紫叶甘蓝型油菜花青素合成的转录组-代谢组联合分析

doi:10.7668/hbnxb.20194754

华北农学报 ›› 2024, Vol. 39 ›› Issue (3): 25-34. doi: 10.7668/hbnxb.20194754

所属专题： 油料作物；生物技术；

• 作物遗传育种·种质资源·生物技术 • 上一篇下一篇

紫叶甘蓝型油菜花青素合成的转录组-代谢组联合分析

周弘峰¹, 朱思颖¹, 贺丹¹, 刘丽莉¹^,²^,³, 陈道宗⁴, 谭晨⁴, 张大为¹^,²^,³, 严明理²^,³

¹ 经济作物遗传改良与综合利用湖南省重点实验室,湖南科技大学生命科学与健康学院,湖南湘潭 411201
² 岳麓山实验室,湖南长沙 410128
³ 油菜杂种优势利用湖南省工程研究中心,湖南省作物研究所,湖南长沙 410125
⁴ 赣南师范大学生命科学学院,江西赣州 341000

收稿日期:2023-12-22 出版日期:2024-06-28
通讯作者:
张大为(1988—),男,湖北枣阳人,博士,硕士生导师,主要从事油菜遗传育种研究。
严明理(1979—),男,湖南邵阳人,研究员,博士,博士生导师,主要从事油菜遗传与分子生物学研究。
作者简介:
周弘峰(1999—),男,湖南株洲人,硕士,主要从事油菜遗传育种研究。
基金资助:
国家自然科学基金项目(U19A2029); 湖南省自然科学基金项目(2023JJ40279); 湖南省教育厅优秀青年项目(21B0490)

Transcriptomic and Metabolomic Analysis of Anthocyanin Synthesis in Purple Leaf Brassica napus

ZHOU Hongfeng¹, ZHU Siying¹, HE Dan¹, LIU Lili¹^,²^,³, CHEN Daozong⁴, TAN Chen⁴, ZHANG Dawei¹^,²^,³, YAN Mingli²^,³

¹ Hunan Key Laboratory of Economic Crops Genetic Improvement and Integrated Utilization,School of Life and Health Sciences,Hunan University of Science and Technology,Xiangtan 411201,China
² Yuelushan Laboratory,Changsha 410128,China
³ Hunan Research Center of Heterosis Utilization in Rapeseed,Crop Research Institute of Hunan Province,Changsha 410125,China
⁴ School of Life Sciences and Health,Gannan Normal University,Ganzhou 341000,China

Received:2023-12-22 Published:2024-06-28

摘要/Abstract

摘要：

通过筛选甘蓝型油菜绿叶和紫叶突变体叶片中差异表达基因和差异代谢物,为解析花青素合成机理奠定理论基础。对苗期叶片呈紫色的甘蓝型油菜(PL)和叶片呈绿色的甘蓝型油菜ZS11(GL)幼苗表型观察发现,PL在长出2片真叶(3周)时紫色斑驳分布,随着发育紫色逐渐加深,抽薹期后紫色逐渐变浅最终消失(>16周),GL叶片则全时期呈绿色;转录组KEGG分析在第6,13,16周找到2 523个共差异表达基因,上述差异表达基因在花青素合成途径显著富集。与花青素合成相关的24个基因显著差异表达,包括3个MYBL2、1个C4H、1个F3H、2个F3'H、2个TT8、3个DFR、4个ANS、5个UGT、3个TT19,上述基因在第6,13周PL叶片中表达量均高于GL。选取8个差异表达基因进行qRT-PCR验证,结果与转录组分析数据趋势一致。花青素靶向代谢组共检测到50种花青素,其中29种积累差异显著;PL相对于GL有16种花青素积累上调,13种花青素积累下调。PL叶片中多种矢车菊素积累量显著增加,而矮牵牛素积累量显著下降,花青素合成途径中差异表达基因和差异代谢物的表达量均达到显著水平。TT8及其靶基因(DFR、ANS、UGT、TT19)在发育早期(6~13周)的上调促进了PL中以矢车菊素为主的花青素的积累以及矮牵牛素为主的花青素的减少,导致叶片叶色出现差异。而发育后期(13~16周),这些花青素合成基因的下调和花青素的降解可能导致花青素积累的减少,使PL由紫叶转变为绿叶。

关键词: 甘蓝型油菜, 花青素, 转录组, 代谢组, 叶色

Abstract:

