中国水稻科学 ›› 2024, Vol. 38 ›› Issue (3): 256-265.DOI: 10.16819/j.1001-7216.2024.231012
尹潇潇, 张芷菡, 颜绣莲, 廖蓉, 杨思葭, 郭岱铭, 樊晶, 赵志学*(), 王文明*(
)
收稿日期:
2023-10-30
修回日期:
2023-12-12
出版日期:
2024-05-10
发布日期:
2024-05-13
通讯作者:
*email: zhixuezhao@sicau.edu.cn;
j316wenmingwang@163.com
基金资助:
YIN Xiaoxiao, ZHANG Zhihan, YAN Xiulian, LIAO Rong, YANG Sijia, Beenish HASSAN, GUO Daiming, FAN Jing, ZHAO Zhixue*(), WANG Wenming*(
)
Received:
2023-10-30
Revised:
2023-12-12
Online:
2024-05-10
Published:
2024-05-13
Contact:
*email: zhixuezhao@sicau.edu.cn;
j316wenmingwang@163.com
摘要:
【目的】信号肽对效应因子的分泌至关重要,影响病原微生物的致病进程。前期鉴定到在稻曲病菌侵染水稻花器官过程中显著上调表达的10个效应因子基因,但这些效应因子是否含有功能性信号肽及其表达模式尚不明确。【方法】利用SignaIP-5.0数据库预测、烟草瞬时表达系统和酵母信号肽分泌实验,分析并鉴定这些效应因子的信号肽。利用实时荧光定量PCR技术,检测效应因子基因的表达模式。【结果】UV_44、UV_1548、UV_1567等10个稻曲病菌效应因子均含有信号肽。烟草瞬时表达结果显示,含有信号肽的eYFP融合蛋白在细胞质壁分离后均能分泌到质外体空间。酵母菌信号肽分泌结果显示,分别携带这10个信号肽的酵母菌株可在以棉子糖为唯一碳源的培养基上生长,且与TTC发生显色反应。实时荧光定量结果表明,除了UV_6754,其余9个效应因子基因的表达模式不尽相同,其中UV_44、UV_1567、UV_3667、UV_4213、UV_4989、UV_5215和UV_8184的表达量在接种后显著升高。【结论】这10个稻曲病菌效应因子均含有分泌功能的信号肽,编码效应因子的基因在稻曲病菌侵染过程中具有不同的表达模式。该结果为深入研究以上稻曲病菌效应因子的功能奠定了基础。
尹潇潇, 张芷菡, 颜绣莲, 廖蓉, 杨思葭, 郭岱铭, 樊晶, 赵志学, 王文明. 多个稻曲病菌效应因子的信号肽验证和表达分析[J]. 中国水稻科学, 2024, 38(3): 256-265.
YIN Xiaoxiao, ZHANG Zhihan, YAN Xiulian, LIAO Rong, YANG Sijia, Beenish HASSAN, GUO Daiming, FAN Jing, ZHAO Zhixue, WANG Wenming. Signal Peptide Validation and Expression Analysis of Multiple Effectors from Ustilaginoidea virens[J]. Chinese Journal OF Rice Science, 2024, 38(3): 256-265.
