基于转录组挖掘胡麻干旱胁迫响应的关键基因

doi:10.7668/hbnxb.20196084

摘要/Abstract

摘要：

为挖掘胡麻响应干旱胁迫的关键基因并解析其内在分子机理,以期为抗旱品种选育提供理论支持。以胡麻抗旱品种天亚13号根系为材料,采用不同浓度(5%,10%,20%,30%)的聚乙二醇(PEG)模拟干旱胁迫。通过分析根长、根粗及干/鲜质量比等生理指标,确定20% PEG为最适胁迫浓度,该浓度处理下上述指标较对照分别显著降低31.63%,30.50%和13.39百分点。在此基础上,利用Illumina转录组测序技术对不同干旱胁迫时间点(8,16,24 h)的根系进行转录组分析,分别鉴定到5 428,5 304和3 603个差异表达基因(DEGs)。KEGG富集分析表明,这些差异表达基因显著富集于碳水化合物代谢途径(12个基因)、甜菜碱生物合成途径(4个基因)及抗氧化防御系统(36个基因)等关键过程。通过qRT-PCR对部分差异表达基因进行验证,其表达趋势与RNA-seq结果一致,证实了数据的可靠性。系统揭示胡麻在干旱胁迫下通过协同调控渗透调节物质(如甜菜碱)合成、能量代谢重编程(碳水化合物代谢)以及活性氧清除机制以增强抗旱能力,为胡麻分子设计育种提供了重要的理论依据和候选基因资源。

关键词: 胡麻, 干旱胁迫, 转录组学, 抗旱基因

Abstract:

In order to uncover key genes involved in flax's response to drought stress and elucidate the underlying molecular mechanisms,thereby providing theoretical support for breeding drought-resistant varieties.Using roots of the drought-resistant flax cultivar Tianya 13 as experimental material,drought stress was simulated with different concentrations(5%,10%,20%,30%)of polyethylene glycol(PEG).Based on the analysis of physiological indicators such as root length,root thickness,and dry/fresh weight ratio,20% PEG was identified as the optimal stress concentration,under which the above indicators were significantly reduced by 31.63%,30.50%,and 13.39 percentage points,respectively,compared to the control.Subsequently,Illumina transcriptome sequencing was performed on roots subjected to drought stress at different time points(8,16,24 h),identifying 5 428,5 304,and 3 603 differentially expressed genes(DEGs)at each time point,respectively.KEGG enrichment analysis revealed that these DEGs were significantly enriched in key pathways such as carbohydrate metabolism(12 genes),betaine biosynthesis(4 genes),and the antioxidant defense system(36 genes).qRT-PCR validation of selected DEGs confirmed expression trends consistent with the RNA-seq results,verifying data reliability.This study systematically reveals,that flax enhances drought tolerance through the coordinated regulation of osmotic adjustment substance(e.g.,betaine)synthesis,energy metabolism reprogramming(carbohydrate metabolism),and reactive oxygen species scavenging mechanisms.It provides important theoretical foundations and candidate gene resources for molecular design breeding in flax.

Key words: Flax, Drought stress, Transcriptomics, Drought resistance genes

中图分类号:

S188
S565

何丽, 付丹丹, 窦晨曦, 王娜, 李军. 基于转录组挖掘胡麻干旱胁迫响应的关键基因[J]. 华北农学报, 2026, 41(4): 99-111. doi: 10.7668/hbnxb.20196084.

HE Li, FU Dandan, DOU Chenxi, WANG Na, LI Jun. Transcriptome-based Identification of Key Genes Involved in Drought Stress Response in Flax[J]. Acta Agriculturae Boreali-Sinica, 2026, 41(4): 99-111. doi: 10.7668/hbnxb.20196084.

