧-25,26 -二羟基达玛烷-3-酮20,24-Epoxy-25,26-dihydroxydammaran-3-one三萜皂苷
(+)-α-蒎烯(+)-alpha-pinene单萜
植基焦磷酸Phytyl diphosphate二萜
款冬花苷Cofaryloside其他萜类
脱氧胸苷二磷酸- L -鼠李糖Deoxythymidine diphosphate-L-rhamnose核苷酸及其衍生物
N1-(5-磷酸-a-D-核糖基)-5,6-二甲基苯并咪唑N1-(5-Phospho-a-D-ribosyl)-5,6-dimethylbenzimidazole核苷酸及其衍生物
水粉蕈素Nebularine核苷酸及其衍生物
苯丙酮酸Phenylpyruvic Acid有机酸及其衍生物
3,4 -二羟基苯丙酮酸3,4-dihydroxyphenylpyruvate有机酸及其衍生物
5-咖啡酰奎宁酸5-Caffeoylquinic acid奎宁酸及其衍生物
L - 2 -氨基- 3 -亚甲基己酸L-2-Amino-3-methylenehexanoic acidAlpha氨基酸
N-[[3-(b-D-吡喃葡萄糖基氧基)-2,3-二氢-2-氧代-1H-吲哚-3-基]乙酰基]天冬氨酸N-[[3-(b-D-Glucopyranosyloxy)-2,3-dihydro-2-oxo-1H-indol-3-yl]acetyl]aspartic acid其他类氨基酸
马齿苋黄素ⅡPortulacaxanthin II其他类氨基酸
酪氨酰基亮氨酸Tyrosyl-Leucine其他类氨基酸
酪氨酰酪氨酸Tyrosyl-Tyrosine其他类氨基酸
3,4,5-三甲氧基苯基6-O-D-芹糖-beta-D-呋喃糖基-beta-D-葡萄糖苷Kelampayoside A其他类碳水化合物
羟基酪醇4'-O-葡萄糖苷Hydroxytyrosol 4'-O-glucoside其他类碳水化合物
葡萄糖菌素Glucocleomin糖苷
茵芋苷Skimmin糖苷
蔷薇甙Multiflorin A糖苷
胡黄连苦甙ⅠPicroside I糖苷
鸡蛋花苷Plumieride糖苷
甘草苷BLicoagroside B糖脂类
24R,25-二羟基维生素D324R,25-Dihydroxyvitamin D3维生素类
骨化二醇Calcidiol维生素类
芫荽异香豆酮DCoriandrone D苯骈吡喃类
桑皮酮QKuwanon Q二苯基庚烷类
麦角日亭Ergocristine其他类生物碱
小蘖红碱Berberrubine异喹啉类生物碱
11β-羟孕酮11beta-Hydroxyprogesterone孕甾烷
7,8-二氢Vomifoliol 9-[芹菜糖基-(1→6)-葡萄糖苷]7,8-Dihydrovomifoliol 9-[apiosyl-(1→6)-glucoside]脂肪酰类
7 -甲基黄嘌呤7-Methylxanthine其他类
依维莫司Everolimus其他类
苦龙苷Amarogentin其他类
10-乙酰氧基苯酚10-Acetoxytoxol其他类
5-苯基-1,3-噁嗪烷-2,4 -二酮5-Phenyl-1,3-oxazinane-2,4-dione其他类
普伐他汀Pravastatin其他类
泽泻醇BAlisol B其他类
质体醌Plastoquinone其他类
双草醚Bispyribac其他类
2-琥珀酰基-5-烯醇丙酮酸-6-羟基-3-环己烯-1-羧酸酯2-succinyl-5-enolpyruvyl-6-hydroxy-3-cyclohexene-1-carboxylate其他类
), ArticleFig(id=1320880583350268851, tenantId=1146029695717560320, journalId=1260987677001138203, articleId=1320880573862753109, language=CN, label=表1, caption=

共有叶色差异代谢物

, figureFileSmall=null, figureFileBig=null, tableContent=
代谢物中文名称代谢物英文名称二级分类
假荆芥甙Nepitrin黄酮糖苷
木犀草素-7-O-葡萄糖苷Luteolin-7-O-Glucoside黄酮糖苷
6''-(4-羟基肉桂酰基)黄芪甲苷4'-葡萄糖苷6''-(4-Hydroxycinnamoyl) astragalin 4'-glucoside黄酮糖苷
山奈酚-3-O-α-L-阿拉伯糖苷Kaempferol-3-O-alpha-L-arabinoside黄酮糖苷
异槲皮苷Isoquercitrin黄酮糖苷
异甘草苷Isoliquiritin黄酮糖苷
根皮苷Phlorizin黄酮糖苷
山奈酚3-[6'''-对香豆糖基-(1→2)-鼠李糖苷]Kaempferol 3-[6'''-p-coumarylglucosyl-(1→2)-rhamnoside]黄酮糖苷
5,7 -二羟基-6-甲氧基黄酮5-鼠李糖苷5,7-Dihydroxy-6-methoxyflavone 5-rhamnoside黄酮糖苷
儿茶素7-葡萄糖苷Catechin 7-glucoside黄酮糖苷
斯皮诺素CSpinosin C黄酮糖苷
柚皮苷Naringin黄酮糖苷
高圣草酚Homoeriodictyol黄酮
针依瓦菊素Acerosin黄酮
矮牵牛素-3-O-Beta-葡萄糖苷Petunidin-3-O-beta-glucopyranoside花青素
矢车菊素-3-O-鼠李糖苷Cyanidin-3-O-Rhamnoside花青素
矮牵牛素3-(6''-对香豆酰基葡萄糖苷)Petunidin 3-(6''-p-coumaroyl-glucoside)花青素
4β-(2-氨基乙基硫代)表儿茶素3-没食子酸酯4beta-(2-Aminoethylthio) epicatechin 3-gallate黄烷醇
二苯甲酮ATheadibenzotropolone A黄烷醇
3'-羟基雌马酚3'-Hydroxyequol异黄酮
染料木素4'-O-葡萄糖醛酸苷Genistein 4'-O-glucuronide异黄酮
黄檀素Dalbergin新黄酮
鼠李醚Rhamnetin黄酮醇
没食子酸-(4alpha→8)-没食子酸-(4alpha→8)-没食子酸Gallocatechin-(4alpha→8)-gallocatechin-(4alpha→8)-gallocatechin双黄酮和多黄酮
苞叶素Bracteatin其他黄酮类
4-p-香豆酰基-1,5-喹啉内酯4-p-Coumaroyl-1,5-quinolactone肉桂酸及其衍生物
桂叶苷AOsmanthuside A肉桂酸及其衍生物
2-O-对-香豆酰羟基柠檬酸2-O-p-Coumaroylhydroxycitric acid肉桂酸及其衍生物
4R,5R,6S-三羟基-2-羟甲基-2-环己烯-1-酮6-(2-羟基-6-甲基苯甲酸酯)4R,5R,6S-Trihydroxy-2-hydroxymethyl-2-cyclohexen-1-one 6-(2-hydroxy-6-methylbenzoate)苯甲酸及其衍生物
2,3,6-三-O-没食子酰基-Beta-D-葡萄糖硫苷Methyl 2,3,6-tri-O-galloyl-beta-D-glucopyranoside苯甲酸及其衍生物
甲基-6-O-没食子酰基-Beta-D-葡萄糖硫苷Methyl 6-O-galloyl-beta-D-glucopyranoside苯甲酸及其衍生物
足叶草毒素Podofilox木脂素类及其衍生物
杨梅苷IMyricatomentoside I木脂素类及其衍生物
芹菜香豆精苷Apiumoside香豆素类及其衍生物
8-羟基-R-醋硝香豆素8-Hydroxy-R-acenocoumarol香豆素类及其衍生物
2-O-(4-羟基肉桂酰基)-1,6-二-O-没食子酰基-β-D-吡喃葡萄糖2-O-(4-Hydroxycinnamoyl)-1,6-di-O-galloyl-beta-D-glucopyranose鞣质
6 -肉桂酰-1-没食子酰葡萄糖6-Cinnamoyl-1-galloylglucose鞣质
山茶酮二醇Camellenodiol三萜
3-羟基-8'-Apo-ε-胡萝卜素-8'-醛3-Hydroxy-8'-apo-epsilon-caroten-8'-al三萜
大豆甾醇ASoyasapogenol A三萜
甘草次酸CGanolucidic acid C三萜
D-吡喃葡萄糖醛酸Tragopogonsaponin A三萜皂苷
20,24 -环氧-25,26 -二羟基达玛烷-3-酮20,24-Epoxy-25,26-dihydroxydammaran-3-one三萜皂苷
(+)-α-蒎烯(+)-alpha-pinene单萜
植基焦磷酸Phytyl diphosphate二萜
款冬花苷Cofaryloside其他萜类
脱氧胸苷二磷酸- L -鼠李糖Deoxythymidine diphosphate-L-rhamnose核苷酸及其衍生物
N1-(5-磷酸-a-D-核糖基)-5,6-二甲基苯并咪唑N1-(5-Phospho-a-D-ribosyl)-5,6-dimethylbenzimidazole核苷酸及其衍生物
水粉蕈素Nebularine核苷酸及其衍生物
苯丙酮酸Phenylpyruvic Acid有机酸及其衍生物
3,4 -二羟基苯丙酮酸3,4-dihydroxyphenylpyruvate有机酸及其衍生物
5-咖啡酰奎宁酸5-Caffeoylquinic acid奎宁酸及其衍生物
L - 2 -氨基- 3 -亚甲基己酸L-2-Amino-3-methylenehexanoic acidAlpha氨基酸
N-[[3-(b-D-吡喃葡萄糖基氧基)-2,3-二氢-2-氧代-1H-吲哚-3-基]乙酰基]天冬氨酸N-[[3-(b-D-Glucopyranosyloxy)-2,3-dihydro-2-oxo-1H-indol-3-yl]acetyl]aspartic acid其他类氨基酸
马齿苋黄素ⅡPortulacaxanthin II其他类氨基酸
酪氨酰基亮氨酸Tyrosyl-Leucine其他类氨基酸
酪氨酰酪氨酸Tyrosyl-Tyrosine其他类氨基酸
3,4,5-三甲氧基苯基6-O-D-芹糖-beta-D-呋喃糖基-beta-D-葡萄糖苷Kelampayoside A其他类碳水化合物
羟基酪醇4'-O-葡萄糖苷Hydroxytyrosol 4'-O-glucoside其他类碳水化合物
葡萄糖菌素Glucocleomin糖苷
茵芋苷Skimmin糖苷
蔷薇甙Multiflorin A糖苷
胡黄连苦甙ⅠPicroside I糖苷
鸡蛋花苷Plumieride糖苷
