‘朝阳椿’叶片类黄酮合成的转录组—代谢组联合分析

doi:10.16420/j.issn.0513-353x.2025-0011

摘要/Abstract

摘要：

以‘朝阳椿’臭椿（Ailanthus altissima‘Chaoyang’）和普通臭椿（A. altissima）的上、中、下3个叶位的叶片为试材，采用超高效液相色谱串联质谱（UPLC-MS/MS）和高通量RNA测序技术，分别测定类黄酮代谢组和转录组。分析两种臭椿不同叶位叶片中起呈色作用的类黄酮组分及其含量变化，并运用皮尔森相关性分析法挖掘关键类黄酮代谢物的潜在调控基因。结果表明，2种查尔酮代谢物和4种黄酮醇代谢物为直接促使‘朝阳椿’叶片呈现黄色的关键类黄酮代谢物，2个CHS和1个PGT1是调控查尔酮代谢物的关键候选结构基因，2个FLS、1个FG2和1个FG3是催化合成黄酮醇代谢物的关键候选结构基因；矢车菊素-3-O-芸香糖苷是导致两种臭椿上叶位叶片比中、下叶位更红、且‘朝阳椿’叶片较普通臭椿更加红艳的关键花色苷，2个CHS、3个F3H、2个DFR和1个CHI为正向调控矢车菊素-3-O-芸香糖苷合成的关键候选结构基因，编码转录因子的调节基因（2个MYB和1个bHLH）极可能对这些正调控结构基因具有激活作用，而1个MYB和1个bHLH极可能起抑制作用。

关键词: 臭椿, 叶色, 类黄酮, 代谢组, 转录组

Abstract:

The leaves from the upper，middle，and lower positions of Ailanthus altissima‘Chaoyang’and common A. altissima were utilized as experimental materials in the study. Ultra- performance liquid chromatography-tandem mass spectrometry（UPLC-MS/MS）and high-throughput RNA sequencing were employed to determine the flavonoid metabolome and transcriptome，respectively. The flavonoid components and their content changes responsible for coloration in the leaves of the two types of A. altissima at different leaf positions were analyzed. Pearson correlation analysis was used to identify potential regulatory genes of key flavonoid metabolites. The results indicated that two chalcone metabolites and four flavonol metabolites were the key flavonoids directly responsible for the yellow coloration in‘Chaoyang’leaves. Two CHSs and one PGT1 were identified as key candidate structural genes regulating chalcone metabolites，while two FLSs，one FG2，and one FG3 were found to be key candidate structural genes catalyzing the synthesis of flavonol metabolites. Cyanidin-3-O-rutinoside was identified as the key anthocyanin that makes the upper leaves of both types of A. altissima redder than the middle and lower leaves，and‘Chaoyang’leaves redder than those of common A. altissima. Two CHSs，three F3Hs，two DFRs，and one CHI were the main candidate structural genes that positively regulated the synthesis of cyanidin-3-O-rutinoside. Regulatory genes encoding transcription factors（two MYBs and one bHLH）are highly likely to activate these above-mentioned positively regulating structural genes，while one MYB and one bHLH are extremely likely to have inhibitory effects.

Key words: Ailanthus altissima, leaf color, flavonoid, metabolome, transcriptome

郑聪慧, 徐振华, 张蔓蔓, 王玉忠, 李向军, 杜克久, 孙然, 李亚. ‘朝阳椿’叶片类黄酮合成的转录组—代谢组联合分析[J]. 园艺学报, 2025, 52(11): 2913-2930.

ZHENG Conghui, XU Zhenhua, ZHANG Manman, WANG Yuzhong, LI Xiangjun, DU Kejiu, SUN Ran, LI Ya. Integrated Transcriptomic and Metabolomic Analysis of Flavonoid Biosynthesis in the Leaves of Ailanthus altissima‘Chaoyang’[J]. Acta Horticulturae Sinica, 2025, 52(11): 2913-2930.

图/表 16

图1 ‘朝阳椿’（A）和普通臭椿（B）的一年生苗木

Fig. 1 One-year-old clonal seedlings of Ailanthus altissima‘Chaoyang’（A）and common A. altissima（B）

表1 叶片颜色描述

Table 1 Summary of leaf color

材料 Material	叶位 Leaf position	样品组代号 Sample group	颜色 Color	颜色代码 Color code
朝阳椿 Chaoyang	1	Y1	中度红橙Moderate reddish orange	N34D
	2	Y2	浅绿黄Light greenish yellow	6D
	3	Y3	亮黄绿Brilliant yellow green	149B
普通臭椿 Common Ailanthus altissima	1	G1	中度橄榄棕色Moderate olive brown	199A
	2	G2	中绿Moderate green	131C
	3	G3	深绿Dark green	136A

图2 PCA得分图 Y：‘朝阳椿’；G：普通臭椿；1、2、3表示叶位。下同

Fig. 2 PCA score plot Y：Ailanthus altissima‘Chaoyang’；G：Common Ailanthus altissima. 1，2，3 indicated leaf position. The same below.

