CN112626039B - 一种氧化还原酶GliT及其在抵抗真菌毒素中应用 - Google Patents
一种氧化还原酶GliT及其在抵抗真菌毒素中应用 Download PDFInfo
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- CN112626039B CN112626039B CN202011528161.XA CN202011528161A CN112626039B CN 112626039 B CN112626039 B CN 112626039B CN 202011528161 A CN202011528161 A CN 202011528161A CN 112626039 B CN112626039 B CN 112626039B
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- glit
- oxidoreductase
- mycotoxin
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Abstract
本发明公开了一种氧化还原酶GliT及其在抵抗真菌毒素中的应用。氧化还原酶GliT,其氨基酸序列如SEQ ID NO.2所示。鉴于目前尚未有关于深海真菌G.pallida FS140抗多种真菌毒素的研究。因此本发明从深海真菌FS140的cDNA文库中获得了氧化还原酶GliT基因序列,并成功导入到酿酒酵母S.cerevisiae BJ5464中进行了抗毒功能验证,从而为后期提高酿酒酵母抗胶霉毒素及其衍生物、单端孢霉烯类毒素、Eupenifeldin和Dicerandrol C毒素的能力,提升相关真菌毒素的异源表达水平,解析相关真菌毒素作用机制奠定分子生物学基础,此外,有利于下一步分离纯化新型氧化还原酶GliT,明确其对相关真菌毒素是否具有降解效果,为开发成经济、可行的降毒试剂、防控真菌毒素添加剂等提供新思路。
Description
技术领域
本发明属于基因工程领域,具体涉及一种深海真菌G.pallida的氧化还原酶GliT及其在抵抗真菌毒素中的应用。
背景技术
真菌毒素是产毒真菌在生长过程中产生的一类次生代谢物质,主要包括黄曲霉毒素、脱氧雪腐镰刀菌烯醇(呕吐毒素)、玉米赤霉烯酮、伏马毒素和赭曲霉毒素等。如何安全合理地利用被毒素污染的粮食作物,维护国家粮食食品质量安全,是一个亟待解决的问题。特别是单端孢霉烯族毒素,被认为是最危险的自然发生粮食污染物,已成为国际研究热点。单端孢霉烯族毒素是一大类由镰刀菌产生的化学结构相似的毒性物质的统称,其基本结构为四环的倍半萜,根据取代基的不同,可以分为A、B、C、D四种类型。迄今为止已经发现了大概170种单端孢霉稀族毒素,而A型和B型单端孢霉稀族毒素污染率最高,包括DON、3-ADON、15-ADON、NIV、T-2和DAS毒素等,其中污染率和含量最高的是DON,因此,阻止和减少DON毒素进入人和动物的食物链以防止其危害为主要研究内容。
胶霉毒素(gliotoxin,GT)是一类二酮哌嗪类化合物(epipolythiodioxopiperazine,ETP),ETP作为重要的毒力因子,其通过不同途经对各种细胞产生特异性毒性,在侵袭性曲霉病中发挥重要协同作用。ETP发挥毒副作用主要通过二硫键,巯基与靶蛋白的交联进而灭活该蛋白活性,并能通过氧化还原循环产生具有毒害的活性氧(reactiveoxygenspecies,ROS),ROS生成机制被认为是GT产生细胞毒性的一种机制。胶霉毒素对宿主的毒性主要包括:诱导细胞凋亡;导致氧化还原反应失衡;抑制蛋白酶体活性,导致NF-κB活性被抑制,降低免疫活性等。它不仅对动植物细胞,而且对宿主菌也有毒性(Kamei K,Watanabe A.Aspergillus mycotoxins and their effect on thehost.Medical Mycology,2005,43:S95–S99.)。
Dicerandrol C和Eupenifeldin毒素是本课题组从深海真菌Phomopsis sp.FS479和Phomopsis tersa FS441中分离得到的真菌次级代谢产物,具有很好的抗肿瘤活性,但是对于人体和动植物的危害尚不清楚。
发明内容
本发明的第一个目的是提供一种氧化还原酶GliT及其在抵抗真菌毒素中的应用,所述的氧化还原酶基因GliT,其氨基酸序列如SEQ ID NO.2所示。
本发明的氧化还原酶基因GliT通过以下方法获得的:通过转录组测序结果预测编码氧化还原酶GliT的序列,在其上下游设计特异性引物,其引物序列为GliT-F:5'-ATGTCCATCGGAAAACTTCTCGC-3';GliT-R:5'-CTATGGCTCCCAATCAATCCCAAAT-3',以由深海真菌FS140转录组反转录而得的cDNA文库为模板,通过PCR扩增获得产物并纯化回收片段,获得氧化还原酶基因GliT,其核苷酸序列如SEQ ID NO.1所示,其编码的氨基酸序列如SEQ IDNO.2所示。