甘草苷BLicoagroside B糖脂类
24R,25-二羟基维生素D324R,25-Dihydroxyvitamin D3维生素类
骨化二醇Calcidiol维生素类
芫荽异香豆酮DCoriandrone D苯骈吡喃类
桑皮酮QKuwanon Q二苯基庚烷类
麦角日亭Ergocristine其他类生物碱
小蘖红碱Berberrubine异喹啉类生物碱
11β-羟孕酮11beta-Hydroxyprogesterone孕甾烷
7,8-二氢Vomifoliol 9-[芹菜糖基-(1→6)-葡萄糖苷]7,8-Dihydrovomifoliol 9-[apiosyl-(1→6)-glucoside]脂肪酰类
7 -甲基黄嘌呤7-Methylxanthine其他类
依维莫司Everolimus其他类
苦龙苷Amarogentin其他类
10-乙酰氧基苯酚10-Acetoxytoxol其他类
5-苯基-1,3-噁嗪烷-2,4 -二酮5-Phenyl-1,3-oxazinane-2,4-dione其他类
普伐他汀Pravastatin其他类
泽泻醇BAlisol B其他类
质体醌Plastoquinone其他类
双草醚Bispyribac其他类
2-琥珀酰基-5-烯醇丙酮酸-6-羟基-3-环己烯-1-羧酸酯2-succinyl-5-enolpyruvyl-6-hydroxy-3-cyclohexene-1-carboxylate其他类
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基于非靶向代谢组学解析茶树新品种‘龙蕊1号’的夏季紫化表型
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黄静 , 吴玲玲 , 巫虹颖 , 农玉琴 , 骆妍妃 , 王明释 , 阳景阳 , 覃潇敏 , 陈远权 *
食品工业科技 | 现代分析与智能检测 2026,47(18): 284-294
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食品工业科技 |现代分析与智能检测 2026 , 47 (18) : 284 -294
基于非靶向代谢组学解析茶树新品种‘龙蕊1号’的夏季紫化表型
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黄静 , 吴玲玲, 巫虹颖, 农玉琴, 骆妍妃, 王明释, 阳景阳, 覃潇敏, 陈远权*
作者信息
  • 广西南亚热带农业科学研究所,广西崇左 532415
通讯作者:
陈远权(1982−),男,硕士,高级农艺师,研究方向:茶树育种栽培与茶叶加工,E-mail:。
作者简介:

黄静(1998−),女,硕士,研究方向:茶树育种与茶叶品质,E-mail:

Revealing Summer Leaf Purpling Phenotype of Camellia sinensis cv. 'Longrui 1' with Untargeted Metabolomics
Jing HUANG , Lingling WU, Hongying WU, Yuqin NONG, Yanfei LUO, Mingshi WANG, Jingyang YANG, Xiaomin QIN, Yuanquan CHEN*
Affiliations
  • Guangxi South Subtropical Agricultural Sciences Research Institute, Chongzuo 532415, China
出版时间: 2026-09-15 doi: 10.13386/j.issn1002-0306.2025060101
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为阐明特异茶树种质资源‘龙蕊1号’芽叶颜色随季节变化的分子机制,本研究以稳定绿色品种‘中茶108’和紫色品种‘紫娟’为对照,综合运用生化测定、非靶向与靶向代谢组学技术,系统解析了‘龙蕊1号’季节性紫化表型在叶片色泽、品质成分与代谢谱等方面的动态规律与形成基础。结果表明,‘龙蕊1号’芽叶由春季的紫绿色转变为夏季的紫色,该过程伴随着游离氨基酸含量显著(P<0.05)下降及总多酚和儿茶素含量显著(P<0.05)上升。非靶向代谢组学在紫色与绿色表型间共筛选出116个共有差异代谢物,其中类黄酮占比最高(31种)。‘龙蕊1号’在季节性转变中表现出独特的代谢重编程,其单萜类生物合成等通路被特异性激活,进一步靶向分析的结果显示,‘龙蕊1号’的季节性紫化是由矢车菊素和飞燕草素-O-半乳糖苷在夏季的爆发式增长所驱动的,此积累模式与‘紫娟’(夏季含量下降)形成鲜明对比。综上,‘龙蕊1号’的季节性紫化是一种由环境因子诱导、特定花青苷(尤其是矢车菊素)大量从头合成所介导的动态响应过程,其代谢调控模式与组成型紫色品种存在本质区别。该特异种质为了解茶树环境适应性及培育特色新品种提供了宝贵的遗传资源与理论依据。

非靶向代谢组  /  南山白毛茶  /  ‘龙蕊1号’  /  紫化表型  /  紫化机理

To elucidate the molecular mechanism underlying the seasonal variation in leaf color of the unique tea plan germplasm 'Longrui 1' (Camellia sinensis cv. 'Longrui 1', Lr1), the dynamic patterns and underlying mechanisms of the seasonal purple phenotype in 'Longrui 1', particularly regarding leaf coloration, quality components, and metabolic profiles, were systematically investigated. A combined approach of biochemical assays, untargeted and targeted metabolomics was employed, using the constitutively green cultivar 'Zhongcha 108' and the constitutively purple cultivar 'Zijuan' as controls. The results indicated that the young shoots of Lr1 transitioned from purple-green in spring to purple in summer, a process accompanied by a significant decrease (P<0.05) in the content of free amino acids and a significant increase (P<0.05) in total polyphenols and catechins. Untargeted metabolomics identified 116 common differentially expressed metabolites (DEMs) between the purple and green phenotypes, with flavonoids being the most predominant class (31 metabolites). Notably, Lr1 exhibited a unique metabolic reprogramming during its seasonal transition, characterized by the specific activation of pathways such as monoterpenoid biosynthesis. Further targeted quantitative approach revealed that the seasonal purpling of Lr1 was driven by an explosive accumulation of cyanidin and delphinidin-O-galactoside in summer, an accumulation pattern that stood in stark contrast to that of 'Zijuan', whose levels decreased in summer. In conclusion, this study demonstrated that the seasonal purpling of Lr1 was a dynamic response process mediated by the massive de novo synthesis of specific anthocyanins (particularly cyanidin), induced by environmental factors. Its metabolic regulatory model was fundamentally different from that of constitutively purple cultivars. This unique germplasm provides a valuable genetic resource and a theoretical basis for understanding the environmental adaptability of tea plants and for breeding novel specialty cultivars.

untargeted metabolomics  /  Nanshan Baimaocha  /  'Longrui 1'  /  purpling phenotype  /  purpling mechanism
黄静, 吴玲玲, 巫虹颖, 农玉琴, 骆妍妃, 王明释, 阳景阳, 覃潇敏, 陈远权. 基于非靶向代谢组学解析茶树新品种‘龙蕊1号’的夏季紫化表型. 食品工业科技, 2026 , 47 (18) : 284 -294 . DOI: 10.13386/j.issn1002-0306.2025060101
Jing HUANG, Lingling WU, Hongying WU, Yuqin NONG, Yanfei LUO, Mingshi WANG, Jingyang YANG, Xiaomin QIN, Yuanquan CHEN. Revealing Summer Leaf Purpling Phenotype of Camellia sinensis cv. 'Longrui 1' with Untargeted Metabolomics[J]. Science and Technology of Food Industry, 2026 , 47 (18) : 284 -294 . DOI: 10.13386/j.issn1002-0306.2025060101
茶树(Camellia sinensis (L) O.Kuntze)是中国原生的经济作物,起源于中国西南地区[1]。茶叶从最初的药用发展到现今的饮用经历了数千年的历史,其栽培利用及茶文化已遍及世界多个国家和地区,成为世界三大非酒精饮料之一。特异叶色茶树是不同于常规绿色表型茶树的稀有资源,其内含物质与表型绿色的茶树也有很大不同[2−3]。其中,紫化茶树由于其独特的外观和内涵成分[4],其特色产品的开发利用逐渐受到关注。目前,紫化茶树在中国、日本[5−6]、肯尼亚等国家作为一种特异性茶树资源广泛种植。
紫化茶树的独特之处在于其富含花青素类物质,花青素是一类重要的水溶性天然色素,广泛存在于几乎所有植物的根、茎、叶、花与果实中,在分类上属于黄酮类化合物。通常而言,茶树紫色芽叶中的花青素含量可高达普通品种的数倍,主要包括飞燕草素、矢车菊素、天竺葵素及其糖苷[7]。花青素在茶树中的合成主要涉及苯丙烷代谢途径和类黄酮代谢途径,以苯丙氨酸作为底物,经一系列反应合成二氢黄酮醇(如二氢杨梅素、二氢槲皮素等),随后经过二氢黄酮醇4-还原酶(Dihydroflavonol 4-reductase,FDR)和花青素合成酶(Anthocyanidin synthase,ANS)的接连催化合成花青素[2]。花青素在类黄酮3-O-葡萄糖基转移酶(flavonoid 3-O-glucosyltransferase,UFGT)的催化下进一步糖苷化,合成花青苷[4]。茶树中导致紫芽表型的花青素积累受遗传因素和环境因素的共同调节。一方面,如‘紫娟’‘紫嫣’等品种在正常栽培条件下即具有比一般品种更高的花青素含量[8];另一方面,茶树中的花青素积累受到温度、光照强度、光质等因素的影响[4]。
本课题组前期对南山白毛茶群体进行了表型与生化组分分析,发现这些资源的遗传多样性丰富,在品质、抗性、产量相关性状上具有很好的优势[9−10],在南山白毛茶群体中发现并选育了一个紫化茶树资源‘龙蕊1号’,其形态特征明显区别于当下较出名的育成品种‘紫娟’。‘龙蕊1号’的新梢在春季为紫绿色,而到夏季则转变成紫红色。目前,针对其他类型的茶树叶色变异,如白化、黄化等,已鉴定出光敏型(浮梁黄叶突变体)、温敏型(安吉白茶)、生态不敏感型(炎陵银边茶)等[11],并对其叶色转变机制有了一定的研究[12−13]。而茶树中季节性紫化的现象鲜有报道,这类叶色转变的物质基础和分子机制也不明确。因此,本研究采用非靶向代谢组学技术,结合表型观察和生化成分测定,对新选育品种‘龙蕊1号’的季节型叶色变化进行全面表征,进一步为分子机制研究和开发利用打下基础。
南山白毛茶‘龙蕊1号’(Lr1)、‘紫鹃’(ZJ)、‘中茶108’(ZC108) 来自广西南亚热带农业科学研究所茶树种质资源圃,每个样本设置3个重复;甲醇(分析纯)、茚三酮(分析纯)、碳酸钠、福林酚、磷酸氢二钠、磷酸二氢钾、磷酸、盐酸、甲酸、无水乙醇 国药集团化学试剂有限公司;HPLC级甲醇、乙腈 纯度≥99.9%,OCEANPAK;茶氨酸(纯度≥99%)、儿茶素(C)标准品(纯度≥98%) 上海源叶生物科技有限公司;飞燕草素-O-半乳糖苷、锦葵色素-O-半乳糖苷、天竺葵色素、矢车菊素标准品 纯度≥98%,武汉天植。
Waters2695高效液相色谱仪 沃特世科技(上海)有限公司;Agilent1100高效液相色谱仪 安捷伦科技(中国)有限公司;HT165离心机 湖南湘仪实验室仪器开发有限公司;PX124ZH/E分析天平 奥豪斯仪器(常州)有限公司;KQ-400E超声波清洗器 昆山市超声仪器有限公司;ZGDCY-24干式氮吹仪 绍兴上虞祥达仪器制造有限公司;HH600恒温水浴锅 绍兴上虞祥达仪器制造有限公司;XW-80A旋涡混合器 上海沪西分析仪器厂有限公司;Alpha1-2LD plus冷冻干燥机 德采实仪器设备(北京)有限公司;Wonbio-96c多样品冷冻研磨仪 上海万柏生物科技有限公司。
鲜叶样品:于2024年4月(春)、7月(夏)取生长良好、无病虫害的茶树一芽二叶鲜叶用液氮固定,运回实验室置于−80 ℃冰箱保存。
儿茶素(C)及总多酚测定方法参照GB/T 8313-2018《茶叶中茶多酚和儿茶素类含量的检测方法》;氨基酸测定方法参照GB/T 8314-2013《茶 游离氨基酸总量的测定》。
经液氮研磨的茶鲜叶粉末0.1 g,加入400 μL冷甲醇溶液(甲醇:水=4:1)于样品中;低温下,高通量组织破碎仪破碎;涡旋混匀后,冰上超声萃取10 min,重复3次;将样品于−20 ℃静置30 min;离心13000×g,4 ℃,15 min,离心后取上清液,用微孔滤膜(0.22 μm)过滤样品后,转移至LC-MS带内衬管的进样小瓶中上机检测。
色谱柱为BEH C18柱(100 mm×2.1 mm,1.7 μm);流动相A为水(含0.1%甲酸),流动相B为乙腈/异丙醇(1/1,含0.1%甲酸);梯度洗脱程序为0~3 min:0~20% B,3~9 min:20%~60% B,9~11 min:60%~100% B,100% B维持2.5 min,13.5~13.6 min:100%~0% B,0% B维持2.4 min。流速为0.40 mL/min,进样量为20 μL,柱温为40 ℃。
