冬枣果皮着色相关类黄酮靶向代谢组学及潜在MYB转录因子分析

doi:10.16420/j.issn.0513-353x.2021-0326

摘要/Abstract

摘要：

以冬枣白熟期（W）、半红期（着色部分SR_R,未着色部分SR_W）、全红期（R）果皮为试材,通过靶向代谢组学分析不同着色期冬枣果皮类黄酮组分及变化,构建冬枣MYB家族进化树,通过转录组分析潜在调控类黄酮合成MYB转录因子。结果表明,着色期4个比较组果皮差异代谢物分别为291（R vs W）个、196（R vs SR_W）个、297（SR_R vs SR_W）个和201（SR_R vs W）,其中共有差异代谢物98个。通过KEGG分析发现共有的差异代谢物主要富集在次生代谢物、苯丙烷类和类黄酮生物合成途径;126个类黄酮物质在果皮着色期被检测到,其中包括13个花青苷类物质、53个黄酮物质、28个黄酮醇类物质、8个异黄酮、5个原花青素和19个黄烷醇类物质;随着冬枣果皮着色,3个花翠素、3个黄酮和3个黄酮醇类物质显著增加,而原花青素和黄烷醇类物质儿茶素、表儿茶素显著减少。冬枣MYB家族聚类共筛选出14个潜在调控类黄酮生物合成的MYB转录因子。进一步通过转录组和qRT-PCR分析发现有12个MYB转录因子随冬枣果皮着色呈下调表达趋势,仅LOC107434709和LOC107404750两个MYB转录因子在着色冬枣果皮中表达水平较高,暗示其可能在冬枣着色过程中发挥积极作用。

关键词: 枣, 着色期, 类黄酮, MYB转录因子

Abstract:

In our study,the white（W）,semi-red（SR_R,SR_W）and full-red（R）pericarps of winter jujube were used as materials,we analyzed components and changes of flavonoids in different coloring periods using targeted metabolomics. The phylogenetic tree of MYB family in winter jujube was constructed,and the potential MYB regulators associated with flavonoid synthesis were analyzed using transcriptomic analysis. The results showed that the differential metabolites between colored and uncolored pericarps were 291（R vs W）,196（R vs SR_W）,297（SR_R vs SR_W）,and 201（SR_R vs W）,respectively. There were 98 differential metabolites in the four comparison groups. Further analysis of KEGG revealed that the shared differential metabolites of four comparison groups mainly enriched in secondary metabolites,phenylpropanoids and flavonoid biosynthetic pathways. The 126 flavonoids were detected during the coloring period of winter jujube pericarp,including thirteen anthocyanins,fifty-three flavones,twenty-eight flavonols,eight isoflavones,five proanthocyanidins and nineteen flavanols. Along with the coloration of winter jujube pericarp,three anthocyanins,three flavones and three flavonols increased significantly,while procyanidins and flavanols（catechin,epicatechin）decreased significantly. Fourteen potential MYB regulators associated with flavonoid synthesis were screened via cluster analysis of MYB family. The analyses of transcriptome and qRT-PCR showed that twelve MYB transcription factors were down-regulated with the coloration of winter jujube pericarp,while LOC107434709 and LOC107404750 had higher expression levels in colored pericarp of winter jujube. This suggested that they might play positive roles in the coloring process of winter jujube.

Key words: jujube, coloring period, flavonoid, MYB transcription factors

中图分类号:

S665.1

许海峰, 王中堂, 陈新, 刘志国, 王利虎, 刘平, 刘孟军, 张琼. 冬枣果皮着色相关类黄酮靶向代谢组学及潜在MYB转录因子分析[J]. 园艺学报, 2022, 49(8): 1761-1771.

XU Haifeng, WANG Zhongtang, CHEN Xin, LIU Zhiguo, WANG Lihu, LIU Ping, LIU Mengjun, ZHANG Qiong. The Analyses of Target Metabolomics in Flavonoid and Its Potential MYB Regulation Factors During Coloring Period of Winter Jujube[J]. Acta Horticulturae Sinica, 2022, 49(8): 1761-1771.