引物名称 Primer name | 引物序列(5′-3′) Primer sequence(5′-3′) | 用途 Usage | |
---|---|---|---|
Psuc2-44-F | AATTCATGAAGGCCTCGACGCTGCTGGCTCTCCTGCCCTTTGCCGCGGCCC | 酵母系统鉴定信号肽功能 Functional validation of the signal peptides using yeast | |
Psuc2-44-R | TCGAGGGCCGCGGCAAAGGGCAGGAGAGCCAGCAGCGTCGAGGCCTTCATG | ||
Psuc2-1548-F | AATTCATGAACCCAATCCTGCTTCTCCTCGCCGCATCAGTCCCCGCCCGCAGCC | ||
Psuc2-1548-R | TCGAGGCTGCGGGCGGGGACTGATGCGGCGAGGAGAAGCAGGATTGGGTTCATG | ||
Psuc2-1567-F | AATTCATGCAGTTGTCCTACCTTCTCTCGCTCGTCTCCCTTGTGGCCGCTC | ||
Psuc2-1567-R | TCGAGAGCGGCCACAAGGGAGACGAGCGAGAGAAGGTAGGACAACTGCATG | ||
Psuc2-1990-F | AATTCATGCGCCCAGCAACCGTCATCTTCGCACTGGCCGTCTCCGTCAAGGCTC | ||
Psuc2-1990-R | TCGAGAGCCTTGACGGAGACGGCCAGTGCGAAGATGACGGTTGCTGGGCGCATG | ||
Psuc2-3667-F | AATTCATGAAGTACACTGCTGCTCTCGTCGCCCTGGCTGCTGCCGTCATGGCCC | ||
Psuc2-3667-R | TCGAGGGCCATGACGGCAGCAGCCAGGGCGACGAGAGCAGCAGTGTACTTCATG | ||
Psuc2-4213-F | AATTCATGATTCTGCTGCTCCTCTTGCTATGCACGACAACATGTTTAGGAC | ||
Psuc2-4213-R | TCGAGTCCTAAACATGTTGTCGTGCATAGCAAGAGGAGCAGCAGAATCATG | ||
Psuc2-4989-F | AATTCATGCACCAGCTCCTTGCACTACTCCTCGCCAGCCTCACCGCTGCCC | ||
Psuc2-4989-R | TCGAGGGCAGCGGTGAGGCTGGCGAGGAGTAGTGCAAGGAGCTGGTGCATG | ||
Psuc2-5215-F | AATTCATGAAGTTTATTTCCCTCGTAGCCGCGGCCGGTGTGGCGTATGCCC | ||
Psuc2-5215-R | TCGAGGGCATACGCCACACCGGCCGCGGCTACGAGGGAAATAAACTTCATG | ||
Psuc2-6754-F | AATTCATGGCTGCAGCATCAGCCCTTTTCTTCCTCACGGCGGCTGTCTCCTGGACAC | ||
Psuc2-6754-R | TCGAGTGTCCAGGAGACAGCCGCCGTGAGGAAGAAAAGGGCTGATGCTGCAGCCATG | ||
Psuc2-8184-F | AATTCATGCAAATCGTCCTCCCTCTCCTGGCCTGCGCAGCCGTGGCTGCCGCCC | ||
Psuc2-8184-R | TCGAGGGCGGCAGCCACGGCTGCGCAGGCCAGGAGAGGGAGGACGATTTGCATG | ||
P1300-44-F | CATGAAGGCCTCGACGCTGCTGGCTCTCCTGCCCTTTGCCGCGGCCGGTAC | 烟草瞬时表达鉴定信号肽功能 Functional validation of the signal peptides using N. benthamiana | |
P1300-44-R | CGGCCGCGGCAAAGGGCAGGAGAGCCAGCAGCGTCGAGGCCTTCATGGTAC | ||
P1300-1548-F | CATGAACCCAATCCTGCTTCTCCTCGCCGCATCAGTCCCCGCCCGCAGCGGTAC | ||
P1300-1548-R | CGCTGCGGGCGGGGACTGATGCGGCGAGGAGAAGCAGGATTGGGTTCATGGTAC | ||
P1300-1567-F | CATGCAGTTGTCCTACCTTCTCTCGCTCGTCTCCCTTGTGGCCGCTGGTAC | ||