样品质谱信号采集分别采用正负离子扫描模式,电喷雾毛细管电压、进样电压和碰撞电压分别为:1.0 kV、40 V和6 eV。离子源温度和去溶剂温度分别为:120 和500 ℃,载气流量:900 L/h,质谱扫描范围:m/z 50~1000,分辨率为:30000。
本研究中共有3个品种、两个季节的样本,分别命名为Lr1_Spring、Lr1_Summer、ZJ_Spring、ZJ_Summer、ZC108_Spring和ZC108_Summer,每个样本6次重复。为了评价在上机过程中分析系统的稳定性,实验过程中会制备一个质控样品(Quality Control,QC)。QC样品由所有检测样品混合而成,在仪器分析的过程中,每8个分析样品插入一个QC样品。
飞燕草素-O-半乳糖苷和锦葵色素-O-半乳糖苷测定[14−15]:称取约0.2 g真空冷冻干燥后的茶粉,加入1.0 mL 1%盐酸甲醇,研磨仪研磨成浆,4 ℃超声提取30 min。离心取上清液,Nylon针式滤器(滤膜0.22 μm)过滤后待测。采用Waters2695高效液相色谱仪进行检测,波长530 nm,色谱柱为Compass C18-2反相色谱柱(250 mm×4.6 mm,5 μm);柱温为30 ℃;流速为1 mL/min,进样体积10 μL。
天竺葵色素和矢车菊素测定:称取约0.3 g真空冷冻干燥后的茶粉,加入1.5 mL 10%盐酸甲醇,研磨仪研磨成浆,超声提取1 h,8000×g离心10 min,取上清液0.75 mL,加入0.75 mL试剂1(乙醇:水:盐酸=2:1:1),于沸水浴中水解1 h,试剂一定容至2.0 mL,混匀,Nylon 针式滤器(滤膜0.22 μm)过滤后待测。采用Agilent 1100高效液相色谱仪进行检测,波长530 nm,色谱柱为C18-BP反相色谱柱(250 mm×4.6 mm,5 μm);柱温设置为30 ℃;流速为1.0 mL/min,进样体积5 μL。
代谢组样本上机完成后由美吉生物进行原始数据处理以及代谢物注释,得到所有样本的代谢物信息。对于代谢物矩阵,去除每组内缺失值均>20%的代谢物,对于缺失值采用所有样本中的极小值填充。采用总和归一化法对样本质谱峰的响应强度进行归一化,同时去除QC样本的相对标准偏差(Relative standard deviation,RSD)>30%的值,并lg对数化处理。经上述预处理的数据用于后续所有分析。
使用R语言程序包ropls v1.6.2[16]进行正交偏最小二乘判别(Orthogonal partial least squares discriminant analysis,OPLS-DA)分析。基于OPLD-DA模型得到的变量权重值(Variant important in projection,VIP)和t检验的P值来筛选差异代谢物(Differentially expressed metabolites,DEM),筛选阈值为VIP>1,P<0.05。
差异代谢物的植物化学分类注释与KEGG富集分析通过Majorbio云分析平台[17]进行。其余数据的统计分析及可视化则在Rv4.4.2环境下,利用tidyverse[18]、ggplot2[19]和ggpubr[20]等程序包实现。其中,采用Tukey's HSD检验对不同样本间的游离氨基酸、总多酚、儿茶素及四种花青素组分的含量进行差异显著性分析(P<0.05)。
本研究对‘中茶108’‘紫娟’及‘龙蕊1号’三个品种在春、夏两季的芽叶表型进行了观察。‘中茶108’芽叶在两季中均呈绿色,‘紫娟’均呈紫色,而‘龙蕊1号’则由春季的绿色转变为夏季的紫色,表现出季节性表型差异(图1A)。
游离氨基酸、总多酚和儿茶素是影响茶树品种适制性的重要品质指标[21]。本研究对三品种两季鲜叶的上述成分进行测定,结果显示(图1B~图1D):‘中茶108’的游离氨基酸含量在两季中均显著(P<0.05)高于‘龙蕊1号’和‘紫娟’,且所有品种均呈现春季高于夏季的趋势。总多酚含量则呈现相反规律,‘中茶108’在两季均显著(P<0.05)低于另外两个品种,且各品种均在夏季出现含量高峰。对于儿茶素,‘中茶108’和‘龙蕊1号’的含量为夏高春低,而‘紫娟’则相反。从品种间比较,春季儿茶素含量为‘紫娟’>‘龙蕊1号’>‘中茶108’,夏季则变为‘龙蕊1号’>‘紫娟’>‘中茶108’。
‘龙蕊1号’是从南山白毛茶群体中选育的特异单株,其芽叶的季节性紫化(图1A)可能与光照调控有关。茶树花青素的季节性积累受光照影响显著,强光下转录因子CsbZIP1与CsMYBL2的互作会抑制MBW复合体的表达,从而调控花青素合成通路[22]。因此,夏季的高光强可能是导致‘龙蕊1号’紫化的主要原因之一。与前人研究结果一致[23−24]。本研究也发现紫色品种(‘紫娟’)及季节性紫化品种(‘龙蕊1号’)的氨基酸含量低于绿色品种(‘中茶108’),但其总多酚和儿茶素含量更高,且后两者在夏季的积累也更普遍(图1B~图1D)。
在本研究中,应用非靶向代谢组方法对‘中茶108’‘龙蕊1号’和‘紫娟’在春、夏两个季节的新梢样本进行了非靶向代谢谱检测,经过注释后共鉴定出代谢物1434种。经过初步质量控制后,共1402种代谢物用于后续分析。
