基于代谢组和转录组揭示不同颜色鲜食玉米花青素积累的分子机制

doi:10.7668/hbnxb.20196314

摘要/Abstract

摘要：

旨在基于代谢组与转录组联合分析,阐明不同颜色鲜食玉米籽粒花青素的组分差异及其分子调控机制,为彩色鲜食玉米分子育种提供候选基因与理论依据。以白色WN2000、黄色HN、彩色CN1和黑色HTN188为材料,综合采用代谢组学鉴定花青素相关代谢物并筛选差异代谢物,结合转录组测序获得差异表达基因(DEGs),通过GO注释与KEGG通路富集解析代谢—基因关联,并在花青素生物合成通路中挖掘关键结构基因与转录调控因子。结果表明,代谢组共鉴定49种花青素类代谢物,HTN188中矢车菊素、芍药素和天竺葵素等主要花青素显著积累,表现出更强的色素沉积特征。转录组共获得12 557个DEGs,其中F3H、DFR、UFGT等结构基因及MYB、bHLH等转录因子在高花青素品种中显著上调。相关性分析显示,基因Zm00001d011438的表达量与总花色素苷含量呈显著正相关,提示其可能在籽粒花青素积累中发挥关键作用。综上,基于代谢组与转录组联合分析,描述了不同颜色鲜食玉米籽粒花青素组成及相关基因表达差异,筛选出与花青素生物合成通路相关的结构基因、转录因子及候选基因Zm00001d011438,为后续功能验证与遗传改良研究提供参考。

关键词: 玉米, 花青素, 代谢组, 转录组, 结构基因

Abstract:

This study aimed to elucidate the compositional differences and molecular regulatory mechanisms of anthocyanins in kernels of fresh-eating maize with different colors by integrating metabolomic and transcriptomic analyses,thereby providing candidate genes and a theoretical basis for molecular breeding of colored fresh-eating maize.White WN2000,yellow HN,colored CN1,and black HTN188 were used as materials.Metabolomics was employed to identify anthocyanin-related metabolites and screen differential metabolites;transcriptome sequencing was conducted to obtain differentially expressed genes(DEGs).Metabolite gene associations were analyzed through GO annotation and KEGG pathway enrichment,and key structural genes and transcriptional regulators were mined within the anthocyanin biosynthetic pathway.The results showed that 49 anthocyanin-related metabolites were identified in total.Major anthocyanins,including cyanidin,peonidin,and pelargonidin,accumulated significantly in HTN188,indicating stronger pigment deposition.Transcriptome analysis identified 12 557 DEGs,among which structural genes such as F3H,DFR,and UFGT,as well as transcription factors including MYB and bHLH,were significantly up-regulated in varieties with high anthocyanin content.Correlation analysis revealed that the expression level of Zm00001d011438 was significantly and positively correlated with total anthocyanin content,suggesting that it may play a key role in kernel anthocyanin accumulation.Overall,the integrated metabolomic and transcriptomic analyses characterized differences in anthocyanin composition and related gene expression among differently colored fresh-eating maize kernels,and identified structural genes,transcription factors,and the candidate Zm00001d011438 associated with the anthocyanin biosynthetic pathway,providing references for subsequent functional validation and genetic improvement.

Key words: Maize, Anthocyanin, Metabolome, Transcriptome, Structural gene

中图分类号:

S513玉米(玉蜀黍)

蒋超, 闫立楠, 史涵博, 尹文旭, 栗慧, 刘松涛, 卢海博, 赵海超, 黄智鸿. 基于代谢组和转录组揭示不同颜色鲜食玉米花青素积累的分子机制[J]. 华北农学报, 2026, 41(4): 12-23. doi: 10.7668/hbnxb.20196314.

JIANG Chao, YAN Linan, SHI Hanbo, YIN Wenxu, LI Hui, LIU Songtao, LU Haibo, ZHAO Haichao, HUANG Zhihong. Elucidating the Molecular Mechanism of Anthocyanin Accumulation in Different-Colored Fresh Edible Corn Through Metabolomic and Transcriptomic Analyses[J]. Acta Agriculturae Boreali-Sinica, 2026, 41(4): 12-23. doi: 10.7668/hbnxb.20196314.