P1300-1567-R | CAGCGGCCACAAGGGAGACGAGCGAGAGAAGGTAGGACAACTGCATGGTAC | ||
P1300-1990-F | CATGCGCCCAGCAACCGTCATCTTCGCACTGGCCGTCTCCGTCAAGGCTGGTAC | ||
P1300-1990-R | CAGCCTTGACGGAGACGGCCAGTGCGAAGATGACGGTTGCTGGGCGCATGGTAC | ||
P1300-3667-F | CATGAAGTACACTGCTGCTCTCGTCGCCCTGGCTGCTGCCGTCATGGCCGGTAC | ||
P1300-3667-R | CGGCCATGACGGCAGCAGCCAGGGCGACGAGAGCAGCAGTGTACTTCATGGTAC | ||
P1300-4213-F | CATGATTCTGCTGCTCCTCTTGCTATGCACGACAACATGTTTAGGAGGTAC | ||
P1300-4213-R | CTCCTAAACATGTTGTCGTGCATAGCAAGAGGAGCAGCAGAATCATGGTAC | ||
P1300-4989-F | CATGCACCAGCTCCTTGCACTACTCCTCGCCAGCCTCACCGCTGCCGGTAC | ||
P1300-4989-R | CGGCAGCGGTGAGGCTGGCGAGGAGTAGTGCAAGGAGCTGGTGCATGGTAC | ||
P1300-5215-F | CATGAAGTTTATTTCCCTCGTAGCCGCGGCCGGTGTGGCGTATGCCGGTAC | ||
P1300-5215-R | CGGCATACGCCACACCGGCCGCGGCTACGAGGGAAATAAACTTCATGGTAC | ||
P1300-6754-F | CATGGCTGCAGCATCAGCCCTTTTCTTCCTCACGGCGGCTGTCTCCTGGACAGGTAC | ||
P1300-6754-R | CTGTCCAGGAGACAGCCGCCGTGAGGAAGAAAAGGGCTGATGCTGCAGCCATGGTAC | ||
P1300-8184-F | CATGCAAATCGTCCTCCCTCTCCTGGCCTGCGCAGCCGTGGCTGCCGCCGGTAC | ||
P1300-8184-R | CGGCGGCAGCCACGGCTGCGCAGGCCAGGAGAGGGAGGACGATTTGCATGGTAC | ||
Uv_Tublin2α-2F/R | ACCAGCTCGTTGAGAACTCG / AATCAGAGTTGAGCTGGCCG | 实时荧光定量PCR RT-qPCR | |
UV_44-RT-F/R | GCGTTGCCAAGAAGACCAAG / ATTCCAGCAATCACGGCAGA | ||
UV_1548-RT- F/R | GACAACGGGCAGACAAACTG / CTAGCAATCCCAGCAGTCGT | ||
UV_1567-RT-F/R | TGCTAGCGCTATCGTTCCTG / GTCGGCATGGTTTTGCAACT | ||
UV_1990-RT-F/R | ACTCACTTCCAAACACGGCA / GCCTTGTCTTCTCAACACGC | ||
UV_3667-RT-F/R | GCCTCCTCTGCAATGTGGAA / CTTGAGGCAGTTGAGTCCGA | ||
UV_4213-RT-F/R | ATGCGAGAACTTGGTCGTGT / AACCAACGCCTCCAACTTGA | ||
UV_4989-RT-F/R | CCTGCCTCCTTCGTCCAAAT / GGTCGATCTGGCAAAGTGGA | ||
UV_5215-RT-F/R | TAGCGTACTGCGACGGTAAC / GAAACAGCGCCCAATTCCTG | ||
UV_6754-RT-F/R | AAATGCGAGGACTAGGGCAC / CTTTTTGCCGGAGGAGGTCT | ||
UV_8184-RT-F/R | CATCGCTGCGAAGACAAAGG / GCGCCTGTGAACTTTTCCTG |
表1 本研究所需引物
Table 1. Primers used in this study
引物名称 Primer name | 引物序列(5′-3′) Primer sequence(5′-3′) | 用途 Usage | |
---|---|---|---|
Psuc2-44-F | AATTCATGAAGGCCTCGACGCTGCTGGCTCTCCTGCCCTTTGCCGCGGCCC | 酵母系统鉴定信号肽功能 Functional validation of the signal peptides using yeast | |
Psuc2-44-R | TCGAGGGCCGCGGCAAAGGGCAGGAGAGCCAGCAGCGTCGAGGCCTTCATG | ||
Psuc2-1548-F | AATTCATGAACCCAATCCTGCTTCTCCTCGCCGCATCAGTCCCCGCCCGCAGCC | ||
Psuc2-1548-R | TCGAGGCTGCGGGCGGGGACTGATGCGGCGAGGAGAAGCAGGATTGGGTTCATG | ||
Psuc2-1567-F | AATTCATGCAGTTGTCCTACCTTCTCTCGCTCGTCTCCCTTGTGGCCGCTC | ||
Psuc2-1567-R | TCGAGAGCGGCCACAAGGGAGACGAGCGAGAGAAGGTAGGACAACTGCATG | ||
Psuc2-1990-F | AATTCATGCGCCCAGCAACCGTCATCTTCGCACTGGCCGTCTCCGTCAAGGCTC | ||
Psuc2-1990-R | TCGAGAGCCTTGACGGAGACGGCCAGTGCGAAGATGACGGTTGCTGGGCGCATG | ||
Psuc2-3667-F | AATTCATGAAGTACACTGCTGCTCTCGTCGCCCTGGCTGCTGCCGTCATGGCCC | ||
Psuc2-3667-R | TCGAGGGCCATGACGGCAGCAGCCAGGGCGACGAGAGCAGCAGTGTACTTCATG | ||
Psuc2-4213-F | AATTCATGATTCTGCTGCTCCTCTTGCTATGCACGACAACATGTTTAGGAC | ||
Psuc2-4213-R | TCGAGTCCTAAACATGTTGTCGTGCATAGCAAGAGGAGCAGCAGAATCATG | ||
Psuc2-4989-F | AATTCATGCACCAGCTCCTTGCACTACTCCTCGCCAGCCTCACCGCTGCCC | ||
Psuc2-4989-R | TCGAGGGCAGCGGTGAGGCTGGCGAGGAGTAGTGCAAGGAGCTGGTGCATG | ||
Psuc2-5215-F | AATTCATGAAGTTTATTTCCCTCGTAGCCGCGGCCGGTGTGGCGTATGCCC | ||
Psuc2-5215-R | TCGAGGGCATACGCCACACCGGCCGCGGCTACGAGGGAAATAAACTTCATG | ||
Psuc2-6754-F | AATTCATGGCTGCAGCATCAGCCCTTTTCTTCCTCACGGCGGCTGTCTCCTGGACAC | ||
Psuc2-6754-R | TCGAGTGTCCAGGAGACAGCCGCCGTGAGGAAGAAAAGGGCTGATGCTGCAGCCATG | ||
Psuc2-8184-F | AATTCATGCAAATCGTCCTCCCTCTCCTGGCCTGCGCAGCCGTGGCTGCCGCCC | ||
Psuc2-8184-R | TCGAGGGCGGCAGCCACGGCTGCGCAGGCCAGGAGAGGGAGGACGATTTGCATG | ||
P1300-44-F | CATGAAGGCCTCGACGCTGCTGGCTCTCCTGCCCTTTGCCGCGGCCGGTAC | 烟草瞬时表达鉴定信号肽功能 Functional validation of the signal peptides using N. benthamiana | |
P1300-44-R | CGGCCGCGGCAAAGGGCAGGAGAGCCAGCAGCGTCGAGGCCTTCATGGTAC | ||
P1300-1548-F | CATGAACCCAATCCTGCTTCTCCTCGCCGCATCAGTCCCCGCCCGCAGCGGTAC | ||
P1300-1548-R | CGCTGCGGGCGGGGACTGATGCGGCGAGGAGAAGCAGGATTGGGTTCATGGTAC | ||
P1300-1567-F | CATGCAGTTGTCCTACCTTCTCTCGCTCGTCTCCCTTGTGGCCGCTGGTAC | ||
P1300-1567-R | CAGCGGCCACAAGGGAGACGAGCGAGAGAAGGTAGGACAACTGCATGGTAC | ||
P1300-1990-F | CATGCGCCCAGCAACCGTCATCTTCGCACTGGCCGTCTCCGTCAAGGCTGGTAC | ||
P1300-1990-R | CAGCCTTGACGGAGACGGCCAGTGCGAAGATGACGGTTGCTGGGCGCATGGTAC | ||
P1300-3667-F | CATGAAGTACACTGCTGCTCTCGTCGCCCTGGCTGCTGCCGTCATGGCCGGTAC | ||
P1300-3667-R | CGGCCATGACGGCAGCAGCCAGGGCGACGAGAGCAGCAGTGTACTTCATGGTAC | ||
P1300-4213-F | CATGATTCTGCTGCTCCTCTTGCTATGCACGACAACATGTTTAGGAGGTAC | ||
P1300-4213-R | CTCCTAAACATGTTGTCGTGCATAGCAAGAGGAGCAGCAGAATCATGGTAC | ||
P1300-4989-F | CATGCACCAGCTCCTTGCACTACTCCTCGCCAGCCTCACCGCTGCCGGTAC | ||
P1300-4989-R | CGGCAGCGGTGAGGCTGGCGAGGAGTAGTGCAAGGAGCTGGTGCATGGTAC | ||
P1300-5215-F | CATGAAGTTTATTTCCCTCGTAGCCGCGGCCGGTGTGGCGTATGCCGGTAC | ||
P1300-5215-R | CGGCATACGCCACACCGGCCGCGGCTACGAGGGAAATAAACTTCATGGTAC | ||
P1300-6754-F | CATGGCTGCAGCATCAGCCCTTTTCTTCCTCACGGCGGCTGTCTCCTGGACAGGTAC | ||
P1300-6754-R | CTGTCCAGGAGACAGCCGCCGTGAGGAAGAAAAGGGCTGATGCTGCAGCCATGGTAC | ||
P1300-8184-F | CATGCAAATCGTCCTCCCTCTCCTGGCCTGCGCAGCCGTGGCTGCCGCCGGTAC | ||
P1300-8184-R | CGGCGGCAGCCACGGCTGCGCAGGCCAGGAGAGGGAGGACGATTTGCATGGTAC | ||
Uv_Tublin2α-2F/R | ACCAGCTCGTTGAGAACTCG / AATCAGAGTTGAGCTGGCCG | 实时荧光定量PCR RT-qPCR | |
UV_44-RT-F/R | GCGTTGCCAAGAAGACCAAG / ATTCCAGCAATCACGGCAGA | ||
UV_1548-RT- F/R | GACAACGGGCAGACAAACTG / CTAGCAATCCCAGCAGTCGT | ||
UV_1567-RT-F/R | TGCTAGCGCTATCGTTCCTG / GTCGGCATGGTTTTGCAACT | ||
UV_1990-RT-F/R | ACTCACTTCCAAACACGGCA / GCCTTGTCTTCTCAACACGC | ||
UV_3667-RT-F/R | GCCTCCTCTGCAATGTGGAA / CTTGAGGCAGTTGAGTCCGA | ||
UV_4213-RT-F/R | ATGCGAGAACTTGGTCGTGT / AACCAACGCCTCCAACTTGA | ||
UV_4989-RT-F/R | CCTGCCTCCTTCGTCCAAAT / GGTCGATCTGGCAAAGTGGA | ||
UV_5215-RT-F/R | TAGCGTACTGCGACGGTAAC / GAAACAGCGCCCAATTCCTG | ||
UV_6754-RT-F/R | AAATGCGAGGACTAGGGCAC / CTTTTTGCCGGAGGAGGTCT | ||
UV_8184-RT-F/R | CATCGCTGCGAAGACAAAGG / GCGCCTGTGAACTTTTCCTG |
基因名称 Gene name | 蛋白大小(氨基酸) Length of protein (aa) | 信号肽长度(氨基酸) Length of signal peptide (aa) | 信号肽评分 Signal peptide scoring(Sec/SPI) |
---|---|---|---|
UV_44 | 386 | 15 | 0.8501 |
UV_1548 | 104 | 16 | 0.9929 |
UV_1567 | 388 | 15 | 0.9722 |
UV_1990 | 279 | 16 | 0.9845 |
UV_3667 | 89 | 16 | 0.9995 |
UV_4213 | 216 | 15 | 0.9642 |
UV_4989 | 231 | 15 | 0.9924 |
UV_5215 | 88 | 15 | 0.9981 |
UV_6754 | 211 | 17 | 0.4707 |
UV_8184 | 106 | 16 | 0.9982 |
表2 10个稻瘟病菌效应因子的信号肽预测
Table 2. Signal peptide prediction of ten effectors from U. virens
基因名称 Gene name | 蛋白大小(氨基酸) Length of protein (aa) | 信号肽长度(氨基酸) Length of signal peptide (aa) | 信号肽评分 Signal peptide scoring(Sec/SPI) |
---|---|---|---|
UV_44 | 386 | 15 | 0.8501 |
UV_1548 | 104 | 16 | 0.9929 |
UV_1567 | 388 | 15 | 0.9722 |
UV_1990 | 279 | 16 | 0.9845 |
UV_3667 | 89 | 16 | 0.9995 |
UV_4213 | 216 | 15 | 0.9642 |
UV_4989 | 231 | 15 | 0.9924 |
UV_5215 | 88 | 15 | 0.9981 |
UV_6754 | 211 | 17 | 0.4707 |
UV_8184 | 106 | 16 | 0.9982 |
图1 载体的构建及阳性克隆的鉴定 A: pCAMBIA1300-35S-eYFP-RBS和pSUC2质粒示意图;B: PCR和琼脂糖凝胶电泳鉴定阳性克隆。
Fig. 1. Construction of vectors and identification of the positive clones A, Diagrams of the pCAMBIA1300-35S-eYFP-RBS and pSUC2 plasmids; B, Positive clones were identified by PCR and agarose gel electrophoresis.
图2 烟草瞬时表达分析10个稻瘟病菌效应因子信号肽的亚细胞定位0.8 mol/L甘露醇诱导质壁分离前(A)和分离后(B)融合蛋白的亚细胞定位。注射农杆菌36 h后拍摄。EV、AvrL567SP分别为阴性和阳性对照。
Fig. 2. Subcellular localization of the transiently expressed signal peptides of ten effectors from U. virens in N. benthamiana Subcellular localization of the fusion proteins containing the predicted signal peptide before (A) and after (B) plasmolysis induced by 0.8 mol/L mannitol. The images were captured at 36 h after infiltration. EV was used as a negative control, and AvrL567SP was used as a positive control.
图3 利用酵母转化酶实验验证信号肽的分泌功能 A: 样品位置示意图;B, C, D: 酵母信号肽筛选实验验证信号肽的分泌功能;E: TTC显色实验验证信号肽的分泌功能。pSUC2-Avr1b为阳性对照,pSUC2-Mg87和YTK12为阴性对照。
Fig. 3. Functional evaluation of the signal peptides using the yeast invertase assay A, Diagrams showing the sample location on Petri plates; B, C and D, Detection of the secretion of signal peptides by the yeast signal trap assay; E, TTC assay of the signal peptides. pSUC2-Avr1b was used as a positive control. YTK12 and pSUC2-Mg87 were used as the negative controls.
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