CN106754977A - 稻瘟菌农药靶标基因MoR1及其编码蛋白和应用 - Google Patents

稻瘟菌农药靶标基因MoR1及其编码蛋白和应用 Download PDF

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CN106754977A
CN106754977A CN201611140339.7A CN201611140339A CN106754977A CN 106754977 A CN106754977 A CN 106754977A CN 201611140339 A CN201611140339 A CN 201611140339A CN 106754977 A CN106754977 A CN 106754977A
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康厚祥
刘文德
胡培
王
王一
吴奇
李成云
王国梁
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Abstract

本发明属于功能靶标基因的克隆与利用领域，具体公开了稻瘟菌MoR1基因的核苷酸序列以及该基因编码的蛋白质。此外，本发明还公开了该基因序列及其编码蛋白作为稻瘟菌农药靶点，尤其是作为稻瘟菌多抗真菌农药的用途。

Description

稻瘟菌农药靶标基因MoR1及其编码蛋白和应用
技术领域
本发明属于功能靶标基因的克隆与利用领域，具体公开了稻瘟菌的MoR1基因序列及其应用。
背景技术
病原真菌可通过多种机制对真菌农药产生抗性，包括1：解毒，真菌体内迅速产生抗体，通过与农药结合使其丧失抗性；2代谢,在农药结合到其作用靶标之前，通过代谢作用迅速将其降解；3：迅速排出，真菌细胞在农药还未达到其作用标靶时便迅速排至细胞外；4：降低农药的吸收，通过改变吸收系统相关基因可以减少农药的吸收量；5：靶标位点的突变，通过突变农药作用靶标位点，使得药物彻底丧失功能。目前杀真菌农药主要针对真菌中细胞膜或者细胞内亚细胞器中一些非常重要的作用靶标来设计。其中甲氧基丙烯酸酯类(Strobiluin)类广谱杀真菌药物是目前应用最为广泛的杀真菌农药，其通过作用于线粒体中的电子传递链来达到广谱杀真菌的作用，其代表药物如：吡唑醚菌酯和嘧菌酯等。但，稻瘟菌群体中已对甲氧基丙烯酸酯类杀菌剂具有较高抗性的菌株。稻瘟病病是危害水稻最严重的病害之一，造成世界范围内水稻平均减产10％-30％(Skamnioti P et al,2009)给粮食安全带来严重威胁。水稻是中国第一大粮食作物，中国稻瘟病年均发生面积380万hm2以上，造成产量年损失数亿公斤(杨勤忠等，2009)利用真菌农药控制和预防稻瘟病是减少损失的有效途径之一，而耐药性稻瘟菌菌株的出现为稻瘟病的可持续控制带来新的挑战，本研发明中找到的耐药性关键基因MoR1，可为持续控制稻瘟病提供关键靶标基因。
参考文献
Skamnioti P,Gurr SJ(2009)Against the grain:safeguarding rice fromrice blast disease.Trends in biotechnology 27:141-150.
杨勤忠，林菲，冯淑杰，等.水稻稻瘟病抗性基因的分子定位及克隆研究进展。中国农业科学，2009,42(5):1601-1615.
