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热带作物种质资源种类繁多,分布在世界138个热带国家或地区,其中94个属于典型热带国家或地区[1]。这些国家绝大部分是发展中国家,主要以农业人口为主,且热带农业产业是其支柱产业,从事热带农业产业的直接和间接相关人口有10亿左右,世界上约有七分之一人口的生活状况与热带农业的生产水平有直接关系[2]。重要的热带作物包括椰子(Cocos nucifera L.)、橡胶树(Hevea brasiliensis)、木薯(Manihot esculenta Crantz)、椰枣(Phoenix dactylifera L.
)、可可(Theobroma cacao L.)、油棕(Elaeis guineensis Jacq.)和咖啡(Coffea arabica L. )等,这些作物富含维生素、氨基酸、矿物质和抗氧化剂等营养成分,热带作物在人类生活中发挥着重要的作用,并对全球经济、医药和生态平衡产生着深远的影响。丰富的热带药用植物资源对恶性肿瘤、白血病、肝炎等疑难杂症的治疗有其独特的疗效[3]。重要的药用植物包括海南粗榧(Cephalotaxus hainanensis Li)、益智(Alpinia oxyphylla Miq.)、槟榔(Areca catechu L.)、胡椒(Piper nigrum L.)、砂仁(Amomum villosum Lour.)、巴戟天(Morinda officinalis How.)、藿香[Agastache rugosa (Fisch.et.Mey.) O.Kuntze]、铁皮石斛(Dendrobium officinale Kimura et Migo )和千年健[Homalomena occulta ( Lour.) Schott.]等,其中从海南粗榧中提取的三尖杉酯类生物碱具有广谱抗癌作用[4];从胡椒中提取的胡椒碱为主的酰胺类生物碱,具有抗炎镇痛、抗肿瘤、抗惊厥等作用[5];从槟榔中提取的活性成分对消化、神经和心血管系统产生影响,具有抗寄生虫、抗氧化等作用[6]。因此从植物中发现新型的有效活性成分或者药物前体化合物是开发和革新天然药物的有效手段[7]。虽然中国拥有丰富的热带作物和药用植物资源宝库,但是大部分的植物尚未得到充分的研究、广泛的开发和有效的利用。笔者对热带植物中的主要化学成分及代谢物研究现状进行了总结,旨在为热带植物的代谢生物学研究及进一步应用开发提供参考。 -
代谢组学利用高通量的检测技术,可以有效分析植物体中代谢物的构成,是系统生物学不可或缺的组成部分,在各类研究领域发挥着越来越重要的作用。热带植物因其独特的生长环境,含有各类营养物质和具有活性的次生代谢物,这些物质成分对人体具有重要的营养和药用价值,虽然一些重要的植物已经有了一定研究进展(表1),但是大部分的植物还未得到充分的研究与开发利用。
表 1 热带植物代谢组学及多组学的相关研究
植物 已检测代谢物 检测技术 多组学研究进展 参考文献 椰子(Cocos nucifera L.) 脂质、酯类、氨基酸、有机酸、维生素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [35-40] 橡胶树(Hevea brasiliensis) 萜类、有机酸、脂质、核苷酸、氨基酸、木质素、类黄酮、香豆素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [42-46] 油棕( Elaeis guineensis Jacq.) 脂质、氨基酸、有机酸、胺类 LC-MS、GC-MS 基因组、转录组、蛋白质组 [49-54] 木薯(Manihot esculenta Crantz) 有机酸、氨基酸、类胡萝卜素、叶绿素、生育酚 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [58-63] 槟榔(Areca catechu L.) 生物碱、类黄酮、丹宁、三萜、类固醇、脂质、鞣质 LC-MS、GC-MS、NMR 基因组、转录组 [66-69] 胡椒(Piper nigrum L.) 生物碱、皂苷、有机酸、香豆素、类黄酮、生物碱、内酯、酚类 LC-MS、GC-MS、NMR 基因组、转录组 [73-77] 海南粗榧(Cephalotaxus hainanensis Li) 三尖杉类生物碱、1-苯乙基异喹啉 LC-MS、GC-MS 转录组 [82-84] 砂仁(Amomum villosum Lour.) 挥发油、多酚、多糖、有机酸、类黄酮 LC-MS、GC-MS、NMR 转录组、蛋白质组 [87-92] 甘蔗(Saccharum officinarum L.) 类黄酮、有机酸、脂肪酸、糖类、木质素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [93-96] 油梨(Persea americana Mill.) 氨基酸、脂肪酸、植物甾醇、生育酚、角鲨烯 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [97-101] 香蕉(Musa nana Lour.) 酚类、类胡萝卜素、脂肪酸、植物甾醇、胺类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [102-104] 腰果(Anacardium occidentalie L.) 维生素、脂质、有机酸、苯丙烷类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [105-108] 可可(Theobroma cacao L.) 类黄酮、氨基酸、醇类、有机酸、酯类、醛类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [109-112] 咖啡(Coffea arabica L.) 多酚、生物碱、糖类、脂肪酸、有机酸、醇类、氨基酸、萜类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [113-117] 龙眼(Dimocarpus longan Lour.) 类黄酮、氨基酸、生物碱、单宁、酚酸、有机酸、核苷酸、脂质 LC-MS、GC-MS 基因组、转录组、蛋白质组 [118-122] 番木瓜(Carica papaya L.) 氨基酸、维生素、植物激素、有机酸、醇类、酯类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [123-126] 菠萝[Ananas comosus (L.) Merr.] 氨基酸、脂类、类黄酮、糖类、糖苷 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [127-130] 椰枣(Phoenix dactylifera L.) 氨基酸、脂肪酸、维生素、糖类 LC-MS、GC-MS、NMR 基因组、转录组 [131-133] 利用基因组、转录组、蛋白质组等组学原理与方法,研究热带植物种质资源的遗传多样性及其地理分布,阐明野生品种、地方品种和现代栽培品种的进化规律具有重要意义。综合目前的研究情况,笔者认为基于代谢组学的热带植物未来主要的研究方向包括:1.解析重要热带作物营养风味物质的成分;2.探究重要药用活性物质的组成,及其合成代谢途径的解析;3.阐明代谢物合成途径的调控机理,探究进一步提高热带作物产量的策略;4.进一步将代谢组学与其他组学进行整合分析,以便更全面地阐述代谢物生物合成的分子机理。
Research progress of important tropical plants based on metabolomics
