山药品种苏蓣8号抗炭疽病基因的转录组测序与表达分析

doi:10.7668/hbnxb.20192826

摘要/Abstract

摘要：

为深入研究炭疽病抗病机制提供候选基因,从分子水平探索炭疽菌侵染对山药炭疽病基因转录水平的影响。苏蓣8号是高感炭疽病品系024经组培诱变所育成的高抗炭疽病山药新品种,为挖掘苏蓣8号炭疽病抗性基因,采用胶孢炭疽菌菌株4-2分别接种苏蓣8号和品系024,以未接种苏蓣8号的叶片为对照,利用转录组测序技术分析抗感材料在接种炭疽菌后不同时间点的差异表达基因。结果表明,抗感材料接种24,48,72,96 h共同差异表达基因个数分别为197,132,187,313个,去除各接种时间点共同差异表达的基因数,共获得711个差异表达基因。GO功能富集显示,差异基因主要与响应生物刺激、防卫反应、细胞壁代谢和氧化还原过程等有关。KEGG富集分析表明,生长素、茉莉酸和乙烯等防御相关的植物激素信号转导途径多个基因表达出现差异,其中5个生长素早期响应基因、茉莉酸和脱落酸合成关键酶基因均上调表达,乙烯响应因子ERF036下调表达,可能负向调控山药炭疽菌的侵染;参与次生代谢产物修饰的多个细胞色素P450基因、泛素连接酶及植物甾醇合成酶、防御素和凝集素基因上调表达;WRKY类、MYB类和TIFY类转录因子正向或负向调控抗病基因的表达,受转录调控的PR蛋白、NBS-LRR抗病基因、受体激酶等大量表达,抗氧化保护酶系统CAT和SOD成员在活性氧信号刺激下表达。此外,与淀粉和蔗糖合成有关酶上调表达,与淀粉分解有关酶下调表达。LAX4、IAA4、IAA21基因表达趋势与转录组测序结果基本一致,且在抗病品种表达量显著高于感病品种,推测生长素信号途径有利于山药对炭疽病的免疫反应。

关键词: 山药, 苏蓣8号, 炭疽病, 转录组, 差异表达基因

Abstract:

In this study,the transcription level of resistance genes to the infection of Colletotrichum gloeosporioides were explored at the molecular level and provided candidate genes for further study of disease resistance mechanism.Suyu 8 is a new yam variety highly resistant to anthracnose,which was bred by tissue culture mutagenesis of the highly susceptible strain 024.In order to find out the anthracnose resistance genes,Suyu 8 and strain 024 leaves were inoculated for different time by Colletotrichum gloeosporioides strain 4-2,the differentially expressed genes were analyzed by transcriptome sequencing.Concurrently,the uninoculation leaves of Suyu 8 were used as control.The number of differentially expressed genes,inoculation for 24,48,72,96 h,were 197,132,187 and 313,respectively.After removing the common differentially expressed genes at each time point,we obtained 711 differentially expressed genes.Go enrichment showed that the differential genes were mainly related to response to biological stimulation,defense response,cell wall metabolism and oxidation-reduction process.KEGG enrichment analysis revealed that many metabolic pathways related to plant disease resistance were changed.The expression of multiple genes varied which defense-related plant hormone signal transduction pathways,such as auxin,jasmonic acid,and ethylene.Among them,five early auxin-responsive genes,the key genes of jasmonic acid and abscisic acid biosynthesis were up-regulated,while ERF036 was down-regulated,which might negatively regulate the infection of Colletotrichum gloeosporioides.Several cytochrome P450 genes,ubiquitin ligases involved secondary metabolite modification and phytosterol synthetases,defensins and lectins were up-regulated.WRKY,MYB and TIFY transcription factors positively or negatively regulate the expression of disease resistance genes.Under the regulation of transcription factors,PR protein,NBS-LRR disease resistance genes,receptor kinases were highly expressed,the expressions of CAT and SOD genes of antioxidant protective enzyme systems were activated by reactive oxygen species.In addition,the enzymes related to starch and sucrose synthesis were up-regulated,and enzymes related to starch degradation were down regulated.The expression trends of LAX4,IAA4,IAA21 were consistent with that of transcriptome sequencing.The relative expression of LAX4,IAA4 and IAA21 in resistant varieties were significantly higher than susceptible varieties,suggesting that auxin signaling pathway is beneficial to the immune response to anthracnose for yam.

Key words: Yam, Suyu 8, Anthracnose, Transcriptome, Differentially expressed genes

韩晓勇, 蒋璐, 殷剑美, 金林, 张培通. 山药品种苏蓣8号抗炭疽病基因的转录组测序与表达分析[J]. 华北农学报, 2022, 37(2): 211-222. doi: 10.7668/hbnxb.20192826.

HAN Xiaoyong, JIANG Lu, YIN Jianmei, JIN Lin, ZHANG Peitong. Transcriptomic and Expression Analysis of Anthracnose Resistance-related Genes from Yam Suyu 8[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(2): 211-222. doi: 10.7668/hbnxb.20192826.

http://www.hbnxb.net/CN/Y2022/V37/I2/211

图/表 12

表1 Q-PCR基因及引物

Tab.1 The genes and primers used for Q-PCR

基因 Gene	正向引物(5'—3') Forward primer(5'—3')	反向引物(5'—3') Reverse primer (5'—3')	基因功能描述 Description of gene function
LAX4	CAATCTGGTTCTTCGCCATC	ACGAAGATCCATGCCACCAC	生长素转运类蛋白4
IAA4	AGCAAGCCTCTGATTGGCCAC	CATCTCCAACCATCATCCAATC	生长素响应蛋白4
IAA21	GGATGGAGCTCCTTATCTTAG	CACCGATAGTAAAGCAGCTGA	生长素响应蛋白17
Actin1	CTCATTGATCGGCATGGAAGC	GGGGAACATAGTTGAACCACCAC	肌动蛋白

表2 测序数据统计结果

Tab.2 The statistics of transcriptome sequencing data

样本 Sample	高质量序列 read 数/条 Clean reads	高质量序列碱基数/bp Clean data	GC含量/% GC content	质量值≥20 碱基百分比/% Q20	质量值≥30 碱基百分比/% Q30
R0h	20 504 603	6 151 380 900	45.26	96.30	90.56
S24h	20 225 071	6 067 521 400	45.35	96.18	90.34
R24h	20 160 088	6 048 026 300	45.30	96.27	90.56
S48h	18 874 399	5 662 319 600	45.67	96.13	90.25
R48h	19 875 790	5 962 737 000	45.31	96.30	90.57
S72h	18 538 496	5 561 548 900	45.05	96.24	90.46
R72h	20 022 778	6 006 833 400	44.98	96.32	90.62
S96h	18 567 803	5 570 340 900	45.40	96.16	90.33
R96h	19 518 804	5 855 641 100	44.92	96.26	90.49

图1 各样本间差异表达基因数

Fig.1 The statistics of differentially expressed genes between samples

图2 差异表达基因韦恩图
A—D.分别为接种24,48,72,96 h差异表达基因韦恩图;E.接种24,48,72,96 h共同差异表达基因韦恩图。

Fig.2 Venn map of differentially expressed genes
A—D.Venn map of differentially expressed genes after inoculation 24,48,72,96 h, respectively;E.Venn map of common differentially expressed genes after inoculation 24,48,72,96 h.

图3 差异表达基因GO分类
BP.生物过程:BP1.细胞过程;BP2.代谢过程;BP3.单组织过程;BP4.刺激响应;BP5.生物调节;BP6.定位;BP7.细胞组分组织或生物发生;BP8.发展过程;BP9.信号;BP10.多细胞生物过程;BP11.生殖;BP12.生殖过程;BP13.多组织过程;BP14.免疫系统过程;BP15.生长;BP16.脱毒;BP17.节律过程;BP18.运动;BP19.细胞杀伤;BP20.生物附着。CC.细胞组分:CC1.细胞;CC2.细胞部分;CC3.细胞器;CC4.膜;CC5.细胞器部分;CC6.膜部分;CC7.大分子复合物;CC8.胞外区;CC9.膜包围腔;CC10.细胞连接;CC11.超分子复合物;CC12.胞外区部分;CC13.拟核;CC14.其他生物体;CC15.其他生物体部分。MF.分子功能:MF1.结合;MF2.催化活性;MF3.转运活性;MF4.结构分子活性;MF5.信号转导活性;MF6.核酸结合转录因子活性;MF7.分子转导活性;MF8.分子功能调节器;MF9.电子载体活性;MF10.抗氧化活性;MF11.营养库活性;MF12.转录因子活性、蛋白质结合;MF13.金属伴侣蛋白活性;MF14.翻译调节活;MF15.蛋白标签。

Fig.3 GO classification of the differentially expressed genes
BP.Biological process:BP1.Cellular process;BP2.Metabolic process;BP3.Single-organism process;BP4.Response to stimulus;BP5.Biological regulation;BP6.Localization;BP7.Cellular component organization or biogenesis;BP8.Developmental process;BP9.Signaling;BP10.Multicellular organismal process;BP11.Reproduction;BP12.Reproductive process;BP13.Multi-organism process;BP14.Immune system process;BP15.Growth;BP16.Detoxification;BP17.Rhythmic process;BP18.Locomotion;BP19.Cell killing;BP20.Biological adhesion.CC.Cellular component:CC1.Cell;CC2.Cell part;CC3.Organelle;CC4.Membrane;CC5.Organelle part;CC6.Membrane part;CC7.Macromolecular complex;CC8.Extracellular region;CC9.Membrane-enclosed lumen;CC10.Cell junction;CC11.Supramolecular complex;CC12.Extracellular region part;CC13.Nucleoid;CC14.Other organism;CC15.Other organism part.MF.Molecular function:MF1.Binding;MF2.Catalytic activity;MF3.Transporter activity;MF4.Structural molecule activity;MF5.Signal transducer activity;MF6.Nucleic acid binding transcription factor activity;MF7.Molecular transducer activity;MF8.Molecular function regulator;MF9.Electron carrier activity;MF10.Antioxidant activity;MF11.Nutrient reservoir activity;MF12.Transcription factor activity, protein binding;MF13.Metallochaperone activity;MF14.Translation regulator activity;MF15.Protein tag.

图4 差异表达基因的GO富集圈图

Fig.4 GO enrichment circular graph of the differentially expressed genes

图5 前20条显著富集的GO类名

Fig.5 Top 20 of significant enrichment terms

图6 差异表达基因KEGG富集分类

Fig.6 KEGG enrichment classification of the differentially expressed genes

表3 差异表达基因显著富集的KEGG通路

Tab.3 KEGG pathways significantly enriched of differentially expressed genes

Ko号 Ko ID	代谢途径 Pathway	差异基因数量/个 DEG number	P值 P-value
map00040	戊糖和葡萄糖醛酸转化	5	0.000 105 447
map00010	糖酵解/糖异生	6	0.001 331 203
map00100	类固醇生物合成	3	0.002 675 332
map00960	托烷,哌啶和吡啶生物碱合成	3	0.003 124 671
map04075	植物激素信号转导	5	0.004 122 894
map00052	半乳糖代谢	3	0.007 001 598
map00500	淀粉与蔗糖代谢	4	0.017 638 130
map00030	磷酸戊糖途径	3	0.020 066 528
map00561	甘油酯代谢	3	0.034 833 484
map00780	生物素代谢	2	0.043 858 617

图7 差异表达基因热图
A.植物与病原互作;B.淀粉和蔗糖代谢;C.激素类;D.转录因子;E.次生代谢。

Fig.7 Heatmap of the differentially expressed genes
A.Plant-phytopathogen interaction;B.Starch and sucrose metabolism;C.Hormones;D.Transcription factors;E.Secondary metabolism.

图8 差异表达基因Q-PCR验证

Fig.8 Q-PCR verification of differentially expressed genes

图9 差异表达基因Q-PCR表达分析
024-S、 024-R、 JXD-S、JXD-R、Su8-R、Su6-R、HH-R分别表示品系024感病叶、健康叶,江西短紫山药感病叶、健康叶,苏蓣8号、苏蓣6号和红河山药。

Fig.9 Q-PCR analysis of differentially expressed genes
024-S, 024-R, JXD-S, JXD-R, Su8-R, Su6-R, HH-R show strain 024 susceptible leaf and healthy leaf, Jiangxi yam susceptible leaf and healthy leaf, Suyu 8, Suyu 6 and Honghe yam.

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