发明内容
本发明目的包括：
提供一种稻瘟菌耐药性的基因序列，以及该基因序列编号的蛋白质；
提供该基因序列及其编码蛋白的用途
提供一种抑制稻瘟菌耐药性的药物靶点，等
本发明首次公开了稻瘟菌MoR1基因序列，其核苷酸序列如序列表SEQ ID NO.1所示；
进一步的，公开了稻瘟菌MoR1基因作为稻瘟菌农药基因靶标的用途；尤其作为耐药性稻瘟菌的农药基因靶标的用途；
以及稻瘟菌MoR1基因作为耐药性稻瘟菌基因靶标的用途。
另一方面，本发明提供了一种稻瘟菌农药基因靶标，其核苷酸序列如序列表SEQID NO.1所示；
本发明还提供了一种稻瘟菌多抗真菌农药基因靶标，其核苷酸序列如序列表SEQID NO.1所示；
所述稻瘟菌优选为耐药性稻瘟菌。
此外，本发明还提供了稻瘟菌MoR1基因编码的蛋白质，其氨基酸序列如序列表SEQID NO.2所示；
进一步的，提供了稻瘟菌MoR1基因编码的蛋白质作为稻瘟菌多抗真菌农药靶点的用途。
本发明还提供了一种稻瘟菌农药靶点，其核苷酸序列如序列表SEQ ID NO.2所示；
以及一种稻瘟菌多抗真菌农药靶点，其核苷酸序列如序列表SEQ ID NO.2所示。
有益效果
稻瘟菌损失的谷物每年可养活约6千万人口，而耐药性稻瘟菌的出现为稻瘟病为目前真菌农药的有效使用带来严重威胁。本发明中的MoR1基因可作为关键的标靶基因，在稻瘟病菌尤其是耐药性稻瘟病菌的可持续防控过程中起关键作用，为粮食安全提供保障。
附图说明
图1：吡唑醚菌酯对不同稻瘟菌菌株生长的抑制比率
图2：嘧菌酯对不同稻瘟菌菌株生长的抑制比率
图3：携带不同MoR1等位基因的三个菌株在两种真菌农药培养条件下的生长情况图。
其中A，B,C分别为三个不同的菌株027,635和029，A1，B1，C1为在0.01PPM吡唑醚菌酯下培养7天后三个菌株的生长情况，A2，B2，C2为在0.01PPM嘧菌酯下培养7天后三个菌株的生长情况。
图4以226菌株为背景菌株，敲除MoR1基因获得的其中两个独立的转化子的PCR检测图
其中，第1到第4列为利用MoR1基因特异的引物扩增检测转化子1和转化子2的情况，第5列为菌株226未敲除阳性对照，第7列为阴性对照，最后一列为MoR1基因回补后的检测结果。
说明：MoR1只编码一个蛋白，转化子1和2是敲除试验中的两个独立的敲除突变体，都是针对MoR1基因的，为了实验严谨，需要两个以上独立转化子的实验一致才认为可靠，也即为平行实验组。
图5a敲除转化子和对照包含完整的MoR1基因的稻瘟菌菌株在0.01ppm吡唑醚菌酯条件下的菌丝生长图
图5b敲除转化子和对照包含完整的MoR1基因的稻瘟菌菌株在0.01ppm吡唑醚菌酯条件下的菌丝生长情况柱状图
图6敲除转化子和对照包含完整的MoR1基因的稻瘟菌菌株在0.01ppm嘧菌酯条件下的菌丝生长图
图7 MoR1基因编码蛋白的3D结构
其中，1为MoR1基因左侧跨膜α-螺旋跨膜结构区域；
2为MoR1基因右侧跨膜α-螺旋跨膜结构区域；
3为MoR1基因转运结合活性功能区域。
具体实施方式
实施例1不同稻瘟菌菌株对真菌农药吡唑醚菌酯和嘧菌酯的抗性水平实验
实验方法和步骤：将两个敲除转化子的菌株与野生型菌株在燕麦培养基上活化，25℃黑暗条件下培养4-7d后，在菌落边缘同一圆周上打取菌饼，分别接种于终浓度为0.01ppm吡唑醚菌酯、0.02ppm嘧菌酯及未经处理的CM培养基上(配方：1L；酵母提取物6g，酸水解酪蛋白3g，酶水解酪蛋白3g，蔗糖10g，琼脂15g)，每个处理每个菌株设3次重复，25℃黑暗条件下培养7d后，用十字交叉法测量各处理条件下各菌株的菌落生长直径并拍照记录。
实验结果如图1、图2所示。图1，图2为不同稻瘟菌菌株对两种主流真菌农药吡唑醚菌酯和嘧菌酯的抗性水平的表型，表明不同菌株对两种农药的抗性水平具有显著的差异。图3为两个敲除转化子的菌株与野生型菌株在两种农药培养条件下的生长情况。
实施例2稻瘟菌MoR1基因的克隆
克隆和测序方法：PCR引物：(F:TTTGGATTTGTGTCTTCTGCC R:TGCATCACTTAATCTTTGCCAC),采用Sanger测序(北京华大基因测序公司)将PCR产物直接测序拼接，MoR1基因的核苷酸序列如序列表SEQ ID NO.1所示。并进一步通过对100株稻瘟菌MoR1基因测序和分析，发现MoR1基因的突变与两种主要农药的抗性敏感性显著相关。MoR1基因编码一个ABC类型的转运子(转运蛋白)。
实施例3MoR1基因的多抗农药功能验证实验
为进一步验证MoR1的抗药性功能，首先选取包含完整的MoR1基因序列、且对两种农药具备一定抗性的菌株226为受体菌株，敲除了MoR1基因并进行了回补验证(回补方法：，利用MoR1基因全长引物F:TTTGGATTTGTGTCTTCTGCC R:TGCATCACTTAATCTTTGCCAC，从菌株125中通过PCR克隆出完整的MoR1基因并转入载体pyK11，通过pyK11转化226转化子，最终将MoR1基因回补到敲除突变体中)。
使用MoR1基因检测引物(引物MoR1-F:AAGTCACTCTTGCTAAAGTC；MoR1-R:CCTTCTGGATGAGATACAGG.)，对226菌株、两个转化子及回补转化子进行PCR扩增。以226菌株为背景菌株，敲除MoR1基因获得的其中两个独立的转化子的PCR检测图及对比图，如说明书附图4所示。
敲除转化子和对照包含完整的MoR1基因的稻瘟菌菌株在0.01ppm吡唑醚菌酯条件下的菌丝生长情况图，培养基为CM完全培养基(配方：1L；酵母提取物6g，酸水解酪蛋白3g，酶水解酪蛋白3g，蔗糖10g，琼脂15g)。实验结果如说明书附图5a、图5b、图6所示；实验表明已成功获得MoR1基因敲除的转化子及MoR1基因回补的转化子。
图5a、图5b表明，226菌株野生型(MoR1+)与敲除转化子在完全培养基生长条件7天后，菌丝直径大小无明显差异；当在完全培养基中包含0.01ppm的吡唑醚菌酯条件下，虽然226菌株野生型(MoR1+)生长也得到一定程度的抑制(图左第一列，图右第一列)，但在两个独立的敲除转化子中(MoR1-)，表现出对0.01ppm的吡唑醚菌酯非常敏感。表明MoR1参与了菌株耐受吡唑醚菌酯的作用。与图5a、图5b类似，图6中，两个独立的敲除转化子在0.01ppm嘧菌酯条件下，与对照相比，表现出对农药更为敏感的特征。表明MoR1基因参与了包括吡唑醚菌酯和嘧菌酯在内的多真菌农药抗性，可作为真菌农药的新的靶标基因。
实施例4MoR1基因编码蛋白的3D结构解析实验
通过对MoR1基因蛋白的3D解构建模与解析，发现MoR1基因编码的蛋白具有典型的ABCC转运子结构(图7)，包含两侧对称的跨膜结构，每个跨膜结构包含6个α-螺旋跨膜域(图7，跨膜结构区域1和2)，另外下测包含两个关键的功能区域，包括活性的氨基酸以及大量的β-折叠结构区域。此区域为关键的药物或者离子等结合区域，通过消耗ATP的方式将物质泵出细胞膜外或泵入细胞膜内。
序列表
<110> 中国农业科学院植物保护研究所
<120> 稻瘟菌农药靶标基因MoR1及其编码蛋白和应用
<130> 2016
<160> 2
<170> PatentIn version 3.3
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<400> 1
000
<210> 2
<211> 1446
<212> PRT
<213> 稻瘟菌（拉丁名）
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Leu Leu Ser Ala Val Glu His Arg Arg Ser Ala Arg Pro Ser Ser Leu
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Ala Asn Cys Cys Val Lys Phe Val Leu Leu Ile Leu Glu Glu Gln Lys
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Arg Leu Phe Leu Thr Gly Tyr Arg Arg Ala Phe Thr Thr Thr Asp Leu
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Glu Leu Ile Ser Ser Pro Leu Tyr Val Lys Ser Ile Arg Ala Gln Phe
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His Gly Met Thr Asn Gly Gln Thr Ala Asn Gly His Ser Leu Phe Ser
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Ser Ser Thr Asn Thr Phe Ala Leu Gln Ala Phe Thr Ser Leu Gly Ser
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Tyr Ala Leu Ala Pro Val Ile Pro Arg Leu Ala Val Thr Gly Phe Thr
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Gly Arg Asp Arg Pro Ala Ser His Gly Tyr Gly Leu Leu Gly Ala Tyr