http://www.hbnxb.net/CN/Y2026/V41/I4/12

图/表 8

表1 qRT-PCR引物序列

Tab.1 qRT-PCR Primer sequence

基因 Gene	正向引物 (5'—3') Forward primer (5'—3')	反向引物 (5'—3') Reverse primer (5'—3')
Zm00001d017274	TGCTCTTCTTGGAGGGAAACG	AACGTGCTGGACTCACCTAA
Zm00001d028180	TGGTCTCAACGACACGATGG	CCCTCCATGACACAGGCAAT
Zm00001d014995	AGTCCATCATCGTCGCAGTC	TCGTTCTCGACCTCTCGTCT
Zm00001d034635	GCCAAAGAAGGGTCAGGTCA	TTTGAGGGTCCATTCCCGTG
Zm00001d044122	CGAGACAAACTCACCGGGAT	GAAAGGGAAGGTGCTGGTCA
Zm00001d032961	ATTCAACGGGGGAGGTAAGC	TGGGGACTGAAACGGTGATG
Zm00001d010521	ATTTAGTGGGCCTCGGTGTG	AACCTAATTGTGGGCCTGGG
Zm00001d047107	GGGCGATGAGCAACACATTC	GGACATCCCACAATGGAGCA
Zm00001d050455	TAGTAGTAGACCAGCGGCGT	ATGACGAAGCAAGTGGTGGT
Zm00001d026370	AAAGTCCAGTCCTGATCGCC	GGTACAGGACGCTGGATGAC
GADPH	GGTGGTGCTAAGCGTGTGAA	GCCAAATTCGTTGTCGTACC

图1 不同颜色玉米品种表型与花青素含量

Fig.1 Phenotype and anthocyanin content of different color maize varieties

图2 不同颜色玉米的代谢组分析 A.样品中检测出的花青素代谢物。B—D.3个比较组玉米花青素生物合成中的代谢物:B.HN vs WN2000;C.CN1 vs WN2000;D.HTN188 vs WN2000。

Fig.2 Metabolomic analysis of maize cultivars with different kernel colors A.Anthocyanin metabolites detected in samples.B—D.Differentially accumulated anthocyanins in three pairwise comparisons:B.HN vs WN2000;C.CN1 vs WN2000;D.HTN188 vs WN2000.

图3 GO注释 A.HN vs WN2000;B.CN1 vs WN2000;C.HTN188 vs WN2000。1.多细胞生物过程;2.生物种间相互作用的生物学过程;3.生殖过程;4.发展过程;5.本地化;6.对刺激的反应;7.生物调节;8.代谢过程;9.细胞过程;10.含蛋白质复合物;11.细胞解剖实体;12.分子传感器活性;13.抗氧化活性;14.分子功能调节活性;15.结构分子活性;16.ATP依赖性活性;17.转录调节因子活性;18.转运活性;19.催化活性;20.结合。

Fig.3 GO annotation A.HN vs WN2000;B.CN1 vs WN2000;C.HTN188 vs WN2000.1.Multicellular organismal process;2.Biological process involved in interspecies interaction between organisms;3.Reproductive process;4.Developmental process;5.Localization;6.Response to stimulus;7.Biological regulation;8.Metabolic process;9.Cellular process;10.Protein-containing complex;11.Cellular anatomical entity;12.Molecular transducer activity;13.Antioxidant activity;14.Molecular function regulator activity;15.Structural molecule activity;16.ATP-dependent activity;17.Transcription regulator activity;18.Transporter activity;19.Catalytic activity;20.Binding.

图4 KEGG富集 A.HN vs WN2000:1.丁酸酯代谢;2.β-丙氨酸代谢;3.谷胱甘肽代谢;4.精氨酸生物合成;5.丙氨酸、天冬氨酸和谷氨酸代谢;6.牛磺酸和次牛磺酸代谢;7.不饱和脂肪酸的生物合成;8.果糖和甘露糖代谢;9.精氨酸和脯氨酸代谢;10.花生四烯酸代谢;11.过氧化物酶体;12.光合生物中的碳固定;13.缬氨酸、亮氨酸和异亮氨酸降解;14.植物激素信号转导;15.氮代谢;16.角质、软脂和蜡的生物合成;17.脂肪酸生物合成;18.其他聚糖降解;19.淀粉和蔗糖代谢;20.蛋白质输出。 B.CN1 vs WN2000:1.谷胱甘肽代谢;2.果糖和甘露糖代谢;3.氨基糖和核苷酸糖代谢;4.丁酸酯代谢;5.光合生物中的碳固定;6.脂肪酸生物合成;7.角质、软脂和蜡的生物合成;8.苯并恶嗪类生物合成;9.植物激素信号转导;10.丙氨酸、天冬氨酸和谷氨酸代谢;11.缬氨酸、亮氨酸和异亮氨酸降解;12.卟啉代谢;13.丙酮酸代谢;14.柠檬酸盐循环(TCA循环);15.糖酵解/糖异生;16.甘油酯代谢;17.乙醛酸和二羧酸盐代谢;18.倍半萜类和三萜类生物合成;19.核黄素代谢;20.半乳糖代谢。 C.HTN188 vs WN2000:1.谷胱甘肽代谢;2.倍半萜类和三萜类生物合成;3.不饱和脂肪酸的生物合成;4.光合生物中的碳固定;5.果糖和甘露糖代谢;6.乙醛酸和二羧酸盐代谢;7.丁酸酯代谢;8.光合作用-天线蛋白;9.精氨酸和脯氨酸代谢;10.丙氨酸、天冬氨酸和谷氨酸代谢;11.氨基糖和核苷酸糖代谢;12.角质、软脂和蜡的生物合成;13.其他聚糖降解;14.精氨酸生物合成;15.甘油磷脂代谢;16.β-丙氨酸代谢;17.过氧化物酶体;18.淀粉和蔗糖代谢;19.甘油酯代谢;20.鞘糖脂生物合成-神经节。

