CN104195152A - 一种钙调磷酸酶催化亚基基因及其应用 - Google Patents
一种钙调磷酸酶催化亚基基因及其应用 Download PDFInfo
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- CN104195152A CN104195152A CN201410423663.4A CN201410423663A CN104195152A CN 104195152 A CN104195152 A CN 104195152A CN 201410423663 A CN201410423663 A CN 201410423663A CN 104195152 A CN104195152 A CN 104195152A
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Abstract
本发明公开一种钙调磷酸酶催化亚基基因,其是在土生隐球酵母BSLL1-1菌株中克隆的,其核苷酸序列如SEQIDNO:1所示,编码如SEQIDNO:2所示氨基酸序列的蛋白质;将该蛋白在大肠杆菌中表达,并进一步将该基因在酿酒酵母中成功表达,该基因使酿酒酵母耐铝能力增加,基因工程菌可以吸附(或者吸收)培养介质中的活性铝,该酿酒酵母基因工程菌具有降低酸性土壤中活性铝含量的应用潜力。
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钙调磷酸酶催化亚基的耐铝基因,核苷酸序列和氨基酸序列,以及耐铝蛋白的异源表达,涉及含有该基因的重组质粒和表达该耐铝基因的工程菌株,它们的制备和表达方法,以及它们在吸附环境中活性铝的应用。
背景技术
酸性土壤在世界上广泛存在,其中可耕种面积为1.79亿hm2。我国酸性土壤的分布遍及14个省区,约占全国耕地面积的21% (熊毅和李庆逵,1987)。氮肥的过量施用是导致农田土壤酸化的最主要原因。此外,工业化发展导致的酸雨、人类不良的耕作方式和阳离子在土壤中的淋失,使酸性土壤的面积不断扩大,酸化程度不断加剧。
铝(aluminum,Al)在地壳中分布广泛,约占地壳总质量的7%。在土壤中,铝的主要存在形态是不溶于水的氧化物或铝硅酸盐,这些形态是对植物和环境没有伤害的。但是随着土壤酸化程度的增加,土壤中的铝从不溶于水的形态中溶解出来,转化为可溶于水的无机离子态,如Al3+、(AlOH)2+、(AlOH)2 +,这些形态对植物根系的毒害作用最大,又称为活性Al。因此,在酸性土壤上铝毒害是影响农作物产量的主要限制因素之一。通常,解决铝毒害的方法是大量施用石灰来提高土壤的pH值,使游离铝沉淀。但是这种方法难以彻底解决土壤酸度和铝毒害问题,同时还存在着潜在的环境问题。
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。因为微生物种类多,代谢类型丰富,生长在酸性环境中的微生物在长期的进化过程中,为了保护细胞免受铝的毒害,产生了一系列的抗铝毒机制:如有机酸及其代谢产物对铝的螯合作用;抗氧化胁迫作用;抗细胞程序性死亡等。因此,筛选或者改良土壤微生物使其增加对铝的耐受性或者提高其对土壤中铝的清除能力,是解决酸性土壤上铝毒害的直接而有效的措施。
钙离子(Ca2+)作为动植物及微生物细胞内重要的第二信使,参与了胞内很多生理反应。Ca2+作为离子信号分子,需要下游受体才能将信号传递下去,并调节多种细胞反应和生物进程。钙调素(Calmodulin, CaM)是真核细胞内高度保守的、广泛存在的一种非常重要的钙离子受体蛋白。CaM自身不具有酶的活性,但其能与靶蛋白相互作用,从而对靶蛋白的活性进行调节。当细胞内的Ca2+ 浓度升高时,Ca2+会与CaM的EF-hand结构域结合,直接导致钙调素结合蛋白(calmodulin binding protein, CaMBP)的氢键网络结构被破坏,CaM结合结构域被释放出来,结合了四个Ca2+的CaM再通过CaM结合结构域与CaMBP相结合,从而将信号传导下去。钙调磷酸酶(Calcineurin,CaN)是钙调素下游结合靶蛋白的一种,是特异性的钙/钙调蛋白依赖的苏氨酸/丝氨酸蛋白磷酸酶。钙调磷酸酶是一个异源二聚体蛋白,由一个71KDa的催化亚基A(CNA)和一个19KDa的调节亚基B(CNB)组成。CNA含有四个功能区:分别是催化结构域;CNB结合结构域;钙调素结合结构域(CBD);自抑制域(AID)。钙调磷酸酶的调节亚基 CNB是一个具有 EF-hand 结构的蛋白质,能与 Ca2+结合。Ca2+存在时,B 亚基与 A 亚基结合后,会使调节片段中的自抑制结构域(AID)从催化活性中心上脱落,从而降低了自抑制结构域对酶活性的自抑制作用。
钙调磷酸酶在很多细胞和发育过程中发挥重要作用,其主要作用是通过转录因子Crz1调节基因的表达,从而使胁迫条件下细胞存活。在白色念珠菌中,钙调磷酸酶的主要作用是响应各种刺激或胁迫(Zelteret al., Fungal GenetBiol, 2004, 41: 827-841; Stie and Fox, Eukaryot Cell, 2008, 7: 177-186)。在高温、高pH以及钙盐离子胁迫的情况下,钙调磷酸酶对于酿酒酵母的生长是必需的(Bonilla et al., EMBO, 2002, 21:2343-2353)。在酿酒酵母中乙醇胁迫可以激活钙离子介导的钙调磷酸酶/Crz1途径(Araki et al., J Biosci Bioeng, 2009, 107(1):1-6)。通过研究钙调磷酸酶对人类主要病原菌念珠菌毒力的影响,结果发现编码钙调磷酸酶催化亚基(CMP1/CNA)的白色念珠菌突变体和编码钙调磷酸酶调节亚基(CNB1)的白色念珠菌突变体,对于盐胁迫、碱胁迫和渗透胁迫是敏感的(Bader et al., Infect Immun, 2003, 71:5344-5354; Blankenship et al., Eukaryot Cell, 2003, 2:422-430)。这些结果表明,钙调素信号途径可能是生物体耐受逆境胁迫的一种普遍机制,过表达该途径上的基因会提高对逆境胁迫的耐受能力。因此,克隆这些基因对研究它们在抗铝毒中的作用具有非常重要的意义。同时,对克隆到的基因进行改造或者构建基因工程菌株,还可以为治理酸性土壤上的铝毒害问题提供理论和应用基础。
发明内容
本发明旨在提供一种钙调磷酸酶催化亚基基因,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1917bp,与黑白轮枝菌(Verticillium alboatrum)丝氨酸/苏氨酸蛋白磷酸酶(PP2B)基因同源性达到81%,编码钙调磷酸酶催化亚基(CNA)蛋白由638个氨基酸组成,分子量约为77KDa。
本发明的另一目的在于提供一种钙调磷酸酶催化亚基基因的重组表达载体pYES3/CT- CNA。
本发明另一目的是提供一种含有钙调磷酸酶催化亚基基因或上述重组表达载体的酿酒酵母工程菌株。
本发明另一目的是将钙调磷酸酶催化亚基基因应用在吸附环境中活性铝中。
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CNA基因组序列。根据该序列设计引物,用土生隐球酵母的cDNA为模板,用设计好的引物扩增CNA基因片段。将扩增片段连接到pMD-18T载体上并转化大肠杆菌DH5a,然后涂布含氨苄青霉素的平板,挑取阳性菌落测序,鉴定出编码CNA蛋白的基因,其序列如SEQ ID NO:1所示,该蛋白由638个氨基酸编码,氨基酸序列如SEQ ID NO:2所示;
2、以土生隐球酵母cDNA为模板,用特异引物扩增CNA片段,然后与经限制性内切酶酶切的pET-32a载体连接并转化大肠杆菌BL21(DE3)菌株,获得重组质粒pET-32a-CNA和含重组表达质粒的重组大肠杆菌菌株BL21(DE3)- pET-32a-CAN;
3、从测序正确的pMD18-T-CNA质粒上酶切回收目的片段,连接到经限制性内切酶酶切的酵母表达载体pYES3/CT质粒上,得到重组质粒pYES3/CT-CNA,将重组质粒用电击法转化酿酒酵母(INVSc1)感受态细胞,用Western blotting法在转基因酵母中检测到了目的条带的存在,从而成功得到含pYES3/CT-CNA表达载体的重组酵母工程菌株INVSc1-pYES3/CT-CNA。
本发明的优点和技术效果如下:
本发明的钙调磷酸酶催化亚基基因可以增加酵母的耐铝能力,钙调磷酸酶催化亚基基因工程菌株可以通过吸附作用降低环境中活性铝的含量,这种利用微生物修复环境中活性铝的技术费用低,操作简便,对环境影响小,不会造成二次污染。
附图说明
图1为本发明中验证CNA和CaM之间相互作用的GST-pull down检测图;图中:GST为纯化的GST蛋白;GST-CAM为纯化的GST-CaM融合蛋白;
图2为本发明原核表达载体 pET-32a-CNA质粒及其BamHI/HandIII双酶切鉴定电泳检测图,其中A图中对照为对照质粒;1,2,3为pET-32a-CNA质粒;B图中marker为DNA分子量标准;2为2号质粒BamHI/HandIII双酶切;
图3为本发明 CNA重组蛋白在大肠杆菌中的表达电泳检测图,其中Marker为蛋白分子量标准;0h, 2h, 4h, 6h, 8h为诱导时间;对照为转空载体的菌株;上清为诱导菌体上清液;沉淀为诱导菌体沉淀;
图4为本发明转基因酵母的构建及检测图,其中图A为 pYES3/CT-CNA质粒的酶切检测图,M为 DNA分子量标准;1, 2, 3, 4为质粒BamHI/HandIII酶切;图B为 Western blotting 检测转基因酵母中CNA蛋白的表达;
图5为本发明转基因酵母耐铝能力检测图;
图6为本发明转基因酵母吸附(或吸收)铝能力的检测图;其中:图A为0.2mM铝时转基因酵母和对照酵母培养基中剩余铝含量;图B为2mM铝时转基因酵母和对照酵母培养基中剩余铝含量。
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CNA基因的克隆和鉴定
提取土生隐球酵母菌基因组DNA(奥斯伯等,精编分子生物学实验指南(第五版),北京:科学出版社,2008),送上海人类基因组研究中心进行基因组测序,得到CNA基因组序列,根据该序列设计引物,引物序列如下:正向引物CNA-F:5’- GGATCCATGACCTCTCCGGCGACCCAGC-3’ (下划线为BamHI酶切位点),反向引物CNA-R:5’- AAGCTTTTAGGCAATAGAGTTCTCGG-3’(下划线为HandIII酶切位点)。用土生隐球酵母的cDNA为模板,用设计好的引物扩增CNA基因片段。PCR反应条件为:94℃预变性3min,然后按照94℃变性30s,65℃退火30s,72℃延伸2min的程序进行30个循环,循环结束后72℃反应10min。CNA基因的PCR产物克隆到pMD18-T载体上,获得TA克隆载体命名为pMD18-T-CNA,然后对获得的TA克隆载体进行酶切验证,对酶切检测正确的pMD18-T-CNA质粒送北京六合华大基因科技股份有限公司测序,CAN核苷酸序列如序列表SEQ ID NO:1所示。
实施例2:验证CNA和CaM的互作
合成CNA的多肽611-627:RLAEVISSPTKGGQGER,免疫雄兔,制备CNA抗体;
1)免疫:选取3.0kg左右的新西兰大白兔,免疫之前,耳静脉取阴性血清,取1000μg钙调磷酸酶催化亚基的多肽,用生理盐水稀释到1000μl,再加入等体积弗氏佐剂(初次免疫用弗氏完全佐剂,加强免疫用弗氏不完全佐剂)。用研钵将溶液和佐剂研磨混匀,使溶液形成油包水的状态。将混匀好的免疫原进行皮下注射,一般免疫三次,前两次免疫间隔两周,最后一次免疫间隔一周。第一次免疫在脚掌部注射打2-3个点,后两次在背部免疫打4-5个点。
2)心脏采血:将兔仰面,四肢缚于动物固定架上(或由助手抓住四肢)。用乙醚麻醉,剪去左胸部毛发,消毒皮肤。剪开皮肤,将心脏剥离,用50ml注射器(连接16号针头)倾斜进针,对准心搏最强处刺入心脏抽血。将抽取的血液立即注入无菌50ml离心管中,待凝固后分离血清。
3)分离血清:于10000rpm,4℃,离心15 min,取上清,分装后置-20℃冰箱中保存备用。
4)多抗效果检测:用Western blotting方法检测得到的多克隆抗体,用阴性血清作为负对照。检测得到的抗体效果很好,可以用于做后面的实验。