对所有样本的数据计算成对相关系数,并对相关性系数矩阵进行聚类及可视化,结果如图2A所示。QC样本的3个重复高度相关,并紧密地聚为一类,而不同品种、不同季节的样本的各6个重复也各自呈现出最高的相关性和最紧密的聚类关系。这说明在本研究的非靶向代谢组学检测具有良好的技术可重复性和高数据质量,结果准确且能反映出不同样本之间的差异。
进一步的,对筛选后的代谢物进行OPLS-DA建模并进行可视化,以更清晰地反映出不同样本的分布情况。如图2B、图2C所示,OPLS-DA的第一、第二个成分分别解释了总方差的30.1%和21.8%(图2B)。此外,OPLS-DA得分以及样本聚类结果均表明,‘中茶108’‘龙蕊1号’和‘紫娟’品种间差异大于各品种在不同季节间的代谢谱差异。OPLS-DA模型的诊断测试表明,该模型表现出良好的方差解释度和预测能力(R2=0.983,Q2=0.945)。
为筛选与芽叶颜色差异相关的代谢物,本研究对四组紫色/绿色表型(Lr1_Summer vs. ZC108_Summer、ZJ_Spring vs. Lr1_Spring、ZJ_Spring vs. ZC108_Spring、ZJ_Summer vs. ZC108_Summer)进行了非靶向代谢组学分析。各比较组分别鉴定出642、584、671和642种差异代谢物(图3A)。具体而言,Lr1_Summer vs. ZC108_Summer中323个下调、319个上调;ZJ_Spring vs. Lr1_Spring中257个下调,327个上调,ZJ_Spring vs. ZC108_Spring中381个下调,290个上调;ZJ_summer vs. ZC108_summer中326个下调,316个上调(图3A)。
为了确定造成茶树芽叶紫色表型的关键差异代谢物,对这4个组合进行了维恩图分析以寻找交集,共筛选出116个在紫、绿芽叶间稳定变化的共有差异代谢物(图3B)。
对这116个共有差异代谢物的化学分类表明,它们主要由类黄酮(31种)、萜类(12种)以及酚酸及其衍生物(11种)构成(图3C)。具体而言,紫色、绿色芽叶茶树新梢的共有差异代谢物包括31种类黄酮、12种萜类、11种酚酸及其衍生物、4种碳水化合物及其衍生物、3种氨基酸及其衍生物等(部分化合物名称及二级分类见表1,完整差异代谢物列表、化学分类及其在不同样本中的相对含量见附表1(详见本刊官网http://www.spgykj.com/文章的网络版)。其中,多种花色苷和黄酮类物质,如矢车菊素-3-O-鼠李糖苷、山奈酚-3-O-α-L-阿拉伯糖苷及矮牵牛素-3-(6''-对香豆酰基葡萄糖苷)等,在紫色样本(‘紫娟’和‘龙蕊1号’夏季)中的含量均显著高于绿色样本(‘中茶108’和‘龙蕊1号’春季)。此外,还发现部分萜类物质,如20,24-环氧-25,26-二羟基达玛烷-3-酮和3-羟基-8'-Apo-ε-胡萝卜素-8'-醛,在季节性紫化的‘龙蕊1号’中特异性高水平积累,可能与其独特的叶色变化表型相关。
KEGG通路富集分析显示,这些共有差异代谢物显著(P<0.05)富集于类黄酮生物合成、黄酮和黄酮醇生物合成以及泛醌和其他萜类-醌类生物合成等次级代谢途径(附图1A~附图1B,详见本刊官网http://www.spgykj.com/文章的网络版)。该结果与化学分类分析高度一致,表明类黄酮和萜类代谢通路的系统性改变是形成茶树芽叶紫、绿颜色差异的核心生化基础。
为探究代谢物在不同季节的动态变化,本研究分别对三个品种春、夏季节间的差异代谢物进行了鉴定和功能分析。结果显示,在三个品种中分别鉴定出651、754和754个差异代谢物(图4A)。KEGG富集分析表明,这三个品种的季节性代谢谱变化趋势具有高度相似性,其差异代谢物均显著富集于α-亚麻酸代谢、类黄酮生物合成、异黄酮生物合成以及植物激素信号转导等通路(图4B)。这揭示了茶树为适应季节性环境变化,会启动一系列保守的次生代谢及信号转导途径。其中,亚麻酸和类黄酮作为关键响应物质,不仅参与植物的逆境胁迫响应[2],也直接影响茶叶的滋味与香气品质[23,25]。
‘龙蕊1号’展现了其独特的季节性代谢调控模式。在其季节性差异代谢物中,有四个通路是其特有富集的,包括单萜类生物合成、芪类化合物、二芳基庚烷类化合物和姜酚的生物合成、泛醌和其他萜类-醌类的生物合成以及黄酮和黄酮醇生物合成(图4B)。这些特异性变化的代谢通路,尤其是与色素和香气前体相关的萜类和黄酮醇的合成,可能与‘龙蕊1号’独特的季节性叶色紫化表型密切相关。
为阐明‘龙蕊1号’季节性叶色紫化的代谢基础,本文首先对其春、夏两季的651个季节性差异代谢物进行了深入分析(图5A)。这些代谢物主要涵盖类黄酮、脂质及萜类等。为聚焦核心变化,以变化倍数(Fold Change,FC>1.5或<0.67)为标准,筛选得到41个关键差异代谢物(图5B)。包括5种类黄酮、5种脂质、3种萜类、3种碳水化合物及其衍生物、3种甾体及其衍生物、2种鞣质、1种氨基酸及其衍生物、1种酚酸类及其衍生物、1种核苷酸及其衍生物、1种醌类以及16种其他类化合物。其中,多数物质在夏季含量下调,而二氢硫辛酸(脂质)、长蒴黄麻素(甾体)和毛竹酚(二萜)等4种化合物则在夏季显著(P<0.05)上调(图5C),且均表现出显著的季节性差异。在这4种代谢物中,二氢硫辛酸属于脂质代谢途径[26],长蒴黄麻素属于甾体生物合成途径[27],而毛竹酚是一类具有生物活性的二萜类代谢物[28]。
在上述筛选出的核心代谢物中并未包含花青素类物质。考虑到非靶向代谢组学在特定化合物(尤其是低丰度或结构不稳定的花青素)检测上可能存在覆盖不足的局限性[29−30],课题组继而采用靶向定量方法,对四种在紫色茶树中起主要呈色作用的花青苷进行了精确测定[7,31],以验证其在‘龙蕊1号’季节性紫化中的作用。
靶向分析结果(图6)清晰地揭示了‘龙蕊1号’中花青苷的特异性季节性积累模式。与春季相比,其夏季芽叶中的矢车菊素和飞燕草素-O-半乳糖苷含量呈现爆发式增长。这一变化趋势与紫色品种‘紫娟’(夏季含量剧烈下降)形成鲜明对比,也显著(P<0.05)区别于含量始终极低的绿色品种‘中茶108’。此外,锦葵色素-O-半乳糖苷在春季的‘龙蕊1号’中未检出,但在夏季积累显著(P<0.05),且含量远高于‘紫娟’和‘中茶108’。这些结果表明,矢车菊素和飞燕草素-O-半乳糖苷等特定花青苷在夏季的大量从头合成,是驱动‘龙蕊1号’芽叶季节性紫化的直接生化原因。此前基于其他紫化茶树的研究表明,矢车菊素是紫化茶树种含量最高的花青素类物质[32],本研究的结果与之相符。但也有研究声称,在‘紫娟’和紫嫣的杂交子代中发现飞燕草素-O-半乳糖苷是最主要的花青素类物质[8],这与本研究中‘紫娟’样本的高含量飞燕草素-O-半乳糖苷相吻合,但从绝对含量上来说,仍然低于矢车菊素。
茶树芽叶的呈色是决定其种质资源特性、加工适制性及最终产品感官品质的关键农艺性状。本研究以表现出独特季节性紫化表型的新品种‘龙蕊1号’为核心,通过与稳定紫色品种‘紫娟’和稳定绿色品种‘中茶108’的多维度比较,系统探究了其叶色转变的生化基础与代谢调控机制,为理解茶树响应环境变化、调控次级代谢提供了新的见解。
本研究最核心的发现是,‘龙蕊1号’的季节性紫化是由特定花青苷组分在夏季的大量积累导致的。结果清晰地表明,矢车菊素和飞燕草素-O-半乳糖苷在‘龙蕊1号’夏季芽叶中的含量发生剧烈增长,这直接解释了其由春季的紫绿色转变为夏季深紫色的表型。这一积累模式与稳定的紫色品种‘紫娟’形成了鲜明对比,后者在春季已积累了高水平的花青苷,进入夏季后含量反而显著下降。‘龙蕊1号’的合成通路可能受环境因子(如光照)的严格诱导,表现为“动态响应型”。夏季的高光强和高温是驱动这一过程的关键环境因子,这与前人关于光照通过调控MBW转录复合体激活花青素合成通路的报道相符[22]。花青素的积累往往伴随着茶树整体次级代谢网络的系统性改变,并直接影响茶叶的风味品质。本研究证实,与绿色品种‘中茶108’相比,紫色品种(‘紫娟’)和季节性紫化品种(‘龙蕊1号’)均表现出更低的游离氨基酸含量和更高的总多酚及儿茶素含量。这导致了两者具有更高的“酚氨比”,该比值是决定茶叶适制性的关键[33]。‘龙蕊1号’的这一生化特性,决定了其在加工方向上更适于制作具有特殊花果香的特色红茶[9]。
非靶向代谢组学分析进一步揭示,‘龙蕊1号’的季节性代谢变化远不止花青素的积累。在其季节性差异代谢物中,单萜类、芪类及泛醌等多种次级代谢通路被特异性富集,这在‘紫娟’和‘中茶108’中并未出现。萜类化合物是构成茶叶香气的重要前体物质[34],其在夏季的特异性合成通路激活,可能赋予了‘龙蕊1号’夏茶独特的香气特征。这表明‘龙蕊1号’的遗传背景使其在响应季节变化时,启动了更为广泛和独特的代谢重编程,这不仅涉及色素的合成,也可能关乎其独特的香气风味和环境适应能力。
综上所述,本研究结果表明,‘龙蕊1号’的季节性紫化可能是由光照诱导的花青苷积累导致的。未来可通过转录组学(RNA-seq)分析,锁定在夏季‘龙蕊1号’中特异性高表达关键转录因子,并如反义寡核苷酸沉默[35]、转基因等手段,从分子层面确证其调控作用。其次,应对‘龙蕊1号’中特异性富集的萜类等香气物质进行靶向定量和香气活性值分析,并结合感官审评,明确其对茶叶最终风味的实际贡献。本研究不仅丰富了对茶树色素形成与环境互作的理论认知,也为培育兼具优良品质与环境适应性的特色茶树新品种提供了宝贵的种质资源和理论依据。

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2026年第47卷第18期
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doi: 10.13386/j.issn1002-0306.2025060101
  • 接收时间:2025-06-11
  • 首发时间:2026-10-24
  • 出版时间:2026-09-15
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  • 收稿日期:2025-06-11
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    广西南亚热带农业科学研究所,广西崇左 532415

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陈远权(1982−),男,硕士,高级农艺师,研究方向:茶树育种栽培与茶叶加工,E-mail:。
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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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