CN110592107B - 一种燕麦叶枯病菌基因CvDUF1及其应用 - Google Patents

一种燕麦叶枯病菌基因CvDUF1及其应用 Download PDF

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CN110592107B
CN110592107B CN201910950510.8A CN201910950510A CN110592107B CN 110592107 B CN110592107 B CN 110592107B CN 201910950510 A CN201910950510 A CN 201910950510A CN 110592107 B CN110592107 B CN 110592107B
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张祥辉
朱耿林
丛杰
纪旭
王璐
刘金亮
潘洪玉
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Abstract

一种燕麦叶枯病菌CvDUF1基因及其应用属微生物基因工程技术领域，本发明提供的来自燕麦叶枯病菌的控制毒素形成的CvDUF1基因，其DNA序列如SEQ ID No:1所示；CvDUF1基因可在植物抗燕麦叶枯病基因工程领域中应用；通过对燕麦叶枯病菌的控制毒素形成的蛋白质CvDUF1进行缺失、突变或修饰，而使其毒素形成受限，可作为靶标在设计和筛选抗燕麦叶枯病药剂中应用。

Description

一种燕麦叶枯病菌基因CvDUF1及其应用
技术领域
本发明属微生物基因工程技术领域，具体涉及植物保护领域中控制真菌毒素形成的基因的发现及应用。
背景技术
燕麦叶枯病是由一种平脐蠕孢菌Cochliobolusvictoriae引起，燕麦叶枯病主要危害叶片、叶鞘、颖等地上部。叶片染病初期会形成边缘不明显的黄色至黄褐色病斑，后逐渐扩大，中间褐色，四周略带黄色。病斑多时，从叶尖端逐渐向下枯死。该病多从下部叶片开始发生并向上扩展。并且，燕麦叶枯病菌可以产生一种毒素-victorin，毒素victorin稀释一百万倍以后仍对特定基因型的燕麦品种有毒害作用，从此也提出了寄主专化性毒素的概念，并得到广泛研究。毒素victorin是燕麦叶枯病菌成功侵染燕麦的必要元素之一。不能产生victorin的菌株则不能对燕麦产生致病性，而离体的victorin也能侵染燕麦。燕麦对victorin的感病性主要由燕麦Vb基因决定，含有Vb基因的燕麦表现为感病，不含有Vb基因的燕麦表现为抗病。而Vb基因与燕麦抗锈病的Pc2基因是同一个基因，因此燕麦叶枯病的感病性是与一个抗性基因相关联的。并且victorin能够引起燕麦的防卫反应，包括胼胝质积累，氧爆发以及细胞程序性死亡。而victorin主要包括5种结构，分别是B、C、D、E和Victoricine，结构C为主要结构，活性占比达到85％-90％。燕麦叶枯病每年给燕麦生产造成了极大的损失，尤其毒素victorin的产生给人畜安全产生很大影响。而毒素victorin的产生机制目前还不清楚，这给燕麦叶枯病的防治造成了很大的困难。
DUF1(Domain of Unknown Function 1)基因是一个在燕麦叶枯病菌中的未知功能基因，通过分析DUF1基因的功能，评价其在燕麦叶枯病菌毒素产生中的作用，有利于鉴定潜在的防治靶标，用于筛选新型防控燕麦叶枯病菌的药剂。
发明内容
本发明的目的旨在提供一种控制真菌毒素形成的基因。
本发明所提供的控制毒素形成的基因来源于燕麦叶枯病菌(Cochliobolusvictoriae)菌株FI3，名称为CvDUF1，其DNA序列如SEQ ID No:1所示。该DNA序列为CvDUF1基因的开放阅读框，由1004个核苷酸组成，其中包含3个内含子序列。
本发明所述燕麦叶枯病菌(Cochliobolus victoriae)CvDUF1基因可在调控燕麦叶枯病菌毒素产生中应用。
对来自燕麦叶枯病菌(Cochliobolusvictoriae)菌株FI3的控制毒素形成的CvDUF1基因进行缺失、突变或修饰，而使其毒素形成受阻，可作为靶标在设计和筛选抗燕麦叶枯病药剂中应用。