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摘要: 代谢组学是继基因组学、转录组学、蛋白组学之后发展起来的一门新兴的组学技术,近年来已经在植物研究领域中显现其重要作用,同样植物代谢组学的发展也推进了热带植物的深入研究。本文主要综述了植物代谢组学的发展进程及其在重要热带作物[椰子(Cocos nucifera L.)、橡胶树(Hevea brasiliensis)、油棕(Elaeis guineensis Jacq.)、木薯(Manihot esculenta Crantz
)]和药用植物[槟榔(Areca catechu L.)、胡椒(Piper nigrum L.)、海南粗榧(Cephalotaxus hainanensis Li)、砂仁(Amomum villosum Lour.)]中的综合运用,并介绍了结合多组学技术开展相关研究的进展,最后,对未来的研究方向提出了展望,旨在为进一步的热带植物代谢生物学研究及其开发利用提供参考。 Abstract: Metabolomics is an emerging omics technology developed after genomics, transcriptomics and proteomics, and has shown its important role in plant research in recent years, and likewise the development of plant metabolomics has advanced the in-depth study of tropical plants. A review was made mainly of the development of plant metabolomics and its integrated application in important tropical crops (Cocos nucifera L., Hevea brasiliensis, Elaeis guineensis Jacq., Manihot esculenta Crantz) and medicinal plants (Areca catechu L., Piper nigrum L., Cephalotaxus hainanensis Li, Amomum villosum Lour.), and recent advances in research of tropical plants based on multi-omics technologies were also reviewed. An outlook for further research based on omics technologies was put forward to provide a reference for further research and exploitation of metabolic biology of tropical plants.-
Key words:
- tropical plant /
- GC-MS /
- LC-MS /
- metabolomics /
- multi-omics
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表 1 热带植物代谢组学及多组学的相关研究
植物 已检测代谢物 检测技术 多组学研究进展 参考文献 椰子(Cocos nucifera L.) 脂质、酯类、氨基酸、有机酸、维生素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [35-40] 橡胶树(Hevea brasiliensis) 萜类、有机酸、脂质、核苷酸、氨基酸、木质素、类黄酮、香豆素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [42-46] 油棕( Elaeis guineensis Jacq.) 脂质、氨基酸、有机酸、胺类 LC-MS、GC-MS 基因组、转录组、蛋白质组 [49-54] 木薯(Manihot esculenta Crantz) 有机酸、氨基酸、类胡萝卜素、叶绿素、生育酚 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [58-63] 槟榔(Areca catechu L.) 生物碱、类黄酮、丹宁、三萜、类固醇、脂质、鞣质 LC-MS、GC-MS、NMR 基因组、转录组 [66-69] 胡椒(Piper nigrum L.) 生物碱、皂苷、有机酸、香豆素、类黄酮、生物碱、内酯、酚类 LC-MS、GC-MS、NMR 基因组、转录组 [73-77] 海南粗榧(Cephalotaxus hainanensis Li) 三尖杉类生物碱、1-苯乙基异喹啉 LC-MS、GC-MS 转录组 [82-84] 砂仁(Amomum villosum Lour.) 挥发油、多酚、多糖、有机酸、类黄酮 LC-MS、GC-MS、NMR 转录组、蛋白质组 [87-92] 甘蔗(Saccharum officinarum L.) 类黄酮、有机酸、脂肪酸、糖类、木质素 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [93-96] 油梨(Persea americana Mill.) 氨基酸、脂肪酸、植物甾醇、生育酚、角鲨烯 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [97-101] 香蕉(Musa nana Lour.) 酚类、类胡萝卜素、脂肪酸、植物甾醇、胺类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [102-104] 腰果(Anacardium occidentalie L.) 维生素、脂质、有机酸、苯丙烷类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [105-108] 可可(Theobroma cacao L.) 类黄酮、氨基酸、醇类、有机酸、酯类、醛类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [109-112] 咖啡(Coffea arabica L.) 多酚、生物碱、糖类、脂肪酸、有机酸、醇类、氨基酸、萜类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [113-117] 龙眼(Dimocarpus longan Lour.) 类黄酮、氨基酸、生物碱、单宁、酚酸、有机酸、核苷酸、脂质 LC-MS、GC-MS 基因组、转录组、蛋白质组 [118-122] 番木瓜(Carica papaya L.) 氨基酸、维生素、植物激素、有机酸、醇类、酯类 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [123-126] 菠萝[Ananas comosus (L.) Merr.] 氨基酸、脂类、类黄酮、糖类、糖苷 LC-MS、GC-MS、NMR 基因组、转录组、蛋白质组 [127-130] 椰枣(Phoenix dactylifera L.) 氨基酸、脂肪酸、维生素、糖类 LC-MS、GC-MS、NMR 基因组、转录组 [131-133] -
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