CN108165559A - 一种c2h2型转录因子基因及其应用 - Google Patents
一种c2h2型转录因子基因及其应用 Download PDFInfo
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Abstract
本发明公开了一种C2H2型转录因子基因asr1，其是在土生隐球酵母BSLL1‑1菌株中克隆的，其核苷酸序列如SEQ ID NO：1所示，编码如SEQ ID NO：2所示氨基酸序列的蛋白质；本发明将带有潮霉素基因阻断的asr1重组片段导入土生隐球酵母中，土生隐球酵母asr1基因突变，突变菌株对多种离子胁迫的抗性增加，该突变菌株具有降低土壤中多种离子的应用潜力。
Description
技术领域
本发明属于基因工程领域，具体地，涉及土生隐球酵母编码C2H2型转录因子基因的核苷酸序列和氨基酸序列，涉及含有该基因的同源重组质粒和含有该同源重组片段的突变菌株，它们的制备方法，以及它们在抵抗环境中离子胁迫中的应用。
背景技术
逆境下植物细胞内常常靠积累无机离子来降低细胞内的渗透势，而根据植物种类的不同，植物体内积累和吸收的离子种类也不同，主要有K+、Na+、Ca2+、Mg2+等。而离子过多则会产生胁迫，离子胁迫会使植物产生营养亏损、呼吸受阻、渗透胁迫、生长减慢等问题。离子胁迫对植物整体发育的影响表现为缩短植物的生长周期，缩短植物的开花期，抑制植物器官和组织的生长，加速其发育进程。用NaCl处理过的植物幼苗，植物的干重、湿重，植株株高都会降低。盐胁迫对植物生理生化方面的影响主要体现在细胞脱水、膜结构和功能发生紊乱，细胞膜通透性发生改变。植物细胞质膜能维持其完整性和选择透过性，取决于细胞内外的一价离子(Na+，K+)和二价离子(Ca2+)之间的平衡，如果一方过量，会打破这个平衡，从而使细胞膜通透性发生改变。过量的Na+、Cl-和Ca2+渗入植物细胞内会导致植物细胞叶绿体受损，细胞光合强度降低。它在影响植物光合作用的同时也会影响其呼吸作用，在盐胁迫下，植物的呼吸作用先会增强，而后随着植物的生长而减弱。离子过多还会降低植物的物质代谢能力，造成一系列不良反应。
土壤微生物是土壤生态体系的重要组成部分，在植物和土壤的相互作用过程中扮演着重要的角色。土壤微生物参与土壤养分转化、物质代谢、有机物分解、矿化以及污染物降解等多种生化反应。特别是，土壤微生物在土壤中污染物或者是重金属清除中发挥了重要作用，也即微生物修复技术。微生物修复技术是利用微生物（土著菌、外来菌、基因工程菌）对污染物的代谢作用而转化、降解污染物。微生物修复技术已成功应用于煤气厂址PAHs污染修复，石油烃污染土壤修复，农药污染土壤修复等。此外，重金属污染土壤的微生物修复主要是利用土壤中天然的微生物资源，削减、净化土壤中重金属或降低重金属毒性，从而使污染物的浓度降低到可以接受的水平，或将有毒有害的污染物转化为无害的物质，也包括将其稳定化以减少其向周边环境扩散。在重金属污染土壤的生态修复中微生物主要通过以下几种方式起作用：（1）通过微生物的吸附、代谢达到对重金属消减、净化作用和固定作用；（2）通过微生物改变重金属的化学形态，使重金属固定或生物可利用性降低，减少重金属的危害；（3）土壤微生物通过氧化还原作用改变根际重金属形态或产生的有机酸可增加金属的溶解性，提高重金属的有效性，以利于植物吸收；（4）通过促进植物生长，提高植物抗病性、抗逆能力等方式间接影响修复效率。微生物抵抗金属离子毒性的一个重要手段是通过细胞壁与细胞膜的膜表面富集作用。当环境中含金属离子浓度较高时，微生物会先摄取一定量的金属离子，刺激其体内的抗性机制表达，促进胞内过多的金属离子外排，或者使外界金属离子进入不了细胞内，以此来降低环境中金属离子对菌体产生的毒性。
锌指基因广泛存在于生物体内，参与细胞分化、胚胎发育等相关基因的表达。根据半胱氨酸及组氨酸的数目不同，锌指蛋白分为很多亚型，其中最为广泛的是C2H2型锌指结构，它作为生物体内重要的转录因子可调控众多生理反应。对粗糙脉孢菌C2H2家族锌指转录因子基因敲除的57株突变株在以2％结晶纤维素为唯一碳源的培养条件下进行产纤维素酶水平分析,发现突变株在蛋白水平及内切β-1,4-葡聚糖酶酶活水平均比野生型有25％-77％不等的显著提高。运用Blastp比对并结合Pfam和SMART对烟草基因组数据库分析, 鉴定了118条C2H2锌指蛋白家族成员，所有C2H2分为5个亚家族,同一亚家族成员之间在结构域和理化性质上呈现较高一致性，每个成员都含有C2H2结构域,在数量上存在较大差异。组织表达分析表明,每个C2H2亚家族都有成员在不同组织中表达,在叶及根中有些基因的表达量较高。C2H2型锌指蛋白参与基因表达调控的方式是识别并结合特异DNA片段，但具体是怎样识别和结合靶基因的还不完全清楚。对锌指蛋白-DNA复合物进行晶体结构分析，及应用定点突变技术来研究结构中具体是哪些氨基酸在识别并结合DNA中起重要作用，获得锌指蛋白中氨基酸与DNA中碱基相对应的识别码，从而为研究其调控基因表达方式奠定了基础。
发明内容
本发明旨在提供一种C2H2型转录因子asr1基因，其核苷酸序列如SEQ ID NO：1所示，编码如SEQ ID NO：2所示氨基酸序列的蛋白质；其由土生隐球酵母（C. humicolus）BSLL1-1菌株产生，基因序列全长1872bp，编码C2H2型转录因子蛋白由623个氨基酸组成，分子量约为66 kDa。
本发明另一目的是将上述C2H2型转录因子基因asr1进行阻断获得的同源重组基因片段及包含该片段的重组质粒。
本发明另一目的在于提供一种含有上述同源重组基因片段的土生隐球酵母突变菌株。
本发明另一目的是将上述同源重组基因片段应用在增强微生物抗离子胁迫能力中。
为了实现本发明的上述目的，本发明提供了如下的技术方案：
1、本发明提取从云南省保山市龙陵县周边茶园茶树根际酸性土壤中分离的土生隐球酵母BSLL1-1菌株基因组DNA，送上海人类基因组研究中心进行基因组测序，得到asr1基因序列。根据该序列设计引物，以土生隐球酵母cDNA为模板扩增asr1基因片段。将目的片段连接到pMD-18T载体上，得到含有目的片段的重组载体pMD18-T-asr1并转化到大肠杆菌感受态细胞DH5α中，涂布含氨苄青霉素的平板上，挑取阳性菌落测序，鉴定出编码Asr1蛋白的基因，其序列如SEQ ID NO：1所示，该蛋白由623个氨基酸编码，氨基酸序列如SEQ ID NO：2所示；
2、asr1基因同源重组片段构建方法
将asr1基因的前800 bp、后872 bp核苷酸和潮霉素基因三个独立片段分别设计扩增引物，前基因的反向引物与后基因的正向引物都带有一段同源臂；分别以asr1前臂和潮霉素基因为模板，扩增得到asr1前臂：潮霉素重组片段。以潮霉素基因与asr1后臂为模板，扩增得到潮霉素：asr1后臂重组片段。然后以上述两个重组片段为模板进行重叠延伸PCR，得到asr1前臂：潮霉素基因：asr1后臂片段，命名为asr1F：hyg：asr1R。
3、本发明将得到的asr1前臂：潮霉素基因：asr1后臂重组片段asr1F：hyg：asr1R连接到pMD18-T载体上得到质粒pMD18-asr1:hyg并对其进行测序；在土生隐球酵母感受态细胞中加入测序正确的重组片段asr1F：hyg：asr1R，然后进行电击转化，将混合液涂于含200μg·mL-1 潮霉素的GM平板上筛选发生同源重组的克隆；从潮霉素平板上挑取单菌落，用PCR和real-time PCR方法检测敲除是否成功。
4、本发明比较了土生隐球酵母野生型菌株与突变型菌株在多种离子胁迫处理下的生长。配制YPD固体培养基，使各离子终浓度如下：Ca2+ 50 mmol/L，Mn2+ 7.5 mmol/L，Mg2 + 300 mmol/L，Na+ 0.8 mol/L，K+ 0.8 mol/L。将野生型酵母和突变酵母接种到YPD液体培养基中摇床培养过夜。按1%接种量转接至新鲜的YPD 液体培养基中，将菌液浓度调到OD600为1；将菌液分别稀释101、102、103、104倍，每个浓度梯度菌液分别取2 µL接种到含有各离子的YPD固体平板上，在28 ℃培养箱中倒置培养，观察菌落大小；以不加离子的YPD固体平板作为对照，每种设置三个平板作为重复；结果发现在含有K+、Na+、Ca2+、Mn2+、Mg2+的平板上，突变型比野生型菌落大，这说明asr1基因突变后增强了土生隐球酵母菌株抗离子胁迫的能力。
本发明的优点和技术效果如下：
本发明为抵抗土壤中离子胁迫提供了一种方法，asr1基因突变菌株可以增加对多种离子胁迫的抗性，这种利用微生物抵抗环境中离子胁迫的技术费用低，操作简便，对环境影响小，不会造成二次污染。
附图说明
图1为本发明荧光定量分析转录因子asr1基因在铝胁迫下的表达示意图，A图为浓度梯度下基因的定量结果；B图为基因在时间梯度下的定量结果。
图2为本发明转录因子asr1基因扩增示意图；
图3为本发明asr1F:hyg:asr1R同源重组片段的扩增示意图，其中a图为asr1基因后臂（C）、潮霉素基因（B）及asr1基因前臂（A）PCR扩增图；b图为潮霉素基因和asr1基因重组片段（ABC），asr1前臂和潮霉素重组片段（AB），asr1后臂和潮霉素重组片段（CB）的PCR扩增图；c图为pMD18-asr1：hyg质粒酶切检测图；
图4为本发明asr1基因敲除的PCR和real-time PCR检测图，A图为敲除菌株的PCR检测图，其中泳道从左到右是DNA分子量标准，野生型菌株为模板、基因敲除菌株为模板。B图为real-time PCR检测敲除菌株的基因转录情况；
图5为本发明中突变菌株耐离子胁迫能力检测图，其中A图为0.8 mol/L钾离子胁迫；B图为0.8 mol/L钠离子胁迫；C图为50 mmol/L钙离子胁迫；D图为7.5 mmol/L锰离子胁迫；E图为300 mmol/L镁离子胁迫。
具体实施方式
下面通过实施例对本发明作进一步详细说明，但本发明的内容并不局限于此，本实施例中方法如无特殊说明的均按常规方法操作，所用试剂如无特殊说明的采用常规试剂或按常规方法配置的试剂。
实施例1: 土生隐球酵母（C. humicolus）BSLL1-1菌株总RNA提取及cDNA的合成
使用TRIzoL试剂盒 (TaKaRa公司)进行酵母总RNA的提取，步骤如下：取土生隐球酵母菌体约0.2 g，加入研钵中用液氮研磨至粉末状，然后加入1 mL的TRIzoL提取液继续研磨至清澈。将研磨液移到EP管，室温静置约5 min，加入0.2 mL氯仿剧烈振荡1 min，然后将样品置于冰上5 min， 12000 rpm 4℃ 离心15 min；将上清转移至新的EP管中，再用氯仿进行一次抽提。取上清并加入等体积异丙醇，-20℃ 静止0.5 h后，12000 rpm，4℃ 离心30 min；弃上清，用75％的乙醇1 mL清洗两次，12000 rpm，4℃ 离心5 min，倒掉乙醇。自然晾干后用20-40 µL DEPC处理后的水进行溶解，-80℃保存。
用Reverse Transcriptase M-MLV 反转录试剂盒对提取的总RNA进行反转录。
1、在管中配制下列模板RNA/引物混合液；
2、70℃保温10 min后迅速在冰上急冷2 min以上；
3、离心数秒钟使模板/引物的变性溶液聚集于管底部；
4、在上述管中配制下列转录反应液，
5、42℃保温1 h；
6、70℃保温15 min后冰上冷却，得到的cDNA溶液可直接用于PCR扩增。
实施例2：荧光定量PCR分析asr1基因在铝胁迫下的表达
设计土生隐球酵母的18S rRNA和C2H2型转录因子Asr1编码基因的real-time PCR引物，18S rRNA基因作为内参，引物序列如下：
用SYBR Premix Ex TaqII（Tli RNaseH Plus）试剂盒进行real-time PCR，反应体系为20 µL（见下表），用Applied Biosystems 7500 Fast Real-Time PCR System使用两步法PCR反应程序：Stage 1：预变性，Reps：1，95 ℃ 30 s；Stage 2：PCR反应，Reps：40，95 ℃ 5s，60 ℃ 30 s；反应结束后确认real-time PCR的扩增曲线和融解曲线，进行PCR定量时制作标准曲线等。
以18S rRNA基因为内参，浓度梯度实验组不加铝离子培养48 h的基因表达量为对照，时间梯度实验组以不加铝离子培养6 h的基因表达量为对照。应用2-ΔΔCt法对实验结果进行分析，公式如下：
根据公式对照（CK）为“1”，并可算出不同时段或铝浓度下的基因表达差异。铝胁迫下C2H2 转录因子asr1基因real-time PCR分析结果得出，5、20、50、100和150 mM铝的基因表达量分别为对照的1.73、2.22、4.31、5.50和6.51倍，整体呈平稳上升上调表达，在150 mM时达到最大表达量为对照的6.51倍。用铝处理6、12、24、36、48 h后表达量分别为对照的1.05、2.92、3.16、3.34和3.12倍，与对照相比整体呈上调趋势，36 h出现了最大表达量是对照的3.34倍，如图1。
实施例3：Asr1转录因子asr1基因的克隆及测序
以土生隐球酵母cDNA为模板进行asr1基因的PCR扩增，扩增的引物为正向：AAGCTTATGCCGCCTGGACCGTCACCCAAAGAT(下划线为HindⅢ酶切位点)，反向：GGATCCCTAGAATGGGCATGGCCCACATTCGTT（下划线为BamHⅠ酶切位点）。反应条件：首先94 ℃预变性3 min，然后94 ℃、30 S，62 ℃、30 S，72 ℃、2 min进行30个循环，循环结束后72 ℃延伸10 min。得到的PCR扩增产物进行琼脂糖凝胶电泳（图2），用DNA胶回收试剂盒纯化目的条带。将目的片段连接到pMD18-T载体上，得到含有目的片段的重组载体pMD18-T-asr1。用热刺激法将其转化到大肠杆菌感受态细胞DH5α中，然后涂于含有氨苄青霉素的LB固体平板上，37 ℃倒置培养约12 h。挑取单菌落于液体LB培养基中培养约12 h后提取质粒，将质粒进行EcoRⅠ、SalⅠ双酶切检测，酶切检测正确重组载体送到上海生工生物有限公司测序，asr1核苷酸序列如序列表SEQ ID NO：1所示。
实施例4：重叠PCR扩增asr1同源重组基因片段
对asr1基因的前800 bp、后872 bp核苷酸和潮霉素基因分别设计扩增引物F1和R1，F3和R3，F2和R2，其序列如下：
F1：ATGCCGCCTGGACCGTCACCCAAAGAT，
R1：CTTCAATATCATCTTCTGTCGGCGAGAGGAGGTGCATGCC；
F2：GGCATGCACCTCCTCTCGCCGACAGAAGATGATATTGAAG，
R2：CTTGGGAGTCGCAAAGCCAAGATTTCAGTAACGTTAAGTG；
F3：CATTAACGTTACTGAAATCCTTGGCTTTGCGACTCCCAAG，
R3：CTAGAATGGGCATGGCCCACATTCGTT。
下划线部分为同源臂。
扩增各个片段的重叠延伸PCR步骤如下：
1、以asr1前臂和潮霉素基因为模板，先不加入引物在94 ℃ 3 min、94 ℃ 30s、62 ℃30 s、72 ℃ 2 min 15 s进行8个循环。然后加入引物F1、R2在相同的条件下进行25个循环，扩增asr1前臂：潮霉素重组片段；
2、以潮霉素基因与asr1后臂为模版，先不加入引物在94 ℃ 3 min、94 ℃ 30s、62 ℃30 s、72℃ 2 min 20s进行8个循环。再加入引物F2、R3在相同的条件下进行25个循环，扩增潮霉素：asr1后臂重组片段；
3、以前两步得到的重组片段为模板进行重叠延伸PCR，先不加入引物在94℃ 3 min、94℃ 30 s、60℃ 30s、72℃ 3 min 15s进行8个循环；然后加入引物F1、R3在相同的条件下进行25个循环，扩增得到asr1：hyg重组基因片段（图3）。
实施例5：asr1基因的敲除及敲除菌株的检测
将得到的重组片段连接到pMD18-T载体上进行TA克隆，酶切检测正确的质粒进行测序；在土生隐球酵母感受态细胞中加入测序正确的重组片段，置于冰上15 min；然后将混合液加入到0.2 cm冰预冷的电转杯中，进行电击（电击参数：25 µF、1500 V、200 Ω），电击后加入400 µL预冷的山梨醇，在超净工作台将混合液均匀涂于200 μg·mL-1 潮霉素含量的GM平板上，置于30℃培养箱倒置培养2-3 d；从200 μg·mL-1 潮霉素含量的GM平板上挑取单菌落，加入GM液体培养基30℃培养过夜，收集菌体进行PCR和real-time PCR检测；PCR结果显示（图4A），突变菌株扩增条带比目的条带大，这说明带有潮霉素基因的重组片段已经成功发生同源重组；Real-time PCR检测结果表明（图4B），与野生型酵母相比，突变菌株asr1基因的表达量仅是野生型菌株表达量的0.2倍，这也说明了基因敲除成功。
实施例6：突变酵母菌株耐离子胁迫能力的检测
在培养基中加入离子浓度为氯化钙：50 mmol/L、氯化锰：7.5 mmol/L、硫酸镁：300mmol/L、氯化钠：0.8 mol/L、氯化钾：0.8 mol/L，以不加任何离子的培养基作为对照。各配制100 mL的YPD培养基，将灭菌后的离子母液加入到灭菌的培养基中，混匀后倒平板；接种野生型和突变型菌株至20 mL的 YPD液体培养基中，28℃摇床培养过夜；取100 µL菌液转接至含有10 mL YPD 液体培养基的试管中，28℃摇床培养；将初始OD600=0.6的活化菌液进行稀释，稀释倍数分别为101 、102 、103 、104；每个浓度梯度菌液分别取2 µL培养物接种到含有各离子的YPD固体平板上，在28℃培养箱中倒置培养，观察菌落大小；以不加离子的YPD固体平板作为对照，每种设置三个平板作为重复；结果如图5所示，在含有K+、Na+、Ca2+、Mn2+、Mg2+的平板上，突变型比野生型菌落大，长势较好，这说明asr1转录因子突变后增强了土生隐球酵母菌株抗离子胁迫的能力。
序列表
<110> 昆明理工大学
<120> 一种C2H2型转录因子基因及其应用
<160> 10
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agctcagtgt gtcccgcatg cggaaagggc tttgcgcgcc ctgacgtgct ccgcaagcat 180
ctgtttacat cgtgcaaatc acgcaggact ggcgatggcg atccaccacc cccgctcaca 240
accgccgaag aagatgccct ggtaccacgc aagcgtagac ggccacagcg cagactcacc 300
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caccatgccc ctcctcaatc cccaatgcac gggcactctc accctcacca acacccacat 900
caatcagcga accaccgtct cggtcctgcc cgcgcaggtg gcagcctcga agtcctttta 960
gccccggcgt ttgcaacgac gcccgagacg acgtttggtt ttggctttgc gactcccaag 1020
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ctgccaccgc cgcatggcct tgcaggggtt cacggcggcg gcagcggcag cggcagcagc 1140
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ctcgagacct ctgccaatgt ggctgtccag cagggttttg gggtggctgg cctctcaccc 1260
gacctcccgg tgccaacaac tacgggtgat gccagcttaa aaagatactc tagcactggg 1320
aaccccgatg gtggccaagg acgtgacgac agattcggcg ccgtaggaga ctactacgcg 1380
tccacagctc aacgctctac ctctgggaac ttttttggtg atggggttgg gccaccaata 1440
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catcgtcgcc tccccgactt ttgtgcccga gacacgcctt gtgccgcggt catgggagtc 1560
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gcccgctggt ctcccgacaa catttcagca gagcaggtca agactccgga cgaagagacc 1680
cgggtcacga ttctcgaagt cgtggagggc gcgacatttt ccgtcccact caaccatccc 1740
ttggcaaaca acaccttggt gacacccgcc cagaaacctc gcccacctgc gccccgtctg 1800
aactttaccg cgccgccgtt tccgccacac cctgaagatg ggtcaaacga atgtgggcca 1860
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Claims (4)
1.一种C2H2型转录因子基因asr1，其核苷酸序列如SEQ ID NO：1所示，编码如SEQ IDNO：2所示氨基酸序列的蛋白质。
2.对权利要求1所述的C2H2型转录因子基因asr1进行阻断获得的同源重组基因片段。
3.一种土生隐球酵母突变菌株，所述突变菌株含有权利要求2所述的同源重组基因片段。
4.权利要求2所述的同源重组基因片段在增强微生物抗离子胁迫能力中的应用。
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