表2 差异代谢物和差异基因的数量统计

Table 2 Statistical table of differential metabolites and differentially expressed genes

类别 Type	对照组—试验组 Control group— test group	差异代谢物数 Number of differential metabolites		差异基因数 Number of differentially expressed genes
类别 Type	对照组—试验组 Control group— test group	下调 Down-regulated	上调 Up-regulated	下调 Down-regulated	上调 Up-regulated
朝阳椿 Chaoyang	Y1-Y2	50	63	4 266	5 617
	Y1-Y3	69	67	12 219	11 431
	Y2-Y3	39	35	6 763	4 828
普通臭椿 Common Ailanthus altissima	G1-G2	42	117	7 968	8 265
	G1-G3	55	112	13 997	10 841
	G2-G3	56	58	7 932	6 036
普通臭椿和‘朝阳椿’ Common A. altissima and Chaoyang	G1-Y1	51	102	4 969	4 558
	G2-Y2	55	57	6 720	8 249
	G3-Y3	62	55	7 970	10 096

图3 差异代谢物韦恩图

Fig. 3 Veen diagram of differential metabolites

图4 差异代谢物聚类热图

Fig. 4 Cluster heat maps of differential metabolites

表3 基因注释结果

Table 3 Results of gene annotation

数据库 Database	基因数 Number of genes	百分比/% Percentage
KEGG	68 246	41.33
Nr	93 228	56.45
SwissProt	66 292	40.14
TrEMBL	92 691	56.13
KOG	55 423	33.56
GO	77 384	46.86
Pfam	60 460	36.61
至少在一个数据库中有注释的基因 Annotated in at least one database	94 616	57.30
基因总数 Total unigenes	165 137	100.00

图5 差异结构基因韦恩图

Fig. 5 Veen diagram of differentially expressed structual genes

图6 与类黄酮合成相关的差异结构基因热图红色字体表示两种臭椿的共有基因

Fig. 6 Heat map of differentially expressed flavonoid related structual genes Genes in red letters represent the common genes of two types of Ailanthus altissima

图7 与花青素合成相关的差异转录因子编码基因聚类热图

Fig. 7 Cluster heat map of differentially expressed anthocyanidin-related TF encoding genes

图8 类黄酮（花青素）差异代谢物和差异结构基因的相关性网络图圆圈为差异表达结构基因，菱形为差异代谢物，橘色线为两者存在正相关关系，绿色线为两者存在负相关关系。* 为同分异构体。下同

Fig. 8 Correlation network of differential metabolites of flavonoids（anthocyanidins）and differentially expressed structural genes Circles represent differentially expressed structual genes，rhombuses represent differential metabolites，orange solid lines indicate a positive correlation，and green solid lines indicate a negative correlation. * indicate isomers. The same below

表4 类黄酮（花青素）关键差异代谢物和关键差异表达基因

Table 4 Key differential metabolites and key differentially expressed genes of flavonoids（anthocyanidins）

类目 Category	代谢物 Metabolites	结构基因 Structual genes	转录因子基因 TF encoding genes
花青素 Anthocyanidins	矢车菊素-3-O-芸香糖苷 Cyanidin-3-O-rutinoside；飞燕草素-3-O-葡萄糖苷^* Delphinidin-3-O-glucoside	CHS-5；CHS-8；F3H-1；F3H-2；F3H-3；DFR-2；DFR-1；CHI-6	MYB-4；bHLH-2；MYB-6；MYB-5；bHLH-3
查尔酮Chalcones	柚皮素查尔酮 Naringenin chalcone；根皮苷 Phlorizin	CHS5；CHS8；PGT1-1	-
黄酮醇 Flavonols	紫云英苷* Astragalin；异槲皮苷 Isoquercitrin；芦丁* Rutin；白麻苷 Baimaside	FLS-1；FLS-2；FG2-1；FG3-2	-

图9 转录因子编码基因与关键花色苷的相关性网络图三角形为差异表达的转录因子编码基因。下同

Fig. 9 Correlation network of TF encoding genes and key anthocyanidins Triangles represent differentially expressed TF-encoding genes. The same below

图10 转录因子编码基因与花青素相关结构基因的相关性网络图

Fig. 10 Correlation network of TF encoding genes and anthocyanidin-related structural genes

图11 ‘朝阳椿’和普通臭椿叶片类黄酮（花青素）合成通路灰色字体代谢物为本研究中未检测到的代谢物

Fig. 11 Flavonoid （anthocyanin）synthesis pathway in leave of‘Chaoyang’and common Ailanthus altissima Metabolites in gray font represent the metabolites not detected in this study

图12 关键基因 qPCR 验证

Fig. 12 Validation of key genes by qPCR

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