验证CNA和CaM之间的相互作用。Glutathione S-transferase(GST)和GST-CaM在大肠杆菌中诱导表达,并且分别用GST亲和层析柱纯化获得纯度较高的目的蛋白。分别取50 μg纯化的GST和GST-CaM融合蛋白与20 mM Al处理12 h的土生隐球酵母总蛋白500 μg在结合缓冲液 (50 mM Tris-Cl,pH 7.5,100 mM NaCl,0.25% TritonX-100,1mM EDTA,1mM DTT)中共同震荡孵育2 h;加入30 μl GST琼脂糖结合树脂在4℃摇床(40 rpm)孵育过夜;4℃,3500g离心5min收集沉淀物,收集的沉淀蛋白混合物用结合缓冲液洗涤三次;沉降下来的蛋白质混合物加入10 μl SDS凝胶加样缓冲液(Tris-HCl 50 mM,pH 6.8;SDS 2 %;DTT 100 mM;溴酚蓝0.1 %;甘油10 %),用沸水浴煮;在12%的SDS-PAGE胶分离所沉淀的蛋白质;分离后的蛋白转移到PVDF膜上,然后分别用anti-CNA特异性抗体孵育膜,随后在Chemidoc XRS(BIO-RAD)中进行成像观察。结果表明,CNA可以被GST-CaM蛋白沉淀;然而负对照GST蛋白没有沉淀到CNA(图1)。因此,CaM和CNA在体外是可以相互作用的。
实施例3:CNA蛋白的异源表达
以土生隐球酵母的cDNA作为模板,用特异引物CNA-F和CNA-R扩增CNA片段,然后与经限制性内切酶BamHI/HandIII酶切的pET-32a载体连接并转化大肠杆菌BL21(DE3)菌株,获得重组质粒pET-32a-CNA。提取质粒进行BamHI/HandIII双酶切鉴定,检测到与目的条带和载体片段同样大小的两条带,如图2所示,说明原核表达载体pET-32a-CNA构建成功。将重组质粒pET-32a-CNA转化大肠杆菌BL21(DE3),得到含重组表达质粒的重组菌株大肠杆菌BL21(DE3)- pET-32a-CNA。
将重组大肠杆菌BL21(DE3)- pET-32a-CNA菌株和对照菌株即含有pET-32a质粒的大肠杆菌BL21(DE3) - pET-32a,接种于含有氨苄青霉素的LB液体培养基中,200 rpm、37℃培养过夜,再按1%的接种量转接于新鲜的LB液体培养基中,20℃培养至OD600为0.6-0.8时,加入ITPG至终浓度为1mM,对含有CNA目的基因的表达载体进行诱导表达;分别收集诱导0h、2h、4h、6h、8h后的表达菌体2ml,收集后的菌液于4 ℃、12 000 rpm离心1 min,弃上清液,沉淀用2M尿素100μl重悬,之后再加20μl SDS凝胶加样缓冲液,煮沸5 -10min后,12 000 rpm离心1 min,取20μl上清进行SDS-PAGE分析,确定菌体的最优表达时间。本实验以含有目的基因的表达菌诱导0 h和含pET-32a空载体的大肠杆菌诱导表达6 h作为对照。随后,对收集的菌体进行超声波破碎,4℃,12000rpm离心20min,收集上清与沉淀,SDS-PAGE检测蛋白的表达形式。SDS-PAGE分离胶的浓度为12%,浓缩胶浓度为4%。SDS-PAGE结果表明,CNA基因编码的蛋白在大肠杆菌BL21(DE3)中大量表达,且在上清和沉淀中都有表达(图3)。重组蛋白的最佳表达条件为20℃,1mM IPTG,诱导时间为6h。
实施例4:CNA转基因酵母的构建及检测
将测序正确的pMD18-T-CNA质粒用BamHI和HindIII酶切,回收CNA 片段,用BamHI和HindIII酶切pYES3/CT质粒,回收得到带有BamHI和HindIII酶切位点的线性pYES3/CT片段,然后将二者进行连接反应,得到pYES3/CT-CNA 质粒。用BamHI和HindIII对pYES3/CT-CNA 质粒酶切,酶切产物电泳检测到了目的条带(图4A),说明外源基因成功插入到酵母表达载体上。
将检测正确的pYES3/CT-CNA质粒电击转化酿酒酵母INVSc1感受态细胞,在SD-Trp平板上长2-3天,挑取单菌落,30℃培养过夜,收集菌体后提取总蛋白,用CNA抗体进行Western blotting 检测,在转基因酵母中检测到了与目的蛋白大小的条带,结果如图4B所示,这进一步验证了外源基因在酿酒酵母细胞得到了成功表达。
实施例5:转基因酵母耐铝能力检测
配制铝浓度为0、0.1、0.2、2mM 的YPD诱导固体培养基(碳源为半乳糖),将活化的菌体按初始OD600为1,稀释倍数分别为100 、10-1 、10-2 、10-3 、10-4,将稀释的菌液5μl点种到平板上,48h后观察生长情况,从图5中看出,随着铝浓度的增加,转基因酵母INVSc1-pYES3/CT-CNA耐铝能力都明显高于对照酵母INVSc1-pYES3/CT。在含有0.1mM铝的固体平板上,与对照酵母相比,转基因酵母菌落较大,表明对照酵母生长受到抑制。在含有2mM铝的固体平板上,稀释倍数为10-4 时,pYES3/CT-CNA 转基因酵母生长良好,对照酵母基本不生长。随着铝浓度的增加,转基因酵母生长状况比对照酵母好,说明在铝胁迫下CNA基因有促进生长的作用。
实施例6:转基因酵母吸附(或吸收)铝的检测
通过检测培养基中剩余活性铝的含量,进而检测pYES3/CT-CNA转基因酵母吸收或吸附铝的能力。分别用含有0.2mM和2mM铝的培养基作为对照,其培养基中剩余铝含量定义为100%。在含有0.2mM铝时,pYES3/CT-CNA转基因酵母培养基中活性铝剩余量为33.54%,与转空载体pYES3/CT酵母相比,pYES3/CT-CNA转基因酵母的培养基中剩余的活性铝明显减少。在含有2mM铝时,pYES3/CT-CNA转基因酵母培养基中活性铝剩余量为74.19%,与转空载体pYES3/CT酵母相比,pYES3/CT-CNA转基因酵母的培养基中剩余的活性铝也明显减少(图6)。这些结果表明,在转基因酵母中,CNA基因可能通过增加菌体对铝的吸附或者吸收来达到耐铝的目的。
序列表
<110> 昆明理工大学
<120> 一种钙调磷酸酶催化亚基基因及其应用
<160> 4
<170> PatentIn version 3.3
<210> 1
<211> 1917
<212> DNA
<213> 土生隐球酵母BSLL1-1
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attgtcattc cagagatcga cttcacccag cacacgctcg agaatggcga gattgtcagc 120
accaccgagc gcgtggtcaa ggacgtgcaa gcgccggcca tgtatgtgcc cacagacgag 180
cagttctggt ccaagcagga ccccacaaag cccgacattg ccttcctcaa gaaccacttc 240
taccgcgagg gacgcttgtc cgaggagcag gcgctgtaca tcctcgagaa gggaggcgag 300
attctcaagt cggaacccaa cctgctcgag gtcgatgcgc cgatcactgt ctgtggtgac 360
attcacggcc aatactatga cctcatgaag ctgttcgagg tcggtggcaa ccctgccgac 420
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ctgtacctct ggtcgctcaa gatgtggtac cccgacacgc tgttcctcct ccgcggtaac 540
cacgagtgtc gccacttgac cgactacttc acgttcaagc tcgagtgcaa gcacaagtac 600
tcggagacgg tgtacaacgc ctgcatggac acgttttgca acctgcccct tgccgccgtc 660
atgaacaggc agttcctctg cattcacggc ggtttgtcgc ctgagctcca cactctggat 720
gacctcagaa ccattaaccg cttccgcgag ccccccaccc acggcctgat gtgcgacatc 780
ctctgggccg acccattgga agactttggc aatgagaaga acccgaccga gaactttgtg 840
cacaaccatg ttcgcggctg ctcgtacttc ttcacgtaca acgccgcctg ccagttcctg 900
gagcgcaaca acctgttgtc catcatccgt gcccacgagg cccaggacgc tggataccgc 960
atgtacaaga agaccaagac tactggcttc ccctcggtca tgaccatctt ctcggcgccc 1020
aactacctcg acgtgtactc gaacaaggcg gccgtcctca agtacgagtc gaatgtcatg 1080
aacattcgtc agttcaactg cacgccgcac ccctactggc tccccaactt tatggacgtc 1140
ttcacctggt cccttccttt cgtcggcgag aagattaccg acatgctgat cgccatcctc 1200
aactgttgca ccaaggagga gctcgaggag gaggaggagg agacgccgat gatcacgcca 1260
gactcccccg aggagggcga ggacggagac atctcggcgg agaggagaca ggttatcaag 1320
aacaagattc tcgctgttgg tcgcatgtcg cgagtgttct cgttgcttcg tgaggagtcg 1380
gagcgagtgt cggagctcaa gagcatcact gggtctgcca acctgccgca gggtgcgctt 1440
gccaacggtg cggagggtat caaggaggct atccagggct ttgacgacgc gcgtaagagt 1500
gatatcgaga acgagcgcct tcctccggac atcatcgacc ccgatgagga caagcctgcg 1560
tcgccatcgc cgtctgtgcc gtcgtcgcct gctccccaca cgggatcgcc cattgccgac 1620
aagactggca cagccttgcc acctctcaac acgaatgtgt ctgcgagcga tattgcgtcg 1680
cctgtgtcgc cggcgacgcc aggagggacg ggttggcgcc gtggacacgg ccgccaggct 1740
tcgctcggta ccacacgcac gtcgccctcg acgcgtcgcc gttcgctcga gaacacgatg 1800
gagctcatcc gcgacgttgt tggcggccga gacgccaatg ccgactcgaa cgtgcgcgag 1860
cttgccgagg tcatctcgag cccgtcgcgg acccggcccg agaactctat tgcctaa 1917
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aagcttttag gcaatagagt tctcgg 26
Claims (4)
1.一种钙调磷酸酶催化亚基基因,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。
2.一种含有权利要求1所述的钙调磷酸酶催化亚基基因的重组表达载体。
3.一种酿酒酵母工程菌株,所述工程菌株含有权利要求1所述的钙调磷酸酶催化亚基基因或权利要求2所述的重组表达载体。
4.权利要求1所述的钙调磷酸酶催化亚基基因在吸附环境中活性铝中的应用。
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CN108588109A (zh) * | 2018-04-10 | 2018-09-28 | 昆明理工大学 | C2H2型转录因子基因asr1的重组表达载体及应用 |
CN111850140A (zh) * | 2020-08-10 | 2020-10-30 | 西北农林科技大学 | 一种山羊PPP3CA基因InDel标记的检测方法及其应用 |
CN114045294A (zh) * | 2021-11-22 | 2022-02-15 | 昆明理工大学 | 一种脂质转运蛋白基因及其应用 |
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