图/表 5

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引物名称 Primer name	引物序列（5′-3′） Primer sequence
LOC107434532	F：CCCAAAACAACCCTGCAAC;R：AGACTAACGCTCCTTCGACCA
LOC107406469	F：AAACGGCTACCACATCCA;R：CACCAGACTTGCCCTCCA
LOC107434709	F：AAGCTTTGATCACGGTTGG;R：TTAGGTCCCATGAACTCCG
LOC107404750	F：CGGAACTTATGCTGGACAAT;R：AGAAGAGGAACCAACGCAAA
LOC107426233	F：AGTGGGAAGGTTGTGGATAG;R：CTTTTCCCAAAAATGAACAAG
LOC107407717	F：ACCCTAACGGGAAAGACGA;R：GGAGCCTGAGAACAAGATGAA
LOC107433002	F：GATAAGCATTGCAAGCCGGG;R：TCCGAACTGGCTTTAGCTGG
LOC107415546	F：CTCCCGAAGGATCACCAAGG;R：GCAGTCCCAGTCATCAACGA
LOC107423566	F：CGCTCGTCCTCAAGCTAATC;R：GAAGAAAGCTGCGGTTCCTA
LOC107429773	F：AGAGAGCGGGTACAGATGGT;R：CAGGGCCAACATGTGCTAGA
LOC107423630	F：ACACGGGGAGGTAGTGACAA;R：CCTCCAATGGATCCTCGTTA
LOC107414848	F：TAGCCATACCTCTCATTGGCAG;R：TCATAGTACTGTGTCCCCATGA
LOC107420842	F：GTTCCTCAACTTAAAGGAAATG;R：ACAACGTTCCGTAGCAGAAAC
LOC107431136	F：CAAAGGCAGCCACATCAGG;R：ACACCAGCTAGTGTTTCTCC

引物名称 Primer name	引物序列（5′-3′） Primer sequence
LOC107434532	F：CCCAAAACAACCCTGCAAC;R：AGACTAACGCTCCTTCGACCA
LOC107406469	F：AAACGGCTACCACATCCA;R：CACCAGACTTGCCCTCCA
LOC107434709	F：AAGCTTTGATCACGGTTGG;R：TTAGGTCCCATGAACTCCG
LOC107404750	F：CGGAACTTATGCTGGACAAT;R：AGAAGAGGAACCAACGCAAA
LOC107426233	F：AGTGGGAAGGTTGTGGATAG;R：CTTTTCCCAAAAATGAACAAG
LOC107407717	F：ACCCTAACGGGAAAGACGA;R：GGAGCCTGAGAACAAGATGAA
LOC107433002	F：GATAAGCATTGCAAGCCGGG;R：TCCGAACTGGCTTTAGCTGG
LOC107415546	F：CTCCCGAAGGATCACCAAGG;R：GCAGTCCCAGTCATCAACGA
LOC107423566	F：CGCTCGTCCTCAAGCTAATC;R：GAAGAAAGCTGCGGTTCCTA
LOC107429773	F：AGAGAGCGGGTACAGATGGT;R：CAGGGCCAACATGTGCTAGA
LOC107423630	F：ACACGGGGAGGTAGTGACAA;R：CCTCCAATGGATCCTCGTTA
LOC107414848	F：TAGCCATACCTCTCATTGGCAG;R：TCATAGTACTGTGTCCCCATGA
LOC107420842	F：GTTCCTCAACTTAAAGGAAATG;R：ACAACGTTCCGTAGCAGAAAC
LOC107431136	F：CAAAGGCAGCCACATCAGG;R：ACACCAGCTAGTGTTTCTCC