http://www.hbnxb.net/CN/Y2026/V41/I4/99

图/表 12

表1 12个样品相关质量评估

Tab.1 Assessment of the quality associated with 12 samples

样本名称 Sample name	原始序列数 Raw reads	过滤后序列数 Clean reads	过滤后碱基/Gb Clean bases	错误率/% Error rate	Q20/%	Q30/%	GC 含量/% GC content
CK_1	11763638	11585163	0.58	0.01	98.29	96.23	48.99
CK_2	12186381	11994323	0.60	0.01	98.27	96.19	49.65
CK_3	12066987	11844725	0.60	0.01	98.07	96.11	49.21
TY8 h_1	10575274	10392565	0.52	0.01	98.10	95.81	49.41
TY8 h_2	10807289	10703269	0.54	0.01	98.28	96.21	49.44
TY8 h_3	10617452	10604433	0.53	0.01	98.16	96.02	49.42
TY16 h_1	11176167	11071109	0.55	0.01	98.28	96.21	49.75
TY16 h_2	10278785	10169417	0.51	0.01	98.03	95.60	49.72
TY16 h_3	10843682	10665422	0.51	0.01	98.03	95.60	49.74
TY24 h_1	11987425	11817837	0.59	0.01	98.42	96.53	49.45
TY24 h_2	10456025	10401224	0.52	0.01	99.08	98.19	49.27
TY24 h_3	10823044	10772827	0.55	0.01	99.22	97.45	49.36

表2 16个DEGs的qRT-PCR引物序列

Tab.2 qRT-PCR primer sequences for 16 DEGs

基因编号 Gene ID	上游引物(5'—3') Forward primer(5'—3')	下游引物(5'—3') Reverse primer(5'—3')
SDH1	GGCGAACTTCATGTTTGCG	CGCAAACATGAATTCGCC
PetF	TCTGATCATGGCAAAGTCAACA	TAACTGTCCCCCCAGTTAAAGA
TYR	CTACGACATCGATGTCGC	GCGACATCGATGTCGTAG
fumB	TCTTGGCAGCAATTCGCA	TGCGAATTGCTGCCAAGA
5GT	CGTACTGCAACTTCGACA	TGTCGAAGTTGCAGTACG
CCD32	GAACAAGGTCGGCGAAAT	TAAACTGGCCGTTGGTCC
SUCD1	GGTCAGCGACTATCAAGCTG	CAGCTTGATAGTCGCTGACC
APX4	AGGACCATGTACGAATAGATGA	CTTAGGTAACACCACCTCTT
GST4	TCTACTATGGTTAAGATGA	TCATCTTAACCATAGTAGA
bHLH63	GCGTGGCTTCTCTCTGTG	CGCACGATGCCAGAACC
SOD-Mn	CCTGGTTGCCAATCAGGC	GCCTGATTGGCAACCAGG
ERF71	GGAAGCTGTCCAATGGAAT	GGGCATCGACCATCTAGGTA
WRKY49	AGGACCATGTACGAATGATA	CTAAGGTAACACCATCTCAT
CDOPA-O	ACTTCTATGGCCTACTCGAG	TGAAGATACCGGATGAGCTC
CHI1	TGTCATCATGGCAAGTCACT	ATCTGTCCCCAGTTATACA
PetE	CTCACTCCGAACCTGTCTT	AAGACAGGTTCGGAGTGAG

图1 不同浓度PEG对天亚13号幼苗根长、根粗、根系干/鲜质量比的影响不同小写字母表示处理间差异显著(P<0.05)。图8同。

Fig.1 Effects of different concentrations of PEG on root length,root thickness,root dry to fresh weight ratio of Tianya 13 seedlings Different lowercase letters are significantly different(P<0.05).The same as Fig.8.

图2 干旱胁迫后不同时间点下根系中的DEGs A.20% PEG处理下,TY8 h vs CK、TY16 h vs CK、TY24 h vs CK DEGs的韦恩图,0 h为对照(CK);B.利用热图绘制干旱胁迫8,16,24 h后胡麻根系中DEGs的表达谱;C—J.利用STEM软件分析干旱胁迫后胡麻根系中DEGs的不同时间表达模式。每个表达谱使用不同的图示表示。数据使用log₂归一化,错误发现率(FDR)<0.05。

Fig.2 DEGs in roots at different time points following drought stress A.Venn plots of TY8 h vs CK,TY16 h vs CK,and TY24 h vs CK DEGs under 20% PEG treatment,with 0 h as control(CK);B.Expression profiles of DEGs in Flax root system after drought stress at 8,16,24 h were plotted by using heatmap;C—J.Analysis of different temporal expression patterns of DEGs in Flax root system after drought stress by using STEM software.Different temporal expression patterns of DEGs in the root system.Each expression profile was represented using a different graphical representation.Data were normalised using log₂ with a false discovery rate(FDR)<0.05.