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Ala Phe Val Tyr Ile Gly Ile Ala Val Ser Asn Ser Trp Tyr Ser Arg
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His Thr Tyr Lys Ser Val Ser Ile Ile Arg Gly Gly Leu Ile Val Ser
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Thr Met Phe Ser Ser Leu Val Leu Ile Ala Leu Leu Gly Ser Pro Leu
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Ile Lys Asn Glu Pro Glu Thr Met Pro Glu Ser Asn Arg Gly Thr Lys
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Ile Asn Leu Gln Ile Gln Lys Gly Ser Phe Val Ala Leu Val Gly Lys
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Gly Arg Ile Gly Ser Gly Gly Ala Arg Leu Ser Gly Gly Gln Lys Gln
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Arg Leu Ala Leu Ala Arg Ala Ile Ala Thr Arg Lys Asp Ile Val Leu
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Pro Asp His Gly Thr Asn Thr Val Met Thr His Lys Val Glu Asn Gly
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Val Met Gly Cys Ala Val Cys Phe Lys Ile Pro Asp Leu Trp Val Gln
900 905 910
Trp Trp Ser Thr Ala Ile Lys Gln Gly Thr Thr Tyr Ser Ser Ser Tyr
915 920 925
Trp Ile Gly Ile Leu Ala Leu Leu Glu Val Leu Pro Leu Leu Met Leu
930 935 940
Trp Leu Ser Leu Phe His Val Leu Phe Phe Ile Val Pro Arg Ser Ala
945 950 955 960
Ser Thr Met His Asp Ser Leu Leu Arg Thr Val Leu Leu Ala Pro Phe
965 970 975
Gly Phe Ile Ser Arg Val Asp Thr Gly Ser Leu Met Asn Arg Phe Asn
980 985 990
Gln Asp Leu Met Phe Val Asp Thr Arg Leu Pro Ile Asp Leu Phe Asn
995 1000 1005
Thr Ser Ile Asp Phe Phe Ile Thr Ile Ile Gln Leu Ile Leu Val
1010 1015 1020
Val Leu Val Ser Lys Glu Ala Leu Ala Ile Leu Pro Val Val Phe
1025 1030 1035
Gly Ala Leu Tyr Leu Ile Gln Lys Val Tyr Leu Arg Ser Ser Lys
1040 1045 1050
Gln Leu Arg Leu Leu Asp Leu Asp Trp Lys Ala Asp Leu His Thr
1055 1060 1065
Ala Phe Gly Glu Thr Thr Ala Gly Leu Ser Val Ile Arg Ala Asn
1070 1075 1080
Gly Trp Leu Asp Pro Met Arg Ala Lys Phe Ala Glu Lys Leu Asp
1085 1090 1095
Arg Ser Gln Glu Pro Phe Tyr Leu Leu Tyr Met Val Gln Arg Trp
1100 1105 1110
Leu Gln Leu Val Leu Asn Leu Val Val Ala Gly Leu Ala Ile Ala
1115 1120 1125
Ile Ala Gly Val Ala Ile Gly Leu Arg Asp Lys Val Ala Ala Gly
1130 1135 1140
Ala Val Gly Val Ala Leu Leu Asn Thr Thr Thr Leu Gly Glu Thr
1145 1150 1155
Leu Thr Asn Phe Ile Met Ser Trp Thr Ser Leu Glu Thr Ser Leu
1160 1165 1170
Gly Ala Ile Ala Arg Val Cys Thr Phe Glu Gln Asp Thr Pro Arg
1175 1180 1185
Glu Arg Glu Glu Pro Ser Thr Thr Asp Leu Pro Asp Asn Arg Pro
1190 1195 1200
Gly Ala Gly Gln Ile Ser Phe Glu Asn Val Trp Ala Thr Tyr Glu
1205 1210 1215
Asp Glu Gly Cys Gly Ser Asn Trp Gly Leu Ser Gly Ile Thr Leu
1220 1225 1230
Ala Val Gln Pro Gly Glu Arg Val Ala Val Cys Gly Arg Thr Gly
1235 1240 1245
Ser Gly Lys Ser Thr Leu Leu Leu Ala Leu Leu Gly Met Leu His
1250 1255 1260
Thr Pro Ala Gly Ser Ile Arg Ile Asp Gly Val Asp Thr Ser Thr
1265 1270 1275
Leu Pro Ile Asp Val Leu Arg Arg Arg Phe Thr Val Val Ser Gln
1280 1285 1290
Asp Ser Phe Phe Glu Pro Thr Ser Thr Phe Arg Gln Glu Leu Asp
1295 1300 1305
Pro Ser Gly Asp Met Ser Asp Gln Ile Ile Glu Glu Val Leu Arg
1310 1315 1320
Glu Cys Arg Ala Trp Glu Ile Val Asp Gly Ser Gly Gly Leu Gly
1325 1330 1335
Gly Lys Arg Ala Asp Ala Asn Leu Ser Ala Gly Glu Val Gln Leu
1340 1345 1350
Leu Ala Ile Ala Arg Leu Val Leu Gln Trp Gln Ser Gln Pro Ala
1355 1360 1365
Gly Ser Gly Gly Ile Ile Leu Leu Asp Glu Ala Thr Ser Asn Leu
1370 1375 1380
Asp Arg Gln Thr Glu Val Leu Val Glu Ser Ile Met Ala Ala Arg
1385 1390 1395
Leu Gln His Ala Thr Val Val Ser Val Met His Arg Leu Glu Ala
1400 1405 1410
Val Ala Ala Tyr Asp Lys Val Ala Val Leu Asp Lys Gly Val Leu
1415 1420 1425
Val Asp Phe Gly Pro Val Thr Asp Val Met Ala Arg Cys Glu Leu
1415 1420 1425
Phe Thr Gly
1445

Claims (10)

1.稻瘟菌MoR1基因，其核苷酸序列如序列表SEQ ID NO.1所示。
2.权利要求1所述的稻瘟菌MoR1基因作为稻瘟菌农药基因靶标的用途。
3.根据权利要求2所述的稻瘟菌为耐药性稻瘟菌。
4.权利要求1所述的稻瘟菌MoR1基因作为耐药性稻瘟菌基因靶标的用途。
5.一种稻瘟菌农药基因靶标，其核苷酸序列如序列表SEQ ID NO.1所示。
6.一种稻瘟菌多抗真菌农药基因靶标，其核苷酸序列如序列表SEQ ID NO.1所示。
7.稻瘟菌MoR1基因编码的蛋白质，其氨基酸序列如序列表SEQ ID NO.2所示。
8.根据权利要求7所述的蛋白质作为稻瘟菌多抗真菌农药靶点的用途。
9.一种稻瘟菌农药靶点，其核苷酸序列如序列表SEQ ID NO.2所示。
10.一种稻瘟菌多抗真菌农药靶点，其核苷酸序列如序列表SEQ ID NO.2所示。
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