类黄酮 Flavonoid	差异倍数 Log₂ fold change
类黄酮 Flavonoid	SR_R vs W R vs W SR_R vs SR_W R vs SR_W
花青苷成分 Anthocyanin component 花翠素Delphinidin	—	—	—	1.32
花翠素 3-O-葡萄糖苷Delphinidin 3-O-glucoside	—	—	—	1.13
二甲花翠素 3-O-葡萄糖苷Malvidin 3-O-glucoside	1.33	1.04	—	—
花翠素 3-O-芸香糖苷Delphinidin 3-O-rutinoside	—	—	—	—
矢车菊素Cyanidin	—	—	—	—
矢车菊素 3-O-葡萄糖苷Cyanidin 3-O-glucoside	-1.38	—	-1.35	—
矢车菊素 3,5-O-二葡萄糖苷Cyanidin 3,5-O-diglucoside	—	—	—	—
矢车菊素 3-O-丙二酰己糖苷Cyanidin 3-O-malonylhexoside	-1.68	-1.64	—	—
矢车菊素半乳糖苷Cyanidin 3-O-galactoside	—	—	-1.28	—
芍药花青素Peonidin	—	—	—	—
芍药花青素 O-己糖苷Peonidin O-hexoside	—	—	—	—
芍药-3-O-葡萄糖甙氯化物Peonidin 3-O-glucoside chloride	—	—	—	—
天竺葵色素苷Pelargonin	—	—	—	—
黄酮/黄酮醇成分 Flavone/flavonol component 8-C-己糖苷-木犀草素 O-己糖苷 8-C-hexosyl-luteolin O-hexoside	1.35	1.48	—	—
C-己糖基-木犀草素 C-戊糖苷 C-hexosyl-luteolin C-pentoside	1.46	1.48	—	—
木犀草素Luteolin	1.20	1.02	1.41	1.23
异鼠李素 5-O-己糖苷Isorhamnetin 5-O-hexoside	2.12	3.56	2.17	3.60
异鼠李素 O-己糖苷Isorhamnetin O-hexoside	2.25	3.52	2.21	3.49
异鼠李素-3-O-葡萄糖苷Isorhamnetin 3-O-glucoside	2.25	3.48	—	—
原花青素/黄烷醇成分 Procyanidin/Flavanol component 原花青素A1 Procyanidin A1	—	—	-2.96	-3.97
原花青素A2 Procyanidin A2	—	1.25	—	—
原花青素A3 Procyanidin A3	1.18	1.74	-1.38	—
原花青素B2 Procyanidin B2	-1.16	-1.57	—	-1.13
原花青素B3 Procyanidin B3	-1.01	-1.52	—	-1.21
儿茶素Catechin	-2.67	-3.91	-1.61	-2.85
表儿茶素 L-Epicatechin	-1.61	-3.43	-1.13	-2.95

类黄酮 Flavonoid	差异倍数 Log₂ fold change
类黄酮 Flavonoid	SR_R vs W R vs W SR_R vs SR_W R vs SR_W
花青苷成分 Anthocyanin component 花翠素Delphinidin	—	—	—	1.32
花翠素 3-O-葡萄糖苷Delphinidin 3-O-glucoside	—	—	—	1.13
二甲花翠素 3-O-葡萄糖苷Malvidin 3-O-glucoside	1.33	1.04	—	—
花翠素 3-O-芸香糖苷Delphinidin 3-O-rutinoside	—	—	—	—
矢车菊素Cyanidin	—	—	—	—
矢车菊素 3-O-葡萄糖苷Cyanidin 3-O-glucoside	-1.38	—	-1.35	—
矢车菊素 3,5-O-二葡萄糖苷Cyanidin 3,5-O-diglucoside	—	—	—	—
矢车菊素 3-O-丙二酰己糖苷Cyanidin 3-O-malonylhexoside	-1.68	-1.64	—	—
矢车菊素半乳糖苷Cyanidin 3-O-galactoside	—	—	-1.28	—
芍药花青素Peonidin	—	—	—	—
芍药花青素 O-己糖苷Peonidin O-hexoside	—	—	—	—
芍药-3-O-葡萄糖甙氯化物Peonidin 3-O-glucoside chloride	—	—	—	—
天竺葵色素苷Pelargonin	—	—	—	—
黄酮/黄酮醇成分 Flavone/flavonol component 8-C-己糖苷-木犀草素 O-己糖苷 8-C-hexosyl-luteolin O-hexoside	1.35	1.48	—	—
C-己糖基-木犀草素 C-戊糖苷 C-hexosyl-luteolin C-pentoside	1.46	1.48	—	—
木犀草素Luteolin	1.20	1.02	1.41	1.23
异鼠李素 5-O-己糖苷Isorhamnetin 5-O-hexoside	2.12	3.56	2.17	3.60
异鼠李素 O-己糖苷Isorhamnetin O-hexoside	2.25	3.52	2.21	3.49
异鼠李素-3-O-葡萄糖苷Isorhamnetin 3-O-glucoside	2.25	3.48	—	—
原花青素/黄烷醇成分 Procyanidin/Flavanol component 原花青素A1 Procyanidin A1	—	—	-2.96	-3.97
原花青素A2 Procyanidin A2	—	1.25	—	—
原花青素A3 Procyanidin A3	1.18	1.74	-1.38	—
原花青素B2 Procyanidin B2	-1.16	-1.57	—	-1.13
原花青素B3 Procyanidin B3	-1.01	-1.52	—	-1.21
儿茶素Catechin	-2.67	-3.91	-1.61	-2.85
表儿茶素 L-Epicatechin	-1.61	-3.43	-1.13	-2.95