图3 干旱胁迫下DEGs的GO及KEGG分析 A、B、C分别是8 h vs CK、16 h vs CK、24 h vs CK DEGs的GO富集柱状图。纵坐标为富集的GO term,横坐标为该term中DEGs个数。*.显著富集的GO term;A:1.代谢过程;2.生物过程;3.有机合成途径;4.生物合成途径;5.细胞生物合成过程;6.氧化还原;7.有机氮化合物合成途径;8.细胞氮化合物合成途径;9.有机物代谢;10.有机氮化合物代谢;11.单个有机体代谢;12.酰胺生物合成途径;13.转录;14.细胞代谢;15.初级代谢;16.多肽类合成途径;17.细胞酰胺代谢;18.多肽代谢;19.大分子物质合成途径;20.细胞大分子类物质合成途径;21.细胞过程;22.核糖体;23.核糖核蛋白复合物;24.非记忆型细胞器;25.细胞内非膜结构细胞器;26.细胞质;27.核糖体结构组分;28.氧化还原酶活性;29.结构分子活性;30.序列特异性DNA结合。B:1.氧化还原过程;2.碳水化合物代谢;3.钾离子运输;4.细胞内碳水化合物代谢;5.代谢过程;6.单个有机体代谢过程;7.金属离子运输;8.生物过程;9.多糖代谢;10.细胞发育过程;11.碳水化合物分解代谢;12.醌代谢;13.醌合成途径;14.二糖类合成途径;15.细胞突起部分;16.ATP合成酶复合体;17.细胞型鞭毛部位;18.氧化还原酶活性;19.水解酶活性;20.水解酶水解活性;21.磷酸酯突变酶活性;22.淀粉酶活性;23.分子功能;24.ATP激活酶活性;25.拮抗剂活性;26.过渡态金属离子;27.β-淀粉酶活性;28.ATP合成酶;29.催化活性;30.肌动蛋白单体结合。C:1.氧化还原过程;2.单个有机体代谢过程;3.单个有机体;4.碳水化合物代谢;5.碳水化合物分解代谢;6.嘌呤核苷三磷酸代谢;7.核苷酸三磷酸酯代谢;8.嘌呤核糖核苷三磷酸酯代谢;9.生物过程;10.醌代谢;11.醌合成;12.核苷酸单磷酸酯代谢;13.ATP代谢;14.三磷酸核苷代谢;15.钾离子转运;16.多糖代谢;17.嘌呤核苷单磷酸代谢;18.嘌呤核糖核苷酸单磷酸代谢;19.外部包裹结构;20.氧化还原酶活性;21.血红素结合能力;22.磷酸甘油酸突变酶活性;23.四吡咯结合能力;24.水解酶水解活性;25.铜离子结合能力;26.2,3-二磷酸甘油酯内酯;27.水解酶活性;28.ATP质子传送泵;29.铁离子结合;30.ATP活性;D、E、F分别是8 h vs CK、16 h vs CK、24 h vs CK DEGs的KEGG富集散点图。点的大小表示此pathway中差异表达基因个数多少,点的颜色对应不同的Qvalue范围。D:1.新陈代谢;2.萜类骨架的生物合成;3.芪类、二芳基庚烷类和姜酚的生物合成;4.淀粉和蔗糖代谢;5.鞘脂代谢;6.核糖体;7.苯丙醇生物合成;8.苯丙氨酸代谢;9.代谢途径;10.谷胱甘肽代谢;11.葡萄糖苷酸类合成途径;12.黄酮类化合物合成途径;13.脂肪酸降解;14.二萜类物质合成途径;15.芳香族化合物的降解;16.半胱氨酸和蛋氨酸代谢;17.植物的昼夜节律;18.芸苔素类固醇的生物合成;19.次生代谢物的生物合成;20.α-亚麻酸代谢。E:1.维生素B6代谢;2.泛醌和其他萜类醌的生物合成作用;3.过氧化物酶体;4.丙烷、哌啶和吡啶生物碱的合成途径;5.硫胺素代谢;6.淀粉和蔗糖代谢;7.内质网中蛋白质加工;8.光合作用-触角蛋白;9.苯丙氨酸;10.戊糖和葡萄糖醛酸的相互转化;11.甜菜碱合成途径;12.葡萄糖苷酸的合成途径;13.类黄酮类合成途径;14.二萜类合成途径;15.半胱氨酸和蛋氨酸代谢;16.植物的昼夜节律;17.光合作用碳固定;18.次生代谢物的生物合成;19.β-丙氨酸代谢;20.α-亚麻酸代谢。F:1.泛醌和其他萜类醌的生物合成作用;2.酪氨酸代谢;3.丙烷、哌啶和吡啶生物碱的合成途径;4.淀粉和蔗糖代谢;5.内质网蛋白质加工途径;6.甜菜碱合成途径;7.苯丙氨酸合成途径;8.苯丙氨酸代谢;9.戊糖和葡萄糖醛酸的相互转化;10.异喹啉生物碱合成途径;11.糖降解/糖醛酸的生成;12.葡萄糖苷酸合成途径;13.黄酮类合成途径;14.二萜类合成途径;15.半胱氨酸和蛋氨酸代谢;16.植物昼夜节律;17.光合作用碳固定;18.过氧化物酶体;19.精氨酸和脯氨酸代谢;20.α-亚麻酸代谢。

Fig.3 GO and KEGG analysis of DEGs under drought stress A,B,and C are bar charts showing the GO enrichment of DEGs for the comparisons 8 h vs CK,16 h vs CK,and 24 h vs CK,respectively.The vertical axis represents the enriched GO terms,and the horizontal axis represents the number of DEGs within each term.*.Significantly enriched.A:1.Metabolic process;2.Biological regulation;3.Organic substance metabolic process;4.Biosynthetic process;5.Cellular biosynthetic process;6.Oxidation-reduction process;7.Organic nitrogen compound biosynthetic process;8.Cellular nitrogen compound biosynthetic process;9.Organic substance metabolic process;10.Organic nitrogen compound metabolic process;11.Single-organism metabolic process;12.Amide biosynthetic process;13.Transcription,DNA-templated;14.Cellular metabolic process;15.Primary metabolic process;16.Peptide biosynthetic process;17.Cellular amide metabolic process;18.Peptide metabolic process;19.Macromolecule biosynthetic process;20.Cellular macromolecule biosynthetic process;21.Cellular process;22.Ribosome;23.Ribonucleoprotein complex;24.Non-membrane-bounded organelle;25.Intracellular non-membrane-bounded organelle;26.Cytoplasm;27.Structural constituent of ribosome;28.Oxidoreductase activity;29.Structural molecule activity;30.Sequence-specific DNA binding.B:1.Oxidation-reduction process;2.Carbohydrate caetabolic process;3.Potassium ion transport;4.Intracellular carbohydrate metabolic process;5.Metabolic process;6.Single-organism metabolic process;7.Metal ion transport;8.Biological process;9.Polysaccharide metabolic process;10.Cell projection organization;11.Carbohydrate catabolic process;12.Quinone metabolic process;13.Quinone biosynthetic process;14.Disaccharide biosynthetic process;15.Cell projection part;16.ATP synthase complex;17.Cell-type specific flagellum part;18.Oxidoreductase activity;19.Hydrolase activity;20.Hydrolase activity,hydrolyzing O-glycosyl compounds;21.Phosphotransferase activity;22.Amylase activity;23.Molecular function;24.ATPase activity,coupled;25.Antagonist activity;26.Transition metal ion binding;27.Beta-amylase activity;28.ATP synthase;29.Catalytic activity;30.Actin monomer binding.C:1.Oxidation-reduction process;2.Single-organism metabolic process;3.Single-organism process;4.Carbohydrate metabolic process;5.Carbohydrate catabolic process;6.Purine nucleoside triphosphate metabolic process;7.Nucleoside triphosphate metabolic process;8.Purine ribonucleoside triphosphate metabolic process;9.Biological process;10.Quinone metabolic process;11.Quinone biosynthetic process;12.Nucleoside monophosphate metabolic process;13.ATP metabolic process;14.Ribonucleoside triphosphate metabolic process;15.Potassium ion transport;16.Polysaccharide metabolic process;17.Purine nucleoside monophosphate metabolic process;18.Purine ribonucleoside monophosphate metabolic process;19.External encapsulating structure;20.Oxidoreductase activity;21.Heme binding;22.Phosphoglycerate mutase activity;23.Tetrapyrrole binding;24.Hydrolase activity,hydrolyzing O-glycosyl compounds;25.Copper ion binding;26.2,3-bisphosphoglycerate-dependent phosphoglycerate mutase activity;27.Hydrolase activity;28.Proton-transporting ATPase complex;29.Iron ion binding;30.ATPase activity.D,E,and F are scatter plots showing the KEGG enrichment of DEGs for the comparisons 8 h vs CK,16 h vs CK,and 24 h vs CK,respectively.The size of the dots indicates the number of differentially expressed genes in that pathway,and the color of the dots corresponds to different Q-value ranges.D:1.Metabolism;2.Terpenoid backbone biosynthesis;3.Stilbenoid,diarylheptanoid and gingerol biosynthesis;4.Starch and sucrose metabolism;5.Sphingolipid metabolism;6.Ribosome;7.Phenylpropanoid biosynthesis;8.Phenylalanine metabolism;9.Metabolic pathways;10.Glutathione metabolism;11.Glucosinolate biosynthesis;12.Flavonoid biosynthesis;13.Fatty acid degradation;14.Diterpenoid biosynthesis;15.Degradation of aromatic compounds;16.Cysteine and methionine metabolism;17.Circadian rhythm-plant;18.Brassinosteroid biosynthesis;19.Biosynthesis of secondary metabolites;20.alpha-Linolenic acid metabolism.E:1.Vitamin B6 metabolism;2.Ubiquinone and other terpenoid-quinone biosynthesis;3.Peroxisome;4.Biosynthesis of alkaloids derived from ornithine,lysine and nicotinic acid;5.Thiamine metabolism;6.Starch and sucrose metabolism;7.Protein processing in endoplasmic reticulum;8.Photosynthesis-antenna proteins;9.Phenylalanine metabolism;10.Pentose and glucuronate interconversions;11.Betalain biosynthesis;12.Glucosinolate biosynthesis;13.Flavonoid biosynthesis;14.Diterpenoid biosynthesis;15.Cysteine and methionine metabolism;16.Circadian rhythm-plant;17.Carbon fixation in photosynthetic organisms;18.Biosynthesis of secondary metabolites;19.beta-Alanine metabolism;20.alpha-Linolenic acid metabolism.F:1.Ubiquinone and other terpenoid-quinone biosynthesis;2.Tyrosine metabolism;3.Biosynthesis of alkaloids derived from ornithine,lysine and nicotinic acid;4.Starch and sucrose metabolism;5.Protein processing in endoplasmic reticulum;6.Betalain biosynthesis;7.Phenylpropanoid biosynthesis;8.Phenylalanine metabolism;9.Pentose and glucuronate interconversions;10.Isoquinoline alkaloid biosynthesis;11.Ascorbate and aldarate metabolism;12.Glucosinolate biosynthesis;13.Flavonoid biosynthesis;14.Diterpenoid biosynthesis;15.Cysteine and methionine metabolism;16.Circadian rhythm-plant;17.Carbon fixation in photosynthetic organisms;18.Peroxisome;19.Arginine and proline metabolism;20.alpha-Linolenic acid metabolism.