本发明证明了CvDUF1基因的缺失，导致燕麦叶枯病菌毒素不能形成，说明CvDUF1基因是燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3毒素形成所必需的基因。目前，关于CvDUF1基因影响毒素形成方面还没有报道，本研究是第一次报道。因此，筛选能够阻止该基因表达与其蛋白质的表达、修饰及定位的化合物，可以有效控制燕麦叶枯病的发生，从而有助于开发新型杀菌剂，即本发明所提供的CvDUF1基因的一个重要用途是：该基因的表达与其编码的蛋白质产物的表达、修饰及定位，可以作为重要候选靶标位点，用于抗燕麦叶枯病药剂的设计和筛选。
附图说明
图1为燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3CvDUF1基因的敲除策略(通过同源重组进行基因替换)示意图
其中：Cv FI3为燕麦叶枯病菌野生型菌株，ΔCvduf1为CvDUF1基因的缺失突变体；引物F1/R1与F2/R2分别用于扩增CvDUF1基因的上下游序列，用作敲除的同源臂；引物F/R，U/NLC37，NLC38/D用于验证突变体。
图2为CvDUF1基因缺失突变体的PCR验证电泳图
其中：F/R、U/NLC37、D/NLC38为所用引物；1为燕麦叶枯病菌(Cochliobolusvictoriae)FI3野生型菌株，2和3为CvDUF1基因缺失突变体；(1)F/R为部分CvDUF1基因扩增结果，(2)U/NLC37为CvDUF1基因上游序列加部分潮霉素序列扩增结果，(3)D/NLC38为CvDUF1基因下游序列加部分潮霉素序列扩增结果。
图3为CvDUF1基因的缺失突变体、野生型菌株及阴性对照菌株(不能产生毒素菌株)的毒素产生测定对比照片
其中：所用试管中液体为各个菌株的培养液，所用叶片为燕麦幼苗叶片，如果菌株能产生毒素，经过24h培养后燕麦叶片会发生弯曲。1为燕麦叶枯病菌(Cochliobolusvictoriae)FI3野生型菌株，2和3为不能产生毒素的燕麦叶枯病菌菌株(阴性对照),4和5为CvDUF1基因的缺失突变体菌株。
图4为CvDUF1基因的缺失突变体与野生型菌株产生毒素的ESI-EIC谱图
其中：将各个菌株接种在液体Fries’培养基培养15天后，进行毒素测定。标准品为纯毒素标准品，WT为燕麦叶枯病菌野生型菌株，ΔCvDUF1为CvDUF1基因的缺失突变体菌株。
具体实施方式
为了更好地描述本发明，下面通过具体的实施例予以进一步说明，下述实施例中的方法，如无特别说明，均为常规方法。
本发明中燕麦叶枯病菌(Cochliobolusvictoriae)菌株FI3提供者为美国康奈尔大学的Babara Gillian Turgeon。
提供者联系方式为：
Dept.of Plant Pathology&Plant-Microbe Biology,334Plant Science Bldg.,Cornell University,Ithaca,NY 14850,United States.E-mail:bgt1@cornell.edu。
实施例1 CvDUF1基因的相关性分析
CvDUF1基因是通过对燕麦叶枯病菌基因组分析获得。燕麦叶枯病菌CvDUF1基因的开放阅读框由1004个核苷酸组成，包含3个内含子。
实施例2CvDUF1基因的敲除
1)CvDUF1基因上下游以及潮霉素基因的扩增
采用引物F1(5'-GGAGCATGCATTTCTCTACGA-3')与R1(5'-TCCTGTGTGAAATTGTTATCCGCTTGGTATCAAAAGAAAAACAACTGAA-3')，以燕麦叶枯病菌野生型菌株FI3的基因组DNA为模板扩增CvDUF1基因上游702bp片段，采用F2(5'-GTCGTGACTGGGAAAACCCTGGCGTGCTACGAGCAGCAGCTAAA-3')与R2(5'-ATGGTGGAGCTGATTTCTGG-3')扩增燕麦叶枯病菌CvDUF1基因下游850bp片段，采用引物M13F(5'-CGCCAGGGTTTTCCCAGTCACGAC-3')和M13R(5'-AGCGGATAACAATTTCACACAGGA-3')，以载体pUCATPH为模板扩增2549bp潮霉素基因。反应体系为：10mmol/L dNTP Mixture，1μL；5×PCR buffer，10μL；上下游引物各2.5μL(10μmol/mL)；模板DNA，2μL；Phusion polymerase，0.5μL(5U)；ddH2O，31.5μL；扩增程序为：98℃预变性2分钟，然后(1)98℃，变性20秒；(2)65℃，退火30秒；(3)72℃，延伸30秒；(4)循环30次；(5)72℃延伸10分钟。将上述3个片段共同转入燕麦叶枯病菌野生型菌株FI3中。
2)燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3转化
a.燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3的产孢培养
从-80℃冰箱取少量燕麦叶枯病菌FI3菌株分生孢子，滴加于CMX培养基【每升CMX培养基包含：0.1g/mL四水合硝酸钙溶液10mL，10mL溶液B，0.5mL微量元素溶液，1g酵母提取物，0.5g酶解干酪素，0.5g酸解干酪素，10g木糖，20g琼脂粉。(每升溶液B包括：20g磷酸二氢钾，25g七水合硫酸镁，15g氯化钠)(每升微量元素溶液包括：57.2mg硼酸，393mg五水合硫酸铜，13.1mg碘化钾，60.4mg一水合硫酸锰，36.8mg四水合钼酸铵，5.49g一水合硫酸锌，948.2mg六水合氯化铁)】上，置24℃培养2周，用CM液体培养基【每升CM培养基包含：0.1g/mL四水合硝酸钙溶液10mL，10mL溶液B，0.5mL微量元素溶液，1g酵母提取物，0.5g酶解干酪素，0.5g酸解干酪素，10g葡萄糖，20g琼脂粉。(每升溶液B包括：20g磷酸二氢钾，25g七水合硫酸镁，15g氯化钠)(每升微量元素溶液包括：57.2mg硼酸，393mg五水合硫酸铜，13.1mg碘化钾，60.4mg一水合硫酸锰，36.8mg四水合钼酸铵，5.49g一水合硫酸锌，948.2mg六水合氯化铁)】刮取、收集孢子，显微镜观察，利用血球计数器调节孢子浓度为1×106/mL。
b.燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3转化
吸取1mL孢子悬浮液于100mLCM液体培养基中，24℃振荡培养(150rpm)28h，离心收集菌丝于80mL酶解液(3.27g氯化钠，0.8g崩溃酶)中酶解2h，收集原生质体。将原生质体用10mLSTC溶液洗涤3次，并最终溶解于500μLSTC溶液(每100mLSTC溶液包括：21.86g山梨醇，1Mtris-HCL1mL，0.735g二水合氯化钙)中。将25mL准备好的PCR片段与100μL原生质体溶液充分混匀，加入1mLPEG溶液(每50mLPEG溶液中包括：聚乙二醇30g，1Mtris-HCL0.5mL，0.37g二水合氯化钙)。最后用1mLSTC溶液稀释，并与再生培养基混合，30℃过夜培养，每个培养皿中加入含有150μg/mL潮霉素的水琼脂10mL，30℃培养3d后挑取扩展的菌落到含有同样抗生素的CMX培养基上。
3)缺失突变体的验证
选用三对引物通过PCR扩增对转化子进行筛选。扩增结果符合如下结果的，确定为CvDUF1基因缺失突变体：上游同源臂之外基因组上的引物U(5'-ACAACGCGTGGATAGAAACA-3')与潮霉素抗性基因的引物NLC37(5'-GGATGCCTCCGCTCGAAGTA-3')配对可以扩增到预期大小(2.1kb)的重组片段；下游同源臂之外基因组上的引物D(5'-CGGTATATCGCCGTTCAACT-3')与潮霉素抗性基因的引物NLC38(5'-CGTTGCAAGACCTGCCTGAA-3')配对可以扩增到预期大小(3.0kb)的重组片段；而编码区引物F(5'-CGGTATGCTCATCGTCCTTT-3')与R(5'-CATTGAGCAGCCTGTCGTAA-3')无扩增条带(野生型菌株可扩增到644kb片段)(见图2)。结果，从转化子中筛选到2株CvDUF1基因缺失突变体，用于后续功能分析。
实施例3CvDUF1基因在燕麦叶枯病菌(Cochliobolus victoriae)菌株FI3毒素产生中
的作用
将燕麦叶枯病菌野生型菌株FI3和CvDUF1基因缺失突变体分别接种于固体CMX培养基上，生长两周后，用5mLFries’培养基【酒石酸铵5g，硝酸铵1g，磷酸二氢钾1g，七水合硫酸镁0.5g，氯化钠0.1g，两水合氯化钙0.13g，蔗糖30g，酵母浸粉1g，铁溶液1mL，用水定容至1L。(铁溶液：七水合硫酸亚铁2g，EDTA2.41g，用水定容至100mL)】冲洗菌丝及分生孢子，收集菌丝及孢子悬浮液至20mLFries’培养基中，黑暗培养15天后用于毒素测定。
毒素生物测定：将上述培养液用纱布过滤，去除菌丝体，取2mL培养液加入试管中，取生长两周左右的燕麦幼苗叶片放入试管中培养24h。过夜培养后如果燕麦叶片保持直立，则证明菌株不能产生毒素，如燕麦叶片变弯曲，则证明菌株能产生毒素。
毒素的仪器测定：将上述培养液用纱布过滤，去除菌丝体，过反向C18柱进行纯化，除去杂质。然后用乙腈/水(50/50)对C18柱进行淋洗，将淋洗液过凝胶柱分离，后用10mMNaCL溶液淋洗，将淋洗液进行浓缩干燥，并用乙腈进行溶解，上高效液相色谱进行测定。高效液相色谱检测条件为：流动相为25％乙腈水溶液，检测器为紫外检测器，检测波长254nm，色谱柱为C18柱。
Figure BDA0002225512420000051
序 列 表
发明名称：一种燕麦叶枯病菌基因CvDUF1及其应用”
SEQ ID No:1的序列
　　　（i）序列特征：（A）长度：1004 bp；（B）类型：核苷酸；（C）链性：单链
　　　（ii）分子类型：DNA
　　　（iii）序列描述：SEQ ID No:1
1-60 atgagctcag cagcatatag ccgagtagat tcaagttcaa ctgacagaaatagtgatgaa
61-120 tatcacaaag aacggtacct tgcacttaga acgaataatg ctaagggtttcaaaggagtt 121-180 caaaggcgat tatggatagc gattaattcc agtttatggc ttgttacactcggtatgctc 181-240 atcgtccttt tacggcggga gagtttgggc acgagaaacg gcgggtttgagcttgggttc 241-300 tcagctactg agcttggtat gacaaacaac attgatctac cgtattcatccaatcgctaa 301-360 ctagaaaatc agccgctacg tttcgaacca ttagaatctc tactgagatattcaatcagc 361-420 ctgaaggaac atcgttctct acattaccta cgtcgtctta tgtaggtgaaccaacagagg 421-480 agattgacaa tgcttggagg atattagttt cccgtaagca agactttccagaatcaatta 481-540 ctcaggccta tgctcattgc tcttctcact agcaactgtc ctttacctaaaagaggacga 541-600 gattggcgag tataaaaaca gcactaccaa gcaagacaac ctctgggtcgccgggtcagt 601-660 tttataccaa acttggctga acctcatgcc ttatttctag taaaaatagacgttgctaac 661-720 attttcattt ggttagtttg caagtatacc actaccttca ctgcattaacgcactacgcc 721-780 gcgcagcata ccaacatgtt tacggggctc cgacaaagga acatctaaatcacctcgacc 781-840 attgcattga catgttacga caggctgctc aatgtcaatc tgacctgacacctatgttat 841-900 atttcaaccc cgagaatgac ccaaacacta tgctcattaa gtcacaccaacactcttgtc 901-960 gccggtttga tttagtaaat gaatgggcga tggctcggtc tcagtgcaaagggaatacta 961-1004cttgtgccat cgaagtaggg aagcaagtag gcggcgagat gtaa

Claims (1)

1.一种燕麦叶枯病菌（Cochliobolus victoriae）CvDUF1基因作为靶标在设计和筛选抗燕麦叶枯病药剂中应用，其中所述基因的DNA序列如SEQ ID NO.1所示。
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