By screening differentially expressed genes and metabolites in the green and purple leaves of Brassica napus,it lays a theoretical foundation for the analysis of mechanism of anthocyanin synthesis.Phenotypic observation of GL and PL seedlings showed that PL exhibited a mottled distribution of purple when two true leaves were grown(3 weeks),as the purple gradually deepened during development,it gradually became lighter and eventually disappeared after the bolting stage(>16 weeks),while GL leaves remained green throughout the entire period;transcriptome analysis identified 2 523 co-differentially expressed genes at weeks 6,13,and 16,and these differentially expressed genes were significantly enriched in the anthocyanin synthesis pathway.Twenty-four genes related to anthocyanin synthesis were significantly differentially expressed,including 3 MYBL2,1 C4H,1 F3H,2 F3'H,2 TT8,3 DFR,4 ANS,5 UGT,and 3 TT19.The expression levels of these genes in PL leaves were higher than those in GL at 6 and 13 weeks.Eight differentially expressed genes were selected for qRT-PCR validation and the results were consistent with the trend of transcriptome analysis data.A total of 50 anthocyanins were detected in the anthocyanin-targeted metabolome,of which 29 showed significant difference in accumulation;compared to GL,PL had 16 types of anthocyanin accumulation upregulated and 13 types of anthocyanin accumulation downregulated.The up-regulation of TT8 and its target genes(DFR,ANS,UGT,TT19)in early development(6—13 weeks)promoted the accumulation of cyanidin-based anthocyanins and the decrease of petunian-based anthocyanins in PL.The expression levels of differentially expressed genes and metabolites in the anthocyanin synthesis pathway reached significant levels.

Key words: Brassica napus, Anthocyanins, Transcriptome, Metabolome, Leaf color

周弘峰, 朱思颖, 贺丹, 刘丽莉, 陈道宗, 谭晨, 张大为, 严明理. 紫叶甘蓝型油菜花青素合成的转录组-代谢组联合分析[J]. 华北农学报, 2024, 39(3): 25-34. doi: 10.7668/hbnxb.20194754.

ZHOU Hongfeng, ZHU Siying, HE Dan, LIU Lili, CHEN Daozong, TAN Chen, ZHANG Dawei, YAN Mingli. Transcriptomic and Metabolomic Analysis of Anthocyanin Synthesis in Purple Leaf Brassica napus[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(3): 25-34. doi: 10.7668/hbnxb.20194754.

http://www.hbnxb.net/CN/Y2024/V39/I3/25

图/表 10

表1 qRT-PCR相应的引物

Tab.1 The corresponding primers of qRT-PCR

基因名称 Gene ID	正向引物序列(5'—3') Forward primer(5'—3')	反向引物序列(5'—3') Reverse primer(5'—3')
BnaC09G0215200ZS	TCTCACGATGCGACGATTCT	ATTGTCATCACTGCCCGGAA
BnaA09G0187400ZS	ATCCAGAATATAACGTGCCTT	CAGATTTCAAATGTTCCGGTA
BnaC01G0158400ZS	ATGCACATTGGAGATACGTT	TTAGCCGGAGACTCAACACT
BnaA01G0127300ZS	AGATCTCTCACTTTGGCCTA	ACTTCATTCTCTAGACGGTCA
BnaA03G0469300ZS	CCAAAAGAATACATCCGTCCA	CATAGCAGCCTTCTTGAGC
BnaC07G0445600ZS	AAGCCACAGGAAAGATCCAA	CGTGGCTTCTATGTAATCACT
BnaC09G0362400ZS	ATTCCGCTGATTGGATACCTG	CTTCCCGTCAATGATCTCTCG
BnaA06G0190700ZS	CTTGCGGTTGCAATGAACCAA	CTTCCTCACTAGAGGCGGTTTA
BnaA10G0103400ZS	TCCCCAAGAAAGCTCTAGACCA	ACGCAGAATTGTCTCTACCTC
BnaA02G0068400ZS	ACTATGCGACCAAATACTCCA	CGTCACATTCTTCGCCTAACCTG
BnaC05G0016900ZS	AGCTAGGCTTGGTGAGTCTAAGTAT	AACCCAAGCACTGACATGTGG
Actin	CTTCCTCACGCTATCCTCCG	AGCCGTCTCCAGCTCTTGC

图1 紫叶甘蓝型油菜和绿叶甘蓝型油菜不同时期表型以及幼叶总花青素含量 A.PL和GL表型对比(3周);B.PL和GL表型对比(6周);C.PL和GL幼叶花青素总含量积累。不同小写字母表示差异显著(P<0.05)。图4—5同。

Fig.1 Phenotypes of PL and GL in different periods and the contents of total anthocyanin in young leaves A.Comparison of PL and GL phenotypes(3 weeks);B.Comparison of PL and GL phenotypes(6 weeks);C.Accumulation of total anthocyanin contents in PL and GL young leaves.Different lowercase letters indicate significant differences(P<0.05).The same as Fig.4—5.