本发明利用同源重组法将GliT基因插入到酵母载体YEp352-TEF1-CYC1的表达盒内部。首先设计含有同源臂的GliT基因的上下游引物,其引物序列为YEp352-GliT-F:5'-G CAATCTAATCTAAGTCTAGAATGTCCATCGGAAAACTTCTCGC-3';YEp352-GliT-R:5'-TACATGATGCGG CCCGTCGACCTATGGCTCCCAATCAATCCCAAAT-3'(下划线序列为同源臂片段),通过PCR扩增获得产物并纯化回收片段。对已构建的YEp352-TEF1-CYC1载体采用内切酶Sal I和Xba I双酶切,然后使用ClonExpress II One Step Cloning Kit C112(Vazyme)将片段和酶切载体重组连接并转化至大肠杆菌感受态细胞,涂布于氨苄青霉素抗性平板筛选出阳性克隆。经过此轮分子克隆,目的基因GliT(其核苷酸序列如SEQ ID NO.1所示)插入到启动子TEF1和终止子CYC1之间,构建得到YEp352-TEF1-GliT载体,将其电转入酿酒酵母BJ5464-D细胞中,利用尿嘧啶缺陷型的SD培养基平板进行筛选和验证。与转入YEp352-TEF1-CYC1质粒(阴性对照)的酿酒酵母BJ5464-D相比,含有重组载体YEp352-TEF1-GliT的酿酒酵母生长速度明显加快,相同培养时间内菌落密度更高,证明功能基因GliT能有效协助酿酒酵母抵抗外源胶霉毒素及其衍生物、单端孢霉烯类毒素、Eupenifeldin和Dicerandrol C毒素,为在酿酒酵母内重构胶霉毒素及其衍生物、单端孢霉烯类毒素、Eupenifeldin和Dicerandrol C毒素毒素生物合成通路奠定基础。
本发明的第二个目的是提供编码上述氧化还原酶GliT的氧化还原酶基因GliT。优选,其核苷酸序列如SEQ ID NO.1所示。
本发明的第三个目的是提供一种表达载体,含有上述的氧化还原酶基因GliT。
本发明的第四个目的是提供一种宿主细胞,含有上述的表达载体。
所述的宿主细胞优选为酿酒酵母S.cerevisiae BJ5464。
本发明的第五个目的是提供上述的氧化还原酶基因GliT在协助宿主细胞抵抗真菌毒素中的应用。
优选地,所述真菌毒素为胶霉毒素及其衍生物、单端孢霉烯类毒素、Eupenifeldin和Dicerandrol C毒素。
更优选地,所述的单端孢霉烯类毒素为呕吐毒素、伏马毒素B1、玉米赤霉烯酮和杆孢菌素E中的至少一种。
所述的宿主细胞优选为深海真菌G.pallidaFS140或酿酒酵母S.cerevisiaeBJ5464。
本发明的第六个目的是提供一种表达盒,该表达盒含有上述的氧化还原酶基因GliT。
与现有技术相比,本发明具有以下有益效果:
本发明所涉及的深海真菌G.pallida FS140分离自南海沉积物,本课题组前期对该菌株进行了转录组测序并对胶霉毒素生物合成相关基因进行了注释。鉴于目前尚未有关于深海真菌G.pallida FS140抗多种真菌毒素的研究。因此本发明从深海真菌FS140的cDNA文库中获得了氧化还原酶GliT基因序列,并成功导入到酿酒酵母S.cerevisiae BJ5464中进行了抗毒功能验证,从而为后期提高酿酒酵母抗胶霉毒素及其衍生物、单端孢霉烯类毒素、Eupenifeldin和Dicerandrol C毒素的能力,提升以上真菌毒素的异源表达水平,解析以上真菌毒素作用机制奠定分子生物学基础,此外,有利于下一步分离纯化新型氧化还原酶GliT,明确其对以上真菌毒素是否具有降解效果,为开发成经济、可行的降毒试剂、防控真菌毒素添加剂等提供新思路。
本发明的深海真菌G.pallida FS140,其公开于文献:Zhang-Hua Sun,JiangyongGu,Wei Ye,Liang-Xi Wen,Qi-Bin Lin,Sai-Ni Li,Yu-Chan Chen,Hao-Hua Li,Wei-MinZhang.Geospallins A–C:New Thiodiketopiperazines with Inhibitory Activityagainst Angiotensin-Converting Enzyme from a Deep-Sea-Derived FungusGeosmithia pallida FS140.Marine Drugs,2018,16(12),464.https://doi.org/10.3390/md16120464。该菌种本申请人也持有,保证自发明的申请日起20年内向公众提供。
附图说明
图1是氧化还原酶GliT基因序列的获得:以FS140 cDNA文库为模板,基因GliT扩增产物的电泳图;
图2为实验所使用真菌毒素结构式及两种酿酒酵母在YPD平板和各种真菌毒素平板中培养30h的效果图。A、酿酒酵母BJ5464-D(YEp352-TEF1-CYC1);B,酿酒酵母BJ5464-D(YEp352-TEF1-GliT),10-2、10-3、10-4分别代表OD600约为0.01、0.001、0.0001的5μL菌液样品,从第二排开始,左侧是真菌毒素的结构式,右侧是含有该真菌毒素的YPD平板,菌在上面的生长图。
图3为重组载体YEp352-TEF1-GliT的构建;其中A为YEp352-TEF1-CYC1载体图谱;B为YEp352-TEF1-GliT载体图谱;C为基因GliT的菌落PCR扩增产物的电泳图;
具体实施方式
以下实施例是对本发明的进一步说明,而不是对本发明的限制。
本实施例中所用的SD固体培养基的配方为:每升含有葡萄糖20g、Do supplement0.62g(-Leu/-Trp/-Ura,Clontech)、无氨基酵母氮源YNB 6.7g(普博欣)、亮氨酸0.06g、色氨酸0.04g和琼脂粉20g,余量为蒸馏水,其配制方法是将各成分混合均匀,灭菌制得。
本实施例中所用的YPD固体培养基的配方为:每升含有酵母粉10g、蛋白胨20g、葡萄糖20g和琼脂粉20g,余量为蒸馏水,其配制方法是将各成分混合均匀,灭菌制得。
本实施例中所用真菌毒素:伏马毒素B1(FUM)和玉米赤霉烯酮(ZEN)购买于Sigma-aldrich;呕吐毒素(DON)购买于Sigma;胶霉毒素(Gliotoxin)购买于上海源叶生物科技有限公司;以下毒素为本课题组分离得到:胶霉毒素衍生物(FS140-12-2)分离自深海真菌Geosmithia pallida FS140;Dicerandrol C分离自Phomopsis sp.FS479;Eupenifeldin从Phomopsis tersa FS441菌株中分离得到。
实施例1新型氧化还原酶GliT基因序列的获得
基因GliT的扩增:将深海真菌Geosmithia pallida FS140接种于YPD培养基平板,于37℃培养72h,挑取新鲜的菌丝体,利用真菌RNA提取试剂盒提取RNA,再用All-in-one RTMaster Kit逆转录获得cDNA。根据转录组测序结果预测编码氧化还原酶基因GliT序列,设计上下游引物GliT-F和GliT-R,其引物序列为GliT-F:5'-ATGTCCATCGGAAAACTTCTCGC-3';GliT-R:5'-CTATGGCTCCCAATCAATCCCAAAT-3',以cDNA文库为模板扩增,获得PCR产物(图1)。回收产物并用pEASY-T1试剂盒进行TA克隆,转化至大肠杆菌感受态细胞,涂布于氨苄青霉素抗性平板筛选出阳性克隆,以通用引物M13-F(5′-GTAAAACGACGGCCAGT-3′)和M13-R(5′-CAGGAAACAGCTATGAC-3′)进行菌液PCR验证阳性克隆并测序,获得目的基因GliT序列(其核苷酸序列如SEQ ID NO.1所示,atgtccatcggaaaacttctcgccaacggagccctgttggttgatgtcctcatcatcggtgcaggcccctcgggtctgtctaccgcaaccggactggcccgtcagcttcataccgcggtcgtctttgactccggagtgtatcgcaacgcaaagacacagcacatgcacaatgtcctaggctgggaccaccggaatccgtccgagctacgggccgccggtcgagctgatctcgctgcgcggtactcgacgatccagttccagaatgccaccgtcgagacgatcaagaggatcggggagaagcaactcttcgaggcgcgtgacacggacggtaagcgctggtatggtcggaaggtcgtgttggccacgggagtccgagacattcctctggatattgagggttactcggaatgctgggccaatggaatctaccactgcctgttctgtgacggctatgaagaacgaggccaggagaccgtcggtgtcctcgccatgggccccatcgccaatcctccacgagccctacacttggcccgaatggcccatcgactctctgaatctgtcaccgtctacacccacggcgatgagcaactggccaaggagattcagcaggcggccgggggtgattcctcgtggctgaagctggagacccggcccatcgtgcgattcgagaagggggatgttgccaaaaccgttatcgtccatttctccgagacgacagacacgaagcaagaaggcttcctggcctataaccccaagacggagatcaacggcccctttgccaaccagctctcattgcagttgtccgaagtcggggacatccagacctcggctccgttctatgagaccagtgtgcccggggtattcgccgttggagactgtgccaccccgttgaaggccgtcagtccggcgattgcaatgggatcgttggctgctggaggtctggttgctcagctgcaggcccagccagtgatggaatttgggattgattgggagccatag)。
实施例2新型氧化还原酶GliT的功能验证
利用同源重组法将新型氧化还原酶基因GliT插入到酵母载体YEp352-TEF1-CYC1中(YEp352-TEF1-CYC1为早期构建质粒,携带有组成型启动子TEF1和终止子CYC1,载体图谱见图3A,为现有技术中的已知产品:Xiaodan Ouyang,Yaping Cha,Wen Li,Chaoyi Zhu,Muzi Zhu,Shuang Li,Min Zhuo,Shaobin Huang and Jianjun Li.Stepwise engineeringof Saccharomyces cerevisiae to produce(+)-valencene and its relatedsesquiterpenes,RSC Adv.,2019,9,30171,DOI:10.1039/c9ra05558d)。首先设计针对基因GliT(SEQ ID NO.1)扩增的上下游引物YEp352-GliT-F和YEp352-GliT-R,其引物序列为YEp352-GliT-F:5'-GCAATCTAATCTAAGTCTAGAATGTCCATCGGAAAACTTCTCGC-3';YEp352-GliT-R:5'-TACATGATGCGGCCCGTCGACCTATGGCTCCCAATCAATCCCAAAT-3'(下划线序列为同源臂片段),以深海真菌Geosmithia pallida FS140的cDNA为模板,通过PCR扩增获得产物。对载体YEp352-TEF1-CYC1采用Sal I和Xba I双酶切并回收产物,然后使用ClonExpress II OneStep Cloning Kit C112(Vazyme)将两个产物重组连接并转化至DH5α中筛选阳性克隆。采用引物YEp352-GliT-F和YEp352-GliT-R进行菌落PCR验证,结果表明基因GliT成功插入YEp352-TEF1-CYC1载体中(图3C),并通过测序予以确认,得到YEp352-TEF1-GliT载体(载体图谱见图3B)。
制备毒素敏感型酿酒酵母Saccharomyces cerevisiae BJ5464-D(relevantgenotype:Δpdr5Δpdr10Δpdr15)的感受态细胞(为现有技术中的已知产品,该菌株对毒性化合物更加敏感:Wolfgang Schweiger,Jayanand Boddu,Sanghyun Shin,BrigittePoppenberger,Franz Berthiller,Marc Lemmens,Gary J.Muehlbauer,and GerhardAdam.Validation of a Candidate Deoxynivalenol-Inactivating UDP-Glucosyltransferase from Barley by Heterologous Expression in Yeast,MPMI,2010,Vol.23,No.7,DOI:10.1094/MPMI-23-7-0977)。将YEp352-TEF1-GliT质粒载体以及YEp352-TEF1-CYC1质粒载体(阴性对照)分别电转入酿酒酵母BJ5464-D细胞中(1500V,5ms),均匀涂布于尿嘧啶缺陷型的SD平板中,在30℃培养2d,利用菌落PCR筛选阳性克隆,获得分别含有YEp352-TEF1-GliT质粒以及YEp352-TEF1-CYC1质粒的酿酒酵母BJ5464-D细胞。
分别将酿酒酵母BJ5464-D(YEp352-TEF1-CYC1)、酿酒酵母BJ5464-D(YEp352-TEF1-GliT)接种于相应缺陷型的SD培养基中,在30℃培养2d。用分光光度计测量各菌液OD600,将各菌液用无菌水稀释到OD600≈1.0作为原液,再以100μL的原液加900μL的无菌水的方式稀释成10-1,以同样的方式稀释成10-2、10-3、10-4。各取5μL不同菌株的10-2、10-3、10-4的稀释液分别在YPD平板和YPD-含各种真菌毒素平板上点板,在30℃培养并实时观察。培养30h的平板结果显示(图2),酿酒酵母BJ5464-D(YEp352-TEF1-CYC1)、酿酒酵母BJ5464-D(YEp352-TEF1-GliT)在不添加任何毒素的YPD平板上生长状况近乎一致,但在含有真菌毒素(包括2.5μM FS140-12-2、100μM Gliotoxin、250μM DON、250μM ZEN、250μM FUM、200μMMR4、500μM Dicerandrol C和400μM Eupenifeldin)的YPD平板上,阴性对照BJ5464-D(YEp352-TEF1-CYC1)明显生长受阻,几乎都不能长。而导入新型氧化还原酶GliT的酿酒酵母则生长良好,其不同稀释度下的菌体密度与正常酿酒酵母相当,说明新型氧化还原酶GliT部分或全部恢复了酿酒酵母BJ5464-D对外源添加毒素的耐受性,有效帮助酿酒酵母在含有毒素的环境下正常生长。
以上仅是本发明的优选实施方式,应当指出的是,上述优选实施方式不应视为对本发明的限制,本发明的保护范围应当以权利要求所限定的范围为准。对于本技术领域的普通技术人员来说,在不脱离本发明的精神和范围内,还可以做出若干改进和润饰,这些改进和润饰也应视为本发明的保护范围。
序列表
<110> 广东省微生物研究所(广东省微生物分析检测中心)
<120> 一种氧化还原酶GliT及其在抵抗真菌毒素中应用
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 999
<212> DNA
<213> 深海真菌FS140(Geosmithia pallida)
<400> 1
atgtccatcg gaaaacttct cgccaacgga gccctgttgg ttgatgtcct catcatcggt 60
gcaggcccct cgggtctgtc taccgcaacc ggactggccc gtcagcttca taccgcggtc 120
gtctttgact ccggagtgta tcgcaacgca aagacacagc acatgcacaa tgtcctaggc 180
tgggaccacc ggaatccgtc cgagctacgg gccgccggtc gagctgatct cgctgcgcgg 240
tactcgacga tccagttcca gaatgccacc gtcgagacga tcaagaggat cggggagaag 300
caactcttcg aggcgcgtga cacggacggt aagcgctggt atggtcggaa ggtcgtgttg 360
gccacgggag tccgagacat tcctctggat attgagggtt actcggaatg ctgggccaat 420
ggaatctacc actgcctgtt ctgtgacggc tatgaagaac gaggccagga gaccgtcggt 480
gtcctcgcca tgggccccat cgccaatcct ccacgagccc tacacttggc ccgaatggcc 540
catcgactct ctgaatctgt caccgtctac acccacggcg atgagcaact ggccaaggag 600
attcagcagg cggccggggg tgattcctcg tggctgaagc tggagacccg gcccatcgtg 660
cgattcgaga agggggatgt tgccaaaacc gttatcgtcc atttctccga gacgacagac 720
acgaagcaag aaggcttcct ggcctataac cccaagacgg agatcaacgg cccctttgcc 780
aaccagctct cattgcagtt gtccgaagtc ggggacatcc agacctcggc tccgttctat 840
gagaccagtg tgcccggggt attcgccgtt ggagactgtg ccaccccgtt gaaggccgtc 900
agtccggcga ttgcaatggg atcgttggct gctggaggtc tggttgctca gctgcaggcc 960
cagccagtga tggaatttgg gattgattgg gagccatag 999
<210> 2
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<213> 深海真菌FS140(Geosmithia pallida)
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Gly Asp Val Ala Lys Thr Val Ile Val His Phe Ser Glu Thr Thr Asp
225 230 235 240
Thr Lys Gln Glu Gly Phe Leu Ala Tyr Asn Pro Lys Thr Glu Ile Asn
245 250 255
Gly Pro Phe Ala Asn Gln Leu Ser Leu Gln Leu Ser Glu Val Gly Asp
260 265 270
Ile Gln Thr Ser Ala Pro Phe Tyr Glu Thr Ser Val Pro Gly Val Phe
275 280 285
Ala Val Gly Asp Cys Ala Thr Pro Leu Lys Ala Val Ser Pro Ala Ile
290 295 300
Ala Met Gly Ser Leu Ala Ala Gly Gly Leu Val Ala Gln Leu Gln Ala
305 310 315 320
Gln Pro Val Met Glu Phe Gly Ile Asp Trp Glu Pro
325 330
Claims (1)
1.氧化还原酶GliT在协助宿主细胞抵抗真菌毒素中的应用,所述的氧化还原酶GliT的氨基酸序列如SEQ ID NO.2所示;所述的真菌毒素为呕吐毒素、伏马毒素B1、玉米赤霉烯酮、Eupenifeldin或Dicerandrol C;所述的宿主细胞为深海真菌Geosmithia pallida FS140或酿酒酵母Saccharomyces cerevisiae BJ5464。
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