CN102603873B - 一种重金属镉抗性相关蛋白DbsCzcA及其编码基因和应用 - Google Patents
一种重金属镉抗性相关蛋白DbsCzcA及其编码基因和应用 Download PDFInfo
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Abstract
本发明公开了一种金属镉抗性相关蛋白DbsCzcA及其编码基因和应用。该镉抗性相关蛋白DbsCzcA氨基酸序列如SEQIDNO:1所示,其编码基因如SEQIDNO:2所示。通过转化大肠杆菌实验证明该基因在大肠杆菌中的表达可以提高其对重金属镉的抗性。本发明的DbsCzcA编码基因可用于提高微生物和植物对重金属镉的抗性,进一步用于清除环境重金属污染,为生物修复提供性能优良的生物资源。
Description
技术领域
本发明属于微生物基因工程,和生物修复领域。具体地说,本发明提供了生活在酸性矿山废水(acid mining drainage,AMD)的一种未知微生物所含有的一种对重金属镉抗性的基因即阳离子外排系统蛋白基因(DbsCzcA)的序列,以及该序列推测编码蛋白质分子的氨基酸序列。本发明在提高微生物和植物对重金属镉的抗性方面有着重大的应用价值。
背景技术
随着矿产资源的开发利用、工业发展,重金属对环境造成的污染日趋严重,土壤重金属污染已经成为一个危害全球环境质量的问题。土壤重金属会影响植物的生长发育,降低农作物的产量和质量,带来了严重的经济损失。此外,受土壤重金属污染的作物在植物体中积累,并通过食物链富集到人体和动物体中,危害人畜健康,引发癌症和其他疾病等。治理重金属污染刻不容缓,各种修复技术和措施正在研究和应用中。各国政府和科学家着力通过两个途径解决这一问题:一为利用物理的,化学的方法试图清除土壤或水体的重金属污染:二为利用现代生物技术清除污染。自从20世纪80年代以来,生物修复技术因其具有处理费用低、对环境影响小、效率高等优点,越来越受到广大科技人员的广泛关注。生物修复一般分为植物修复、动物修复和微生物修复三种类型,其中植物修复和微生物修复是研究的热点。微生物修复就是利用微生物将环境中的污染物降解或转化为其他无害物质的过程。近年来,基于微生物对重金属的作用机理,以修复有毒有害金属污染或回收有经济价值重金属为目的的生物处理技术日趋成熟。植物修复指利用植物去治理水体、土壤和底泥等介质中的污染的技术。然而用于重金属污染修复的生物往往会受到重金属的毒害,生长缓慢、生物量小,甚至不能生存,所以重金属对植物和微生物的毒害作用是生物修复的主要限制因素。
解决生物修复中重金属对生物的毒害作用的根本途径在于研究耐受重金属的分子生物学机制,克隆对重金属耐受的关键基因,通过基因工程手段获得用于生物修复中性能优良的转基因工程生物。
由于技术上的原因,直到最近,对微生物基因资源的利用主要局限于可培养微生物。然而,已培养微生物仅占自然界中微生物的不到1%,因此各种生境中的微生物宏基因组是一个巨大而未发掘的基因资源库。极端环境具有丰富的微生物资源,当中许多与逆境和关键生命过程相关的基因在长期的适应进化中获得了更强的耐性潜能,发掘这些抗性基因已成为国际重要的研究热点。酸性矿山废水(AMD)是极端生境微生物学研究的重要系统。AMD来源于采矿活动使含硫矿物(主要为黄铁矿,FeS2)暴露于空气和水中,在微生物催化作用下迅速氧化产酸所致,其pH值一般在1-4左右,而且富含硫酸盐以及Pb、Zn、Cu、Cd和Ni等重金属,是采矿业面临的最严重环境问题之一。在AMD中生存的原核微生物在长期的进化过程中逐渐形成了一些独特机制,以应对低pH值、高盐度以及高重金属等多种极端环境协迫。因此,AMD生境成为极具特色和丰富的抗逆基因库。
镉(Cd)是一种极其重要的工业和环境化学污染物,因其对环境水、空气和土壤的污染而在动植物体内蓄积,最终导致对人类健康的危害,并由于它在体内能长期蓄积,不易排除,故易产生慢性的和远期的病理效应,可损伤许多组织和系统。把生物体内的镉离子快速排出是一种有效的解毒方式。
Czc是一种膜结合的蛋白复合体,它通过主动运输把细胞内的Co2+,Zn2+和Cd2+排出到细胞外,从而解除这些重金属离子的毒性。在这个蛋白复合体中,CzcA编码一个阳离子/质子对流泵,CzcB编码一个亚基与阳离子结合,CzcB主要与Zn2+结合,CzcC与CzcB结合,使CzcB构象发生改变,能与Co2+和Cd2+结合。,CzcD、CzcE、CzcS和CzcR编码四个CzcCBA操纵子,对Czc复合体起着调控的作用。但CzcD、CzcE、CzcS和CzcR对这个Czc蛋白复合体的表达并不是必需的。
Nies等发现CzcR and CzcD影响Alcaligenes eutrophus通过Czc重金属外排系统对钴,锌,镉的抗性。Legatzki研究表明Ralstonia metallidurans的Czc系统与另外两个P型ATP酶相互影响,对提高它对重金属的抵抗力都有一定的作用。在Ralstonia metallidurans中czcNp启动子控制着Czc复合体的表达,从而决定着它对重金属的抵抗能力。
发明内容
本发明的目的在于提供AMD微生物中的阳离子外排系统蛋白CzcA,以及编码该蛋白的新基因,为今后开发基因工程产品奠定物质基础。
在本发明的一个方面,提供了一种酸性矿山废水微生物中阳离子外排系统蛋白DbsCzcA,它为如下蛋白分子(i)或(ii):
(i)具有入SEQ ID NO:1所述的氨基酸序列;
(ii)在SEQ ID NO:1限定的氨基酸序列经过取代,缺失或叠加一个或几个氨基酸衍生的蛋白质与(i)的蛋白质具有相同的功能。
在本发明的另一个方面,提供了一种DNA分子,它包括:编码所述的阳离子外排系统蛋白CzcA的核苷酸序列。
发明发现DbsCzcA蛋白具有金属镉抗性作用,可以在治理环境镉污染中应用。
发明同时保护了该蛋白的编码基因,具有如SEQ ID NO:2所示的序列;以及该基因在治理环境镉污染中的应用。
发明用表达DbsCzcA的基因工程菌作为试验对象,发现在镉胁迫下,其镉耐受性大大高于非基因工程菌。在镉离子浓度为200μM以下时,尤其是150M以下时,仍有较高的存活率。
与现有技术相比,本发明具有以下有益效果:
本发明从AMD微生物中克隆了一个新的CzcA基因,并对其对重金属镉的抗性进行了功能分析。通过大肠杆菌转化实验证明该基因可以提高大肠杆菌中对镉的抗性。
应用本发明的基因序列或氨基酸序列进行转基因开发基因工程产品面具有重大的应用价值,具体来说可用于培育高生物量的重金属超富集植物或微生物,用于重金属污染土壤和水体的生态修复。
附图说明
附图1为DbsCzcA在大肠杆菌中的表达时相;其中:
M:蛋白分子量标准
1:携带空载体pET28a的BL21(DE3)菌株
2-9:携带pET28a-DbsCzcA的BL21(DE3)菌株分别诱导表达0、1、2、3、4、5、6、7小时。
附图2为表达DbsCzcA的大肠杆菌在不同镉浓度下培养12小时后BL21(DE3)的生长情况。
附图3为表达DbsCzcA的大肠杆菌在150μM镉胁迫下的生长曲线。
具体实施方式
实施例1DbsCzcA基因全序列的克隆
野外微生物样品采集:
在云浮铅/锌矿选取不同酸化阶段(以pH为标准)的AMD,使用0.22μm的滤膜收集20LAMD里面的细胞。为了保持核酸的完整保存,保存样品冻于液氮中,24小时内带回实验室,并放于-70℃冰箱长期保存。
核酸的提取:
DNA提取采用SET方法,过程如下:向SET buffer中加入溶菌酶及蛋白酶K,消化30min后,15,000rpm离心15min后用氯仿抽提2次,用异丙醇沉淀过夜后,75%乙醇清洗2次,最后溶于灭菌水中。用Qiagen tip-100柱纯化回收基因组DNA,用核酸蛋白分析仪检测DNA质量及浓度。
基因组测序:
用Roche公司的GS FLX Titanium General Library Preparation Kit制备上机样品。使用Roche 454Genome Seqencer FLX测序仪,通过进行测序,用Pyrobayer软件获取碱基序列。
基因组序列分析:
去除低质量的测序结果后,对基因组进行如下分析:
序列拼接:使用Euler-SR及454公司的GS De Novo Assembler Software进行序列拼接。用N50指数评价拼接的效果;
基因组注释:将拼接好的微生物的全基因组序列用IMG和SEED系统进行基因组的注释,发现新的基因。
DbsCzcA基因片段的克隆:
通过PCR方法:以含目的基因的克隆质粒为模板,按基因序列设计一对引物(在上游和下游引物分别引入不同的酶切位点),PCR获得一条3kb大小的条带,并克隆到PCR2.1(购自invitrogen公司)载体中,序列委托invitrogen公司测定。
DbsCzcA基因序列分析:
测序结果表明CzcA基因大小为3240bp(见SEQ ID NO:2),编码1079个氨基酸(见SEQ ID NO:1)。
实施例2DbsCzcA基因的功能分析
本实施例中利用大肠杆菌转化实验分析DbsCzcA基因的功能。
构建重组表达载体:
设计以下一对引物,在DbsCzcA基因5’引入BamHI酶切位点,3’引入XhoI酶切位点。
DbsCzcA-F(SEQ ID NO:3):
5’CGCGGATCCATGCTCAAAGCCATTTTAGCCT3’
DbsCzcA-R(SEQ ID NO:4):
5’CCGCTCGAGCTGATCTTCCTCATCCATCTGG3’
以PCR2.1-DbsCzcA载体为模板进行PCR。分别将PCR产物和酵母穿梭表达载体pET28a用BamHI和XhoI进行酶切,将酶切产物回收、连接、并转化大肠杆菌DH5α。经测序并酶切鉴定,得到了pET28a-DbsCzcA重组子。
蛋白表达:
将重组子pET28a-DbsCzcA转化大肠杆菌表达菌株BL21(DE3),通过PCR验证筛选阳性克隆。挑取含重组质粒的菌体单斑至10ml LB(含Kan50μg/ml)中37℃过夜培养。将1ml菌液加入到含100ml LB培养基(含Kan50μg/ml),37℃震荡培养至OD600约为0.4-1.0(最好0.6,大约需2hr)。
加入IPTG至终浓度为1mM进行诱导,每隔1小时收集1ml菌液,离心12000g×60s收获沉淀,用80μl ddH2O重悬,加入20μl 5×SDS-PAGE上样缓冲液,混匀,沸水浴10min。取上清作为样品做SDS-PAGE分析(见附图1)。
转化子对重金属镉抗性的测定:
挑取含重组质粒的菌体单斑至10ml LB(含Kan 50μg/ml)中37℃过夜培养。将100ul菌加入到10ml Cd2+浓度分别为0μM、50μM、100μM、150μM、200μM的LB培养基(含Kan 50μg/ml、IPTG 1mM)中,37℃、200rpm震荡培养12小时,分别测定OD600值。
根据实验结果,选取Cd2+浓度为150μM作为胁迫条件,进行了生长曲线的测定。挑取含重组质粒的菌体单斑至10ml LB(含Kan 50μg/ml)中37℃过夜培养。将100ul菌加入到10ml Cd2+浓度为150μM的LB培养基(含Kan 50μg/ml、IPTG 1mM)中,37℃、200rpm震荡培养,每隔2小时测定OD600值。
实验结果表明,表达DbsCzcA基因的BL21(DE3)在0-200μM Cd2+浓度下都可以正常生长,而空载体对照在Cd2+浓度超过150μM时便不能生长(见附图2)。在150μM Cd2+浓度下,表达DbsCzcA基因的BL21(DE3)在培养过程中都可以生长,而空载体对照的生长一直受到抑制(见附图3)。实验结果说明,DbsCzcA对重金属镉的防御起着重要作用。
SEQ ID NO:1
MLKAILAFVLTRRPIILLGFAVFIGAGLVAVNKLNIEAYPNPAPVILEITAQAPGLSAEEMERYYTIPMEIGLYSALGIDNIRSTSFYGLSFVRVTFKYGVDFHFAYEQAALALQQNVTLPQNQIPQIQQSSLVGEVYRYQVVGPPHFGLTNLRSVQDWIILRRLSTIPGVVQVNSWGGTTKEFQVEADLNKLQAYNVTVQQLMAALGNGNINVGGREITIGQQSVNIRGIGLIDDGGADDLTKGYHVDDIEKIVLSQTNGVPVRVRDVAKVRVGF VPRLGIAGRDHDDDIAASIVVMSRVMHTNDIVPKIKAMVEQMNHDGSLPPGVHIEPYYDRLNLVSTTTHTVFHNLIFGCLLVFAIQWIFLGDLKSAIIVGINIPFALLFSIIILVLQGEDANLLSIRAVDFGIIIDSAVILVENIFRNLQAKKEERQRLLNDLSEGQMGHDPTTDRSVDSNVTWTDQLRLIFISSMQINRSVLFSVMITVAAFVPLFTMQGVEGQIFGPMARTYGYALAGALISSFTITPVLSSFLLRGNIQEVETFLVRHLHNLYIPILRWSIVNRKGVVLSGLLFLILSGILGTRLGSEFLPALEEGNFWIRASMPLTMGLDAGTEATRKMREILLRHPEVITVVSQHGRPDNGSDASAFSNVELFVPLKPYDQWPSGLTKEKLTQTLQQEFSDALPGVGFNFSQYIQDNVEEALSGVKGANSVKIIGHDLPTLERLAGQVMDQMNHVEGVADLGIFRVMGQPNLNITIDRDKAARYGLNTGDINTVIQATMGDAVASTVLEGDRQFNLTVRLPLQQRDNVEVIGNIKVGYTTPTGGTAFIPLRELATITLDSGASYIYHETTQRYIPIKF SVRGRDLGSTVSEAQKRIAENIQLPNGYRIIWAGEFEDLEKAKKRLAIIVPISLMLIAVPLYGLFNSARDSMLALLGIPFAVGGGILSLFITGIPFSVSAAIGFISLLGVSVMDRILIITYFNQLRMSGVSPIKALVEASEKRMRPLLMTALSACIGLFPAAISHGIGSQVQRPLATVVVGGLAIGSLLLLIVVPALHALLLPKEKRSS F G KAQMDEEDQ
SEQ ID NO:2
ATGCTCAAAGCCATTTTAGCCTTCGTGCTGACTCGACGACCGATTATCTTACTGGGGTTTGCTGTGTTTATCGGGGCTGGCCTTGTTGCTGTGAACAAACTCAACATCGAAGCCTATCCCAACCCGGCACCTGTTATTTTGGAAATTACCGCTCAGGCTCCGGGTCTCTCGGCAGAAGAAATGGAGCGTTACTATACGATTCCTATGGAAATCGGCCTGTATTCGGCGTTGGGTATCGACAATATTCGCTCGACTTCTTTTTATGGTTTGTCGTTTGTTCGTGTGACGTTCAAATATGGGGTTGATTTTCATTTTGCCTACGAGCAAGCTGCTTTGGCGCTCCAGCAAAATGTCACCTTGCCGCAAAATCAGATTCCTCAGATTCAGCAGTCCAGTCTGGTTGGTGAGGTATATCGTTATCAAGTGGTAGGGCCACCGCACTTTGGCTTAACAAATCTTCGCTCAGTACAAGACTGGATTATCCTGCGTCGCTTATCCACCATCCCCGGAGTGGTTCAAGTCAACAGTTGGGGGGGGACAACCAAGGAATTCCAAGTTGAGGCAGATCTCAACAAACTACAAGCCTACAATGTCACCGTACAACAGTTGATGGCGGCGCTGGGTAACGGCAATATCAACGTGGGTGGTCGCGAAATCACCATAGGCCAACAGTCGGTCAACATTCGCGGCATCGGCTTGATCGATGATGGAGGAGCGGATGACCTGACCAAAGGCTACCATGTTGATGACATCGAAAAGATTGTCCTGTCACAAACCAATGGGGTTCCAGTTCGAGTTCGGGATGTCGCTAAAGTCAGGGTTGGGTTTGTACCCCGACTGGGAATTGCAGGTCGTGATCATGACGATGACATTGCCGCATCCATCGTGGTCATGAGCCGