中国水稻科学 ›› 2023, Vol. 37 ›› Issue (6): 563-576.DOI: 10.16819/j.1001-7216.2023.230301
收稿日期:
2023-03-30
修回日期:
2023-04-20
出版日期:
2023-11-10
发布日期:
2023-11-14
通讯作者:
*email: zhywang@zju.edu.cn
基金资助:
TONG Qi, WANG Chunyan, QUE Yawei, XIAO Yu, WANG Zhengyi*()
Received:
2023-03-30
Revised:
2023-04-20
Online:
2023-11-10
Published:
2023-11-14
Contact:
*email: zhywang@zju.edu.cn
摘要:
【目的】 探明稻瘟病菌热激蛋白(HSP)40在形态分化和致病过程中的作用。【方法】 利用DNA同源重组方法敲除了稻瘟病菌HSP40编码基因MoMHF6,获得ΔMomhf6突变体,并通过表型分析、基因回补、RNA-seq分析等对MoMHF6的生物学功能进行较系统的研究。【结果】 敲除MoMHF6基因导致稻瘟病菌气生菌丝生长、无性产孢、孢子萌发、子囊壳产生和附着胞形成均显著下降,但与野生菌株相比,ΔMomhf6突变体在CM培养基上的径向生长没有显著差异,在子囊壳中仍可形成子囊和子囊孢子。在1、2和3 mol/L甘油溶液处理条件下,ΔMomhf6突变体附着胞塌陷率显著升高,说明MoMHF6与附着胞膨压形成有关。ΔMomhf6突变体对洋葱内表皮的穿透能力和对感病寄主的致病性完全丧失,甚至在划伤大麦叶片表皮组织中的扩展也受明显限制。而且,ΔMomhf6突变体对氧化胁迫的敏感性增强,附着胞发育过程中糖原的转运和降解缓慢,说明MoMHF6参与氧化胁迫反应和附着胞的糖原代谢。将完整的MoMHF6基因重新导入ΔMomhf6突变体可回补突变体的所有表型缺陷。另外,RNA-seq分析显示,部分已知的稻瘟病菌致病相关基因表达可能受MoMHF6调控,如MoATG4、MoPL1、MoVPR和GAS1等。【结论】 综上所述,稻瘟病菌HSP40编码基因MoMHF6在产孢、附着胞形成、穿透寄主、氧化胁迫应答、致病等过程中起重要作用,研究结果对进一步阐明MoMHF6调控稻瘟病菌形态分化和致病过程的基因网络和分子机制具有重要意义。
童琪, 王春燕, 阙亚伟, 肖宇, 王政逸. 稻瘟病菌热激蛋白(HSP)40编码基因MoMHF6的鉴定及功能研究[J]. 中国水稻科学, 2023, 37(6): 563-576.
TONG Qi, WANG Chunyan, QUE Yawei, XIAO Yu, WANG Zhengyi. Identification and Functional Characterization of the Heat Shock Protein (HSP) 40 Encoding Gene, MoMHF6, in Magnaporthe oryzae[J]. Chinese Journal OF Rice Science, 2023, 37(6): 563-576.
引物 Primer | 序列 Sequence (5′-3′) |
---|---|
MHF6-up-F | CCCCCGGGCTGCAGGAATTCGAGATAACAAAAGGTAT |
MHF6-up-R | GCTCCTTCAATATCATCTTCTCTCGCTCAGTTCGAAATGGGAT |
MHF6-down-F | TAGAGTAGATGCCGACCGAACAAGAATCAATGCCCAGTCTCGTGC |
MHF6-down-R | TACCGGGCCCCCCCTCGAGCGAATACACTTTGGAGAC |
HPT-F | GACAGACGTCGCGGTGAGTT |
HPT-R | GTCCGAGGGCAAAGAAATAG |
MHF6-YW-F | TCGAGGAGATTGAGTGCGTC |
MHF6-YW-R | CTTCTTTTGCTTGGCTTTGC |
MHF6-DX-F | GACTTCAATCAACCCTAACC |
MHF6-DX-R | AAGCCGGACGGAAAGACTTT |
HB-MHF6-F | TCCCCCGGGCTGCAGGAATTCTTCTCCAGAAAATCCCTGGA |
HB-MHF6-R | GATAAGCTTGATATCGAATTCATTCGACCGGCGATCTTCCG |
Actin-RT-F | ATTTACGAGGGTTTCTCCTTGC |
Actin-RT-R | TCTCCTGCTCAAAGTCAAGAG |
HOX2-qRT-F | CGATAATTGCTCCCACACCT |
HOX2-qRT-R | GAAGGAGTCGGTGGTGACAT |
COS1-qRT-F | ATGGATTCCCAGCCTCGTA |
COS1-qRT-R | CGTTGACCAGCAAAGACAA |
HTF1-qRT-F | GGCGACGATACGAAGAAA |
HTF1-qRT-R | TGAACCACCTTGGCTTTG |
CON7-qRT-F | GGCGACGATACGAAGAAA |
CON7-qRT-R | TGAACCACCTTGGCTTTG |
COM1-qRT-F | GAAAGAACCTATCAGGGCG |
COM1-qRT-R | GTTTGCGATTGGCATTAGC |
STU1-qRT-F | CTACGTTAAGTCCGAGATGG |
STU1-qRT-R | CGTGATCAGCCTCATCTTCC |
表1 本研究中使用的引物
Table 1. Primers used in this study.
引物 Primer | 序列 Sequence (5′-3′) |
---|---|
MHF6-up-F | CCCCCGGGCTGCAGGAATTCGAGATAACAAAAGGTAT |
MHF6-up-R | GCTCCTTCAATATCATCTTCTCTCGCTCAGTTCGAAATGGGAT |
MHF6-down-F | TAGAGTAGATGCCGACCGAACAAGAATCAATGCCCAGTCTCGTGC |
MHF6-down-R | TACCGGGCCCCCCCTCGAGCGAATACACTTTGGAGAC |
HPT-F | GACAGACGTCGCGGTGAGTT |
HPT-R | GTCCGAGGGCAAAGAAATAG |
MHF6-YW-F | TCGAGGAGATTGAGTGCGTC |
MHF6-YW-R | CTTCTTTTGCTTGGCTTTGC |
MHF6-DX-F | GACTTCAATCAACCCTAACC |
MHF6-DX-R | AAGCCGGACGGAAAGACTTT |
HB-MHF6-F | TCCCCCGGGCTGCAGGAATTCTTCTCCAGAAAATCCCTGGA |
HB-MHF6-R | GATAAGCTTGATATCGAATTCATTCGACCGGCGATCTTCCG |
Actin-RT-F | ATTTACGAGGGTTTCTCCTTGC |
Actin-RT-R | TCTCCTGCTCAAAGTCAAGAG |
HOX2-qRT-F | CGATAATTGCTCCCACACCT |
HOX2-qRT-R | GAAGGAGTCGGTGGTGACAT |
COS1-qRT-F | ATGGATTCCCAGCCTCGTA |
COS1-qRT-R | CGTTGACCAGCAAAGACAA |
HTF1-qRT-F | GGCGACGATACGAAGAAA |
HTF1-qRT-R | TGAACCACCTTGGCTTTG |
CON7-qRT-F | GGCGACGATACGAAGAAA |
CON7-qRT-R | TGAACCACCTTGGCTTTG |
COM1-qRT-F | GAAAGAACCTATCAGGGCG |
COM1-qRT-R | GTTTGCGATTGGCATTAGC |
STU1-qRT-F | CTACGTTAAGTCCGAGATGG |
STU1-qRT-R | CGTGATCAGCCTCATCTTCC |
图1 MoMhf6与部分真菌同源蛋白的系统发育分析 通过MEGA 7.0 软件对包括稻瘟病菌在内的9个真菌的HSP40同源蛋白构建系统进化树。进化树节点上的数字代表1000株树中有百分之几的树具有这根树枝,即自展值,反映了该树枝的可信度。树枝的长短代表遗传距离。进化树的距离标尺为0.05。
Fig. 1. Phylogenetic analysis of MoMhf6 and its orthologues from several fungal species. The phylogenetic tree was constructed by MEGA version 7.0. Numbers at nodes of the branch represent bootstrapping value in 1000 replications. The length of the branches represents genetic distance. The distance scale=0.05.
图2 MoMHF6基因敲除示意图及ΔMomhf6突变体和互补转化子的PCR验证 A―敲除载体pKO-MoMHF6及MoMHF6靶标基因的敲除,红箭头表示靶基因MoMHF6,蓝箭头代表耐潮霉素基因,图中引物位置被标出;B―确认缺失突变体ΔMomhf6和回补菌株ΔMomhf6-C,左图:用MHF6-YW-F/R引物扩增出524 bp的MoMHF6基因片段;右图:用MHF6-DX-F/R引物验证不同菌株靶向基因缺失或回补,1.5 kb和3.14 kb的条带分别对应HPH和MoMHF6。M−Marker; 泳道1~4分别代表Guy11, ΔMomhf6, MoMHF6-ect(ectopic)和ΔMomhf6-C (complemented)。
Fig. 2. Schematic diagram of gene knockout of MoMHF6 and confirmation of ΔMomhf6 mutants and complemented transformants by PCR. A, Knockout vector pKO-MoMHF6 and target gene deletion of MoMHF6. The red arrow represents the target gene MoMHF6. The blue arrow represents the hygromycin resistant gene. The location of primers is indicated; B, Confirmation of the deletion mutant ΔMomhf6 and the complemented strain ΔMomhf6-C. Left: A 524 bp MoMHF6 gene fragment was amplified by primers MHF6-YW-F/R. Right: the targeted gene deletion or complementation events of different strains were further verified by primers MHF6-DX-F/R. The bands of about 1.5 kb and 3.14 kb were corresponding with HPH and MoMHF6 genes respectively. M, Marker; Lane 1, Guy11; Lane 2, ΔMomhf6; Lane 3, MoMHF6-ect (ectopic); Lane 4, ΔMomhf6-C (complemented).
图3 MoMHF6是气生菌丝生长和子囊壳产生所必需的 A−将野生型Guy11、ΔMomhf6突变体、异位插入菌株MoMHF6-ect和回补菌株ΔMomhf6-C在CM板上25℃下培养10 d,接种10 d后拍照;B−柱状图代表不同菌株在CM培养基上的径向生长,误差线表示标准误;C−ΔMomhf6×TH3杂交组合的子囊壳显著减少,在两种杂交组合中均能观察到子囊和子囊孢子,比例尺为20 μm。
Fig. 3. MoMHF6 is required for aerial hypha growth and perithecia production. A, The wild type Guy11, the ΔMomhf6 mutant, the ectopic insertion strain MoMHF6-ect and the complemented strain ΔMomhf6-C were cultured on CM plates at 25 ℃ for 10 days and photographed at 10 days after inoculation; B, The bar graph shows radical growth of different strains on CM plates. Error bars represent standard error; C, Perithecia production was severely reduced for the ΔMomhf6×TH3 cross. Asci and ascospores in perithecia of both crosses were observed. Bar=20 μm.
图4 MoMHF6在稻瘟病菌致病过程中具有重要作用 A―ΔMomhf6突变体对易感病的大麦叶片无致病性。将不同菌株的分生孢子悬液(浓度为5×104 个/mL)点接于大麦叶段,接种5 d后拍照;B―ΔMomhf6突变体对易感病的水稻叶片无致病性。将不同菌株的分生孢子悬液(浓度为5×104 个/mL)喷雾接种于水稻幼苗上,接种5 d后拍照;C―ΔMomhf6突变体在划伤大麦叶片上的侵染生长受到明显抑制。不同菌株的分生孢子悬液(浓度为5×104个/mL)点接于划伤的大麦叶段上,CK表示H2O,接种5 d后拍照;D―ΔMomhf6突变体不能穿透洋葱表皮质层和细胞壁。将不同菌株的分生孢子悬液(浓度为2×104 个/mL)点接于洋葱表皮上,在黑暗条件下诱导24 h和48 h后,在光学显微镜下进行镜检和拍照,比例尺为20 μm。
Fig. 4. MoMHF6 plays important roles in pathogenesis by M. oryzae. A, The ΔMomhf6 mutant is nonpathogenic to susceptible barley leaves. Conidial suspension (5×104 conidia/mL) of different tested strains were drop-inoculated onto barley leaf segments and photographed at 5 dpi (days post inoculation); B, The ΔMomhf6 mutant is nonpathogenic to susceptible rice leaves. Rice seedlings were sprayed with conidial suspension (5×104 conidia/mL) of different strains and photographed at 5 dpi; C, Invasive growth of the ΔMomhf6 mutant was severely impaired on wounded barley leaf segments. Conidial suspension (5×104 conidia/mL) of different strains was dropped on wounded barley leaf segments. CK represents H2O. Photographs were taken at 5 dpi; D, The ΔMomhf6 mutant was unable to penetrate onion cuticle and cell wall. Conidial suspension (2×104 conidia/mL) of different strains was dropped on onion epidermis. After 24 h and 48 h of incubation in darkness, the onion epidermis was examined and photographed under a light microscope. Bars=20 μm.
图5 MoMHF6在稻瘟病菌无性产孢中起重要作用 A―显微镜下观察气生菌丝生长和分生孢子梗发育,比例尺为100 μm;B―25℃条件下,不同菌株在CM培养基上培养12 d后统计分析产孢量;C―qRT-PCR分析几个产孢相关基因转录表达水平,误差线表示标准误,星号表示显著差异(*P<0.05; **P<0.01)。
Fig. 5. MoMHF6 is important for asexual sporulation. A, Microscopic observation of aerial hypha growth and conidiophore development. Bars=100 μm; B, Statistical analysis of conidiation on CM at 25 ℃ for 12 days; C, qRT-PCR analysis of transcriptional expression of several sporulation related genes. Error bars represent standard error. Asterisks indicate significant difference (*P<0.05; **P<0.01).
图6 敲除MoMHF6导致分生孢子萌发、附着胞形成和附着胞膨压积累显著降低 A―统计分析分生孢子萌发率,分生孢子在疏水表面诱导萌发6 h;B―统计分析附着胞形成率,将分生孢子置于疏水表面诱导形成附着胞,黑暗诱导24 h;C―统计分析附着胞塌陷率,用1 mol/L、2 mol/L、3 mol/L的甘油溶液处理诱导24 h形成的附着胞,处理时间10 min,误差线表示标准误,星号表示显著差异(*P<0.05; **P<0.01)。
Fig. 6. Deletion of MoMHF6 results in significant reduction of conidium germination, appressorium formation and appressorium turgor generation A, Statistical analysis of conidium germination. Conidia were allowed to germinate on hydrophobic surface for 6 hours; B, Statistical analysis of appressorium formation. Conidia were placed on the hydrophobic surface to induce appressorium formation and incubated for 24 hours; C, Statistical analysis of collapsed appressoria. Appressoria induced for 24 hours were incubated in 1 mol/L, 2 mol/L or 3 mol/L glycerol solution for 10 minutes. Error bars represent standard error. Asterisks indicate significant difference (*P<0.05; **P<0.01).
图7 MoMHF6基因的缺失导致稻瘟病菌附着胞发育过程中糖原转运和降解延缓 A-将野生型Guy11、ΔMomhf6和回补菌株ΔMomhf6-C的分生孢子悬液(浓度为5×104 个/mL)分别在疏水表面诱导4、8、12、24 h后,用KI/I2溶液进行染色,并在不同的时间间隔拍照,比例尺为20 μm;B-统计各待测菌株在不同时间段分生孢子内的糖原转移情况;C-统计各待测菌株在不同时间段附着胞内的糖原降解情况。误差线表示标准误,星号表示显著差异(P<0.05)。
Fig. 7. Deletion of MoMHF6 delays glycogen transportation and degradation during appressoria development in M. oryzae. A, Conidial suspension (5×104 conidia/mL) of the wild type Guy11, the mutant ΔMomhf6 and the complemented strain ΔMomhf6-C was incubated on hydrophobic surface for 4, 8, 12, 24 hours respectively and then stained with KI/I2 solution. Photographs were taken at different time intervals. Bars=20 μm. B, Statistical analysis of glycogen mobilization in conidia of different tested strains; C, Statistical analysis of glycogen degradation in appressoria of different tested strains. Error bars represent standard error. Asterisks indicate significant difference (P<0.05).
图8 MoMHF6缺失增强稻瘟病菌对氧化胁迫的敏感性 A-将不同菌株分别接种在含有2.5、5和10 mmol/L H2O2的CM培养基上,培养10 d后观察菌落形态;B-测定菌落直径并计算生长抑制率,误差线表示标准误,星号表示显著差异 (*P<0.05; **P < 0.01)。
Fig. 8. Deletion of MoMHF6 increases sensitivity of M. oryzae to oxidative stress. A, The different tested strains were inoculated on CM supplemented with 2.5, 5 and 10 mmol/L H2O2, respectively. Growth of various tested strains after 10 days is presented; B, Colony diameters were measured and growth inhibition rates were calculated. Error bars represent standard deviation. Asterisks indicate significant difference (*P<0.05; **P<0.01).
图9 ΔMomhf6突变体转录组差异表达基因分析 A—ΔMomhf6差异表达基因数量的统计分析;B—ΔMomhf6突变体DEGs的GO富集散点图,富集比率表示GO项中标注的DEGs数量与同一GO项中标注的所有基因数量的比值,q值是修正后的p值,q值越小表示集中度越高。
Fig. 9. Transcriptome analysis of the ΔMomhf6 mutant for differentially expressed genes. A, Statistical analysis of numbers of differentially expressed genes in the ΔMomhf6 mutant in comparison to the wild type strain Guy11; B, Scatter plot of GO term enrichment of DEGs in the ΔMomhf6 mutant. Rich ratio represents the ratio of numbers of DEGs annotated in the GO term to the numbers of all genes annotated in the same GO term. The q-value is corrected p value, with lower value means greater intensiveness.
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