图4 干旱胁迫下DEGs参与的碳水化合物代谢过程正方形.DEGs表达量,其中左侧代表8 h vs CK比较组中的DEGs表达量;中间代表16 h vs CK比较组中的DEGs表达量;右侧代表24 h vs CK比较组中的DEGs表达量。

Fig.4 DEGs are involved in carbohydrate metabolism process under drought stress The squares.Expression of DEGs,where the left side represents the expression of DEGs in the 8 h vs CK comparison group;the middle represents the expression of DEGs in the 16 h vs CK comparison group;and the right side represents the expression of DEGs in the 24 h vs CK comparison group.

图5 干旱胁迫下胡麻甜菜碱合成相关的DEGs分析圆圈.干旱胁迫下调节的代谢物;长方形.受干旱胁迫调控的基因(其中红色表示上调;蓝色表示下调);3个方块.8,16,24 h下各个基因的表达量。

Fig.5 Analysis of DEGs related to betaine synthesis in flax under drought stress Circles.Metabolites regulated under drought stress;Rectangles.Genes regulated by drought stress(where red indicates up-regulation;blue indicates down-regulation);The three squares.Expression levels of each gene under the conditions of 8,16,and 24 h respectively.

图6 甜菜碱合成途径关键基因与不同时间甜菜碱含量线性关系图

Fig.6 Linear relationship graph of key genes in betaine biosynthesis pathway and betaine content at different times

图7 抗氧化分子和GPX候选基因在4个时间点(0,8,16,24 h)的聚类热图集中校正后,log₂(TPM+1)的值表示表达水平,其中具有相似表达模式的基因聚类。

Fig.7 Clustered heatmaps of antioxidant molecules and GPX candidate genes at four different time points(0,8,16,24 h) After centralised correction,the value of log₂(TPM+1)indicates the expression level,where genes with similar expression patterns are clustered.

图8 通过qRT-PCR验证干旱胁迫下胡麻中16个DEGs 以Actin为内参基因;Y轴表示经log₂转换后DEGs的归一化表达。

Fig.8 Validation of 16 drought stress-responsive DEGs in flax by qRT-PCR Actin was used as an internal reference gene;Y-axis indicates normalised expression of DEGs after log₂ transformation.

表3 RNA-seq与qPCR结果验证

Tab.3 Validation of RNA-seq and qPCR results

基因编码 Gene ID	基因 Gene	FPKM (CK)	FPKM (8)	FPKM (16)	FPKM (24)	8 h vs CK	16 h vs 8 h	24 h vs 16 h
Lus_GLEAN_10000016	SDH1	56.45	103.03	121.24	149.64	上调	上调	上调
Lus_GLEAN_10000033	PetF	23.04	62.73	48.53	89.61	上调	下调	上调
Lus_GLEAN_10000019	TYR	65.49	176.98	40.55	107.64	上调	下调	上调
Lus_GLEAN_10000160	fumB	65.55	72.99	43.88	56.11	上调	上调	下调
Lus_GLEAN_10000376	5GT	100.87	120.66	140.77	24.55	上调	上调	下调
Lus_GLEAN_10000519	CCD32	169.77	122.92	78.98	44.32	下调	下调	下调
Lus_GLEAN_10000561	SUCD1	44.21	78.76	42.44	44.89	上调	下调	下调
Lus_GLEAN_10000711	APX4	56.67	180.88	66.43	69.21	上调	下调	下调
Lus_GLEAN_10000842	GST4	124.32	98.76	134.67	34.55	下调	上调	下调
Lus_GLEAN_10000666	bHLH63	29.88	54.44	87.66	40.98	上调	上调	下调
Lus_GLEAN_10001270	SOD-Mn	38.54	98.87	66.23	60.98	上调	下调	下调
Lus_GLEAN_10001140	ERF71	30.45	56.66	90.34	112.32	上调	上调	上调
Lus_GLEAN_10001414	WRKY49	58.65	26.54	79.45	38.45	下调	上调	下调
Lus_GLEAN_10000945	CDOPA-O	47.66	53.66	62.67	91.77	上调	上调	上调
Lus_GLEAN_10001332	CHI1	24.34	39.67	53.44	75.44	上调	上调	上调
Lus_GLEAN_10001514	PetE	46.45	92.03	111.24	150.64	上调	上调	上调

图9 16种关键基因地上和地下基因表达量测定

Fig.9 Expression levels of 16 key genes in aerial and subterranean part

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