Fig.4 KEGG enrichment A.HN vs WN2000:1.Butanoate metabolism;2.beta-Alanine metabolism;3.Glutathione metabolism;4.Arginine biosynthesis;5.Alanine,aspartate and glutamate metabolism;6.Taurine and hypotaurine metabolism;7.Biosynthesis of unsaturated fatty acids;8.Fructose and mannose metabolism;9.Arginine and proline metabolism;10.Arachidonic acid metabolism;11.Peroxisome;12.Carbon fixation in photosynthetic organisms;13.Valine,leucine and isoleucine degradation;14.Plant hormone signal transduction;15.Nitrogen metabolism;16.Cutin,suberine and wax biosynthesis;17.Fatty acid biosynthesis;18.Other glycan degradation;19.Starch and sucrose metabolism;20.Protein export.B.CN1 vs WN2000:1.Glutathione metabolism;2.Fructose and mannose metabolism;3.Amino sugar and nucleotide sugar metabolism;4.Butanoate metabolism;5.Carbon fixation in photosynthetic organisms;6.Fatty acid biosynthesis;7.Cutin,suberine and wax biosynthesis;8.Benzoxazinoid biosynthesis;9.Plant hormone signal transduction;10.Alanine,aspartate and glutamate metabolism;11.Valine,leucine and isoleucine degradation;12.Porphyrin metabolism;13.Pyruvate metabolism;14.Citrate cycle(TCA cycle);15.Glycolysis/Gluconeogenesis;16.Glycerolipid metabolism;17.Glyoxylate and dicarboxylate metabolism;18.Sesquiterpenoid and triterpenoid biosynthesis;19.Riboflavin metabolism;20.Galactose metabolism.C.HTN188 vs WN2000:1.Glutathione metabolism;2.Sesquiterpenoid and triterpenoid biosynthesis;3.Biosynthesis of unsaturated fatty acids;4.Carbon fixation in photosynthetic organisms;5.Fructose and mannose metabolism;6.Glyoxylate and dicarboxylate metabolism;7.Butanoate metabolism;8.Photosynthesis- antenna proteins;9.Arginine and proline metabolism;10.Alanine,aspartate and glutamate metabolism;11.Amino sugar and nucleotide sugar metabolism;12.Cutin,suberine and wax biosynthesis;13.Other glycan degradation;14.Arginine biosynthesis;15.Glycerophospholipid metabolism;16.beta-Alanine metabolism;17.Peroxisome;18.Starch and sucrose metabolism;19.Glycerolipid metabolism;20.Glycosphingolipid biosynthesis-ganglio series.

图5 玉米花青素生物合成途径基因分析 A.花青素生物合成途径及与结构酶基因相关DEGs的log₂(FPKM值)绘制热图。酶包括4-香豆酰辅酶A连接酶(4CL)、苯丙氨酸解氨酶(PAL)、反式肉桂酸-4-单氧化酶(C4H)、查尔酮合成酶(CHS)、黄烷酮3-羟化酶(F3H)、查尔酮异构酶(CHI)、类黄酮3-羟化酶(F3'H)、二氢黄酮醇还原酶(DFR)、黄酮醇合成酶(FLS)、udp-类黄酮葡萄糖基转移酶(UFGT)、花青素合成酶(ANS)、花青素还原酶(ANR)和无色花青素还原酶(LAR)。颜色梯度包括红色、白色和蓝色,分别代表上调、未调控和下调的基因。B.根据FPKM值分析花青素生物合成途径中所有DEGs的热图。

Fig.5 Analysis of DEGs in the anthocyanin biosynthesis pathway in maize A.Heatmap illustrating log₂(FPKM values)of DEGs involved in the anthocyanin biosynthesis pathway and structural enzyme genes.Enzymes include 4-coumarate-CoA ligase(4CL),phenylalanine ammonia-lyase(PAL),trans-cinnamate 4-monooxygenase(C4H),chalcone synthase(CHS),flavanone 3-hydroxylase(F3H),chalcone isomerase(CHI),flavonoid 3'-hydroxylase(F3'H),dihydroflavonol 4-reductase(DFR),flavonol synthase(FLS),UDP-flavonoid glucosyltransferase(UFGT),anthocyanidin synthase(ANS),anthocyanidin reductase(ANR),and leucoanthocyanidin reductase(LAR).The heatmap color gradient reflects gene regulation:red for upregulation,white for no change,and blue for downregulation.B.Heatmap of all DEGs in the anthocyanin biosynthesis pathway based on FPKM values.

图6 玉米花色素合成关键结构酶和转录因子相互作用网络

Fig.6 Interaction network of key structural enzyme genes and transcription factors(TFs)in maize anthocyanin biosynthesis

图7 qPCR 验证转录组结果

Fig.7 Transcriptome results verified by qPCR

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