表2 油菜转录组测序数据统计

Tab.2 Statistics of transcriptome sequencing of Brassica napus

样本ID Sample ID	Read数量/M Number of read	碱基数量/Gb Number of base	Q30碱基数量/Gb (Q30百分比/%) Q30 bases(Q30 percentage)	Q20碱基数量/Gb (Q20百分比/%) Q20 bases(Q20 percentage)	比对率/% Mapping rate
GL_6Wk_r1	52.068	7.731	7.604(98.35)	7.349(95.06)	96.21
GL_6Wk_r2	49.644	7.391	7.269(98.35)	7.024(95.03)	96.89
GL_6Wk_r3	50.693	7.546	7.418(98.30)	7.162(94.91)	97.04
PL_6Wk_r1	49.655	7.393	7.268(98.31)	7.018(94.92)	93.55
PL_6Wk_r2	45.563	6.785	6.673(98.35)	6.449(95.04)	93.90
PL_6Wk_r3	40.911	6.082	5.948(97.81)	5.684(93.47)	92.93
GL_13Wk_r1	47.623	7.116	6.974(98.00)	6.701(94.16)	96.79
GL_13Wk_r2	51.203	7.649	7.503(98.09)	7.221(94.41)	96.87
GL_13Wk_r3	42.116	6.297	6.175(98.06)	5.939(94.32)	96.90
PL_13Wk_r1	50.268	7.514	7.369(98.07)	7.089(94.34)	92.77
PL_13Wk_r2	45.394	6.781	6.649(98.05)	6.394(94.29)	92.82
PL_13Wk_r3	42.250	6.317	6.193(98.04)	5.953(94.23)	92.72
GL_16Wk_r1	49.853	7.452	7.276(97.64)	6.950(93.26)	96.41
GL_16Wk_r2	41.980	6.280	6.129(97.59)	5.849(93.14)	96.60
GL_16Wk_r3	45.819	6.860	6.695(97.59)	6.388(93.12)	96.72
PL_16Wk_r1	48.443	7.251	7.074(97.56)	6.748(93.07)	92.80
PL_16Wk_r2	43.808	6.554	6.393(97.53)	6.094(92.97)	92.77
PL_16Wk_r3	46.205	6.904	6.745(97.68)	6.442(93.30)	92.90

图2 GL和PL不同时期差异基因统计以及共同差异表达基因功能富集分析 A.GL和PL不同时期差异表达基因的数量:上调表达代表基因在PL中相对于GL上调表达,下调表达代表基因在PL中相对于GL基因下调表达;B.PL和GL在不同发育阶段共有的差异表达基因的维恩图:G0.第6周的DEGs,G1.第13周的DEGs,G2.第16周的DEGs;C.共有差异表达基因Pathway富集分析:1.转录因子,2.囊泡运输中的SNARE相互作用,3.倍半萜类和三萜类生物合成,4.分泌系统,5.真核生物中的核糖体生物发生,6.核糖体,7.嘌呤代谢,8.光合作用的蛋白质,9.光合作用,10.苯丙氨酸、酪氨酸和色氨酸的生物合成,11.核苷酸切除修复,12.错配修复,13.同源重组,14.外来体,15.DNA复制蛋白,16.DNA复制,17.染色体及相关蛋白,18.伴侣和折叠催化剂,19.精氨酸和脯氨酸代谢,20.花青素生物合成。

Fig.2 Statistical analysis of DEGs between GL and PL at different developmental stages and enrichment analysis of co-differentially expressed genes A.The number of DEGs among GL and PL in different periods:Up-regulated expression indicates genes were up-regulated in PL compared with GL,Down-regulated expression indicates genes were Down-regulated in PL compared with GL;B.Venn diagrams of differentially expressed genes between PL and GL shared at different developmental stages:G0.DEGs at week 6,G1.DEGs at week 13,G2.DEGs at week16;C.Enrichment analysis of genes expression in pathway:1.Translation factors,2.SNARE interactions in vesicular transport,3.Sesquiterpenoid and triterpenoid biosynthesis,4.Secretion system,5.Ribosome biogenesis in eukaryotes,6.Ribosome,7.Purine metabolism,8.Photosynthesis proteins,9.Photosynthesis,10.Phenylalanine,tyrosine and tryptophan biosynthesis,11.Nucleotide excision repair,12.Mismatch repair,13.Homologous recombination,14.Exosome,15.DNA replication proteins,16.DNA replication,17.Chromosome and associated proteins,18.Chaperones and folding catalysts,19.Arginine and proline metabolism,20.Anthocyanin biosynthesis.

图3 花青素合成途径相关基因表达水平热图花青素合成相关基因表达倍数分析,Log₂FPKM值;-.log₂FC≤1,+.1<log₂FC≤log₂100,++.log₂FC>log₂100,P.只在PL中表达,G.只在GL中表达。

Fig.3 Heat map of gene expression levels of anthocyanin biosynthesis pathway related gene Fold-fold analysis of genes related to anthocyanin synthesis,Log₂ FPKM;-.log₂FC≤1,+.1<log₂FC≤log₂100,++.log₂FC>log₂100,P.Only expressed in PL,G.Only expressed in GL.

图4 差异基因qRT-PCR验证结果

Fig.4 Verification results of qRT-PCR of differentially expressed gene UGT.BnaC09G0362400ZS;GST.BnaA02G0068400ZS;ANS1.BnaC01G0158400ZS;ANS2.BnaA01G0127300ZS;ANS3.BnaA03G0469300ZS;ANS4.BnaC07G0445600ZS;DFR1.BnaC09G0215200ZS;DFR2.BnaA09G0187400ZS.

图5 花青素代谢组分析 A.花青素合成相关大类代谢物含量,积累不同小写字母表示差异显著(P<0.05);B.花青素差异代谢物热图,Log₂(花青素代谢产物含量(μg/g)):1.矢车菊素-3-(6-O-p-对香豆酰)-葡萄糖苷,2.矢车菊素-3-O-槐糖苷,3.矢车菊素-3-O-5-O-(6-O-对香豆酰)-二葡萄糖苷,4.矢车菊素-3,5,3'-O-三葡萄糖苷,5.矢车菊素-3-(6″-咖啡酰槐糖苷)-5-葡萄糖苷,6.芍药花素-3,5-O-二葡萄糖苷,7.山柰酚-3-O-芸香糖苷,8.矢车菊素-3-O-桑布双糖苷-5-O-葡萄糖苷,9.矢车菊素-3-O-桑布双糖苷,10.飞燕草素-3-O-鼠李糖苷,11.飞燕草素-3-O-5-O-(6-O-对香豆酰)-二葡萄糖苷,12.锦葵色素-3-O-(6-O-丙二酰-β-D-葡萄糖苷),13.天竺葵素-3-O-5-O-(6-O-对香豆酰)-二葡萄糖苷,14.芍药花素-3-(咖啡酰-葡萄糖基-葡萄糖苷)-5-葡萄糖苷,15.芍药花素-3-O-(6-O-丙二酰-β-D-葡萄糖苷),16.原花青素B1,17.矢车菊素-3-O-木糖苷,18.矢车菊素-3-O-葡萄糖苷,19.矮牵牛素-3-O-半乳糖苷,20.矮牵牛素-3-O-葡萄糖苷,21.矮牵牛素-3-O-槐糖苷,22.柚皮素-7-O-葡萄糖苷,23.槲皮素-3-O-葡萄糖苷(异槲皮苷),24.矢车菊素-3-O-芸香糖苷,25.飞燕草素-3-O-(6-O-丙二酰-β-D-葡萄糖苷),26.飞燕草素-3-O-槐糖苷,27.飞燕草素-3-O-葡萄糖苷,28.矮牵牛素-3-O-5-O-(6-O-对香豆酰)-二葡萄糖苷,29.矮牵牛素-3-(6-O-p-对香豆酰)-葡萄糖苷。

Fig.5 Anthocyanin metabolome analysis A.Accumulation of major metabolites related to anthocyanin synthesis,different lowercase letters indicate significant differences(P<0.05);B.Heat map of anthocyanin differential metabolites(Log₂ (anthocyanin metabolites content)):1.Cyanidin-3-(6-O-p-caffeoyl)-glucoside,2.Cyanidin-3-O-sophoroside,3.Cyanidin-3-O-5-O-(6-O-coumaroyl)-diglucoside,4.Cyanidin-3,5,3'-O-triglucoside,5.Cyanidin-3-(6″-caffeylsophoroside)-5-glucoside,6.Peonidin-3,5-O-diglucoside,7.Kaempferol-3-O-rutinoside,8.Cyanidin-3-O-sambubioside-5-O-glucoside,9.Cyanidin-3-O-sambubioside,10.Delphinidin-3-O-rhamnoside,11.Delphinidin-3-O-5-O-(6-O-coumaroyl)-diglucoside,12.Malvidin-3-O-(6-O-malonyl-beta-D-glucoside),13.Pelargonidin-3-O-5-O-(6-O-coumaroyl)-diglucoside,14.Peonidin-3-(caffeoyl-glucosyl-glucoside)-5-glucoside,15.Peonidin-3-O-(6-O-malonyl-beta-D-glucoside),16.Procyanidin B1,17.Cyanidin-3-O-xyloside,18.Cyanidin-3-O-glucoside,19.Petunidin-3-O-galactoside,20.Petunidin-3-O-glucoside,21.Petunidin-3-O-sophoroside,22.Naringenin-7-O-glucoside,23.Quercetin-3-O-glucoside,24.Cyanidin-3-O-rutinoside,25.Delphinidin-3-O-(6-O-malonyl-beta-D-glucoside),26.Delphinidin-3-O-sophoroside,27.Delphinidin-3-O-glucoside,28.Petunidin-3-O-5-O-(6-O-coumaroyl)-diglucoside,29.Petunidin-3-O-(6-O-p-coumaroyl)-glucoside.

图6 PL相对GL差异代谢物差异倍数分析

Fig.6 Analysis of differential metabolite multiples of PL relative GL

图7 差异代谢物KEGG分类

Fig.7 Differential metabolite KEGG classification

图8 花青素生物合成转录组与代谢组联合分析红色基因.该基因在PL中相比GL上调表达;黑色基因.该基因在PL和GL表达差异不显著;红色上行箭头.该代谢物在PL中上调积累;蓝色下行箭头.该代谢物在PL中下调积累。

Fig.8 Joint analysis of anthocyanin biosynthesis in transcriptome and metabolome The red gene.The up-regulated expression of the gene in PL compared with GL;The black gene.There is no significant difference in the expression of PL and GL;The red ascending arrow.The up-regulated accumulation of the metabolite in PL;The blue down arrow.Down-regulated accumulation of this metabolite in PL.

参考文献 32

[1]	Li L, Zhang H, Liu J N, Huang T Z, Zhang X S, Xie H, Guo Y R, Wang Q C, Zhang P, Qin P. Grain color formation and analysis of correlated genes by metabolome and transcriptome in different wheat lines at maturity[J]. Frontiers in Nutrition, 2023, 10:1112497.doi:10.3389/fnut.2023.1112497.
[2]	He D, Zhang D W, Li T, Liu L L, Zhou D G, Kang L, Wu J F, Liu Z S, Yan M L. Whole-genome identification and comparative expression analysis of anthocyanin biosynthetic genes in Brassica napus[J]. Frontiers in Genetics, 2021, 12:764835.doi:10.3389/fgene.2021.764835.
[3]	Lepiniec L, Debeaujon I, Routaboul J M, Baudry A, Pourcel L, Nesi N, Caboche M. Genetics and biochemistry of seed flavonoids[J]. Annual Review of Plant Biology, 2006, 57:405-430.doi:10.1146/annurev.arplant.57.032905.105252. pmid: 16669768
[4]	Grotewold E. The genetics and biochemistry of floral pigments[J]. Annual Review of Plant Biology, 2006, 57:761-780.doi:10.1146/annurev.arplant.57.032905.105248. pmid: 16669781
[5]	Zhang H N, Li W C, Wang H C, Shi S Y, Shu B, Liu L Q, Wei Y Z, Xie J H. Transcriptome profiling of light-regulated anthocyanin biosynthesis in the pericarp of litchi[J]. Frontiers in Plant Science, 2016, 7:963.doi:10.3389/fpls.2016.00963.
[6]	Fukusaki E I, Kawasaki K, Kajiyama S, An C I, Suzuki K, Tanaka Y, Kobayashi A. Flower color modulations of Torenia hybrida by downregulation of chalcone synthase genes with RNA interference[J]. Journal of Biotechnology, 2004, 111(3):229-240.doi:10.1016/j.jbiotec.2004.02.019.
[7]	陈清, 汤浩茹, 董晓莉, 侯艳霞, 罗娅, 蒋艳, 黄琼瑶. 植物Myb 转录因子的研究进展[J]. 基因组学与应用生物学, 2009, 28(2):365-372.doi:10.3969/gab.028.000365.
	Chen Q, Tang H R, Dong X L, Hou Y X, Luo Y, Jiang Y, Huang Q Y. Progress in the study of plant Myb transcription factors[J]. Genomics and Applied Biology, 2009, 28(2):365-372.
[8]	van der Krol A R, Mur L A, de Lange P, Mol J N, Stuitje A R. Inhibition of flower pigmentation by antisense CHS genes:promoter and minimal sequence requirements for the antisense effect[J]. Plant Molecular Biology, 1990, 14(4):457-466.doi:10.1007/BF00027492. pmid: 2102827
[9]	Gonzalez A, Zhao M Z, Leavitt J M, Lloyd A M. Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings[J]. The Plant Journal, 2008, 53(5):814-827.doi:10.1111/j.1365-313X.2007.03373.x. pmid: 18036197
[10]	Chalhoub B, Denoeud F, Liu S Y, Parkin I A P, Tang H B, Wang X Y. Early allopolyploid evolution in the post-neolithic Brassica napus oilseed genome[J]. Science, 2014, 345(6199):950-953.doi:10.1126/science.1253435. pmid: 25146293
[11]	Fujii K, Ohmido N. Stable progeny production of the amphidiploid resynthesized Brassica napus cv.Hanakkori,a newly bred vegetable[J]. Theoretical and Applied Genetics, 2011, 123(8):1433-1443.doi:10.1007/s00122-011-1678-5. pmid: 21861174
[12]	Grant K, Carey N M, Mendoza M, Schulze J, Pilon M, Pilon-Smits E A H, van Hoewyk D. Adenosine 5'-phosphosulfate reductase(APR2)mutation in Arabidopsis implicates glutathione deficiency in selenate toxicity[J]. The Biochemical Journal, 2011, 438(2):325-335.doi:10.1042/BJ20110025.
[13]	Li H B, Zhu L X, Yuan G G, Heng S P, Yi B, Ma C Z, Shen J X, Tu J X, Fu T D, Wen J. Fine mapping and candidate gene analysis of an anthocyanin-rich gene,BnaA.PL1,conferring purple leaves in Brassica napus L.[J]. Molecular Genetics and Genomics, 2016, 291(4):1523-1534.doi:10.1007/s00438-016-1199-7.
[14]	Goswami G, Nath U K, Park J I, Hossain M R, Biswas M K, Kim H T, Kim H R, Nou I S. Transcriptional regulation of anthocyanin biosynthesis in a high-anthocyanin resynthesized Brassica napus cultivar[J]. Journal of Biological Research, 2018, 25:19.doi:10.1186/s40709-018-0090-6.
[15]	Zhang D W, Liu L L, Zhou D G, Liu X J, Liu Z S, Yan M L. Genome-wide identification and expression analysis of anthocyanin biosynthetic genes in Brassica juncea[J]. Journal of Integrative Agriculture, 2020, 19(5):1250-1260.doi:10.1016/s2095-3119(20)63172-0.
[16]	Ma Y J, Duan H R, Zhang F, Li Y, Yang H S, Tian F P, Zhou X H, Wang C M, Ma R. Transcriptomic analysis of Lycium ruthenicum Murr.during fruit ripening provides insight into structural and regulatory genes in the anthocyanin biosynthetic pathway[J]. PLoS One, 2018, 13(12):e0208627.doi:10.1371/journal.pone.0208627.
[17]	Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCT method[J]. Methods, 2001, 25(4):402-408.doi:10.1006/meth.2001.1262. pmid: 11846609
[18]	Yang Z Q, Wang S B, Wei L L, Huang Y M, Liu D X, Jia Y P, Luo C F, Lin Y C, Liang C Y, Hu Y, Dai C, Guo L, Zhou Y M, Yang Q Y. BnIR:a multi-omics database with various tools for Brassica napus research and breeding[J]. Molecular Plant, 2023, 16(4):775-789.doi:10.1016/j.molp.2023.03.007.
[19]	邹婷, 刘丽莉, 向建华, 周定港, 吴金锋, 李莓, 李宝, 张大为, 严明理. 芸薹属植物MYBL2基因的克隆及其在A、B、C基因组中的PCR鉴别[J]. 中国农业科学, 2023, 56(3):416-429.doi:10.3864/j.issn.0578-1752.2023.03.002.
	Zou T, Liu L L, Xiang J H, Zhou D G, Wu J F, Li M, Li B, Zhang D W, Yan M L. Cloning of MYBL2 gene from Brassica and its PCR identification in genomes A,B and C[J]. Scientia Agricultura Sinica, 2023, 56(3):416-429.
[20]	Davies K M, Schwinn K E. Transcriptional regulation of secondary metabolism[J]. Functional Plant Biology, 2003, 30(9):913-925.doi:10.1071/FP03062. pmid: 32689076
[21]	Guo N, Cheng F, Wu J, Liu B, Zheng S N, Liang J L, Wang X W. Anthocyanin biosynthetic genes in Brassica rapa[J]. BMC Genomics, 2014, 15(1):426.doi:10.1186/1471-2164-15-426.
[22]	Ma Z H, Li W F, Mao J, Li W, Zuo C W, Zhao X, Dawuda M M, Shi X Y, Chen B H. Synthesis of light-inducible and light-independent anthocyanins regulated by specific genes in grape Marselan(V.vinifera L.)[J]. PeerJ, 2019, 7:e6521.doi:10.7717/peerj.6521.
[23]	Kim J, Lee W J, Vu T T, Jeong C Y, Hong S W, Lee H. High accumulation of anthocyanins via the ectopic expression of AtDFR confers significant salt stress tolerance in Brassica napus L.[J]. Plant Cell Reports, 2017, 36(8):1215-1224.doi:10.1007/s00299-017-2147-7.
[24]	Sun Y, Li H, Huang J R. Arabidopsis TT19 functions as a carrier to transport anthocyanin from the cytosol to tonoplasts[J]. Molecular Plant, 2012, 5(2):387-400.doi:10.1093/mp/ssr110. pmid: 22201047
[25]	Xu W J, Grain D, Bobet S, Le Gourrierec J, Thévenin J, Kelemen Z, Lepiniec L, Dubos C. Complexity and robustness of the flavonoid transcriptional regulatory network revealed by comprehensive analyses of MYB-bHLH-WDR complexes and their targets in Arabidopsis seed[J]. The New Phytologist, 2014, 202(1):132-144.doi:10.1111/nph.12620.
[26]	He Q, Zhang Z F, Zhang L G. Anthocyanin accumulation,antioxidant ability and stability,and a transcriptional analysis of anthocyanin biosynthesis in purple heading Chinese cabbage(Brassica rapa L.ssp.pekinensis)[J]. Journal of Agricultural and Food Chemistry, 2016, 64(1):132-145.doi:10.1021/acs.jafc.5b04674.
[27]	Lee C, Lee J, Lee J. Relationship of fruit color and anthocyanin content with related gene expression differ in strawberry cultivars during shelf life[J]. Scientia Horticulturae, 2022, 301:111109.doi:10.1016/j.scienta.2022.111109.
[28]	Jiang N, Chung S O, Lee J, Ryu D, Lim Y P, Park S, Lee C, Song J, Kim K, Park J T, An G. Increase of phenolic compounds in new Chinese cabbage cultivar with red phenotype[J]. Horticulture,Environment,and Biotechnology, 2013, 54(1):82-88.doi:10.1007/s13580-013-0136-5.
[29]	Zhao Y, Qi X H, Liu Z J, Zheng W F, Guan J, Liu Z Y, Ren J, Feng H, Zhang Y. Transcriptome and metabolome profiling to explore the causes of purple leaves formation in non-heading Chinese cabbage(Brassica rapa L.ssp.chinensis makino var.mutliceps hort.)[J]. Foods, 2022, 11(12):1787.doi:10.3390/foods11121787.
[30]	唐康, 李桂花, 郭巨先, 罗文龙, 骆善伟, 姜诗政, 符梅. 基于代谢组学的红菜薹和菜心茎部色泽差异成分比较[J]. 现代食品科技, 2023, 39(9):137-143.doi:10.13982/j.mfst.1673-9078.2023.9.0998.
	Tang K, Li G H, Guo J X, Luo W L, Luo S W, Jiang S Z, Fu M. Metabolomics-based comparison of color difference components in the stems of Hongcaitai and Caixin[J]. Modern Food Science and Technology, 2023, 39(9):137-143.
[31]	Xie Y, Tan H J, Ma Z X, Huang J R. DELLA proteins promote anthocyanin biosynthesis via sequestering MYBL2 and JAZ suppressors of the MYB/bHLH/WD40 complex in Arabidopsis thaliana[J]. Molecular Plant, 2016, 9(5):711-721.doi:10.1016/j.molp.2016.01.014. pmid: 26854848
[32]	Hrazdina G, Iredale H, Mattick L R. Anthocyanin composition of Brassica oleracea cv.Red Danish[J]. Phytochemistry, 1977, 16(2):297-299.doi:10.1016/s0031-9422(00)86817-x.

选择文件类型/文献管理软件名称

选择包含的内容

紫叶甘蓝型油菜花青素合成的转录组-代谢组联合分析

Transcriptomic and Metabolomic Analysis of Anthocyanin Synthesis in Purple Leaf Brassica napus

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 10

参考文献 32

相关文章 15

编辑推荐

Metrics

本文评价

[1]	张敬敬, 田鹏, 于洪春, 李冰, 高秀瑞, 刘伟, 吴楠, 赵鑫泽, 宋雪, 刘会茹, 潘秀清, 武彦荣. 基于BSA-seq和RNA-seq挖掘西瓜果皮硬度候选基因[J]. 华北农学报, 2024, 39(5): 63-71.
[2]	陈雨蝶, 张泽荣, 李恒湘, 李天乐, 曾思杰, 邬贤梦, 熊兴华, 肖钢. 甘蓝型油菜BnEXO基因家族功能与表达分析[J]. 华北农学报, 2024, 39(5): 24-32.
[3]	冯韬, 陈勤勤, 杨佳, 谭晖, 尹明智, 胡燕. 外源蔗糖对甘蓝型油菜种子萌发与幼苗发育的影响[J]. 华北农学报, 2024, 39(5): 110-116.
[4]	吴心成, 贺日升, 肖硕定, 张振乾, 杨柳, 刘忠松, 陈浩. 甘蓝型油菜抗茎秆倒伏鉴定与候选基因挖掘[J]. 华北农学报, 2024, 39(4): 54-63.
[5]	杨明煊, 李明玉, 汪波, 王泽, 刘智强, 周广生, 于放, 刘志文. 甘蓝型油菜转录因子BnHY5-2参与盐碱胁迫的响应[J]. 华北农学报, 2024, 39(4): 47-53.
[6]	周嫒婷, 彭睿琦, 葛白瑞雪, 王芳, 伍建榕, 马焕成. 生防菌特基拉芽孢杆菌以不同方式处理油茶后的转录组分析[J]. 华北农学报, 2024, 39(4): 194-205.
[7]	朱凡, 徐芷琪, 段雨洁, 李阳阳, 宋有洪. 玉米花丝响应干旱胁迫的转录组学分析[J]. 华北农学报, 2024, 39(1): 37-47.
[8]	梅艳芳, 刘瑞莉, 韩明轩, 张文琦, 陈振鹏, 王秀源, 苗秀萍, 肖超柱, 张任政, 董雅娟, 柏学进. 基于转录组测序技术挖掘影响布莱凯特黑牛睾丸大小的功能基因[J]. 华北农学报, 2024, 39(1): 191-200.
[9]	时梦, 冀晓昊, 杨兴旺, 王莹莹, 王海波, 王孝娣. 桃叶片低温响应转录组分析[J]. 华北农学报, 2023, 38(S1): 49-57.
[10]	牛早柱, 赵艳卓, 陈展, 宣立锋, 牛帅科, 李亚囡, 杨丽丽. 铺设反光膜促进设施葡萄着色的机理初探[J]. 华北农学报, 2023, 38(S1): 203-210.
[11]	赵明珠, 王丽丽, 马作斌, 唐志强, 郑文静. *水稻DEP1突变体籽粒品质性状及其代谢组分析*[J]. 华北农学报, 2023, 38(S1): 1-7.
[12]	孙晓波, 郭臻昊, 李畅, 周惠民, 刘晓青, 苏家乐. 杜鹃花品种大鸳鸯锦花瓣条纹形成的色素基础和差异表达基因的挖掘[J]. 华北农学报, 2023, 38(6): 26-35.
[13]	陈春, 魏岳, 黄聪, 黎观红, 谢金防, 武艳平. 基于转录组测序筛选鸡皮肤毛囊性状相关基因和关键通路[J]. 华北农学报, 2023, 38(6): 218-227.
[14]	闫尊强, 王鹏飞, 杨巧丽, 黄晓宇, 高小莉, 滚双宝. C型产气荚膜梭菌性腹泻仔猪脾脏转录组分析[J]. 华北农学报, 2023, 38(5): 188-196.
[15]	彭多姿, 黄颢源, 范占煌, 戴悦, 元妙新, 张振乾. 不同杂交类型甘蓝型油菜在重金属污染土中的适应性研究[J]. 华北农学报, 2023, 38(4): 181-189.

模态框（Modal）标题

选择文件类型/文献管理软件名称

选择包含的内容

紫叶甘蓝型油菜花青素合成的转录组-代谢组联合分析

Transcriptomic and Metabolomic Analysis of Anthocyanin Synthesis in Purple Leaf Brassica napus

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 10

参考文献 32

相关文章 15

编辑推荐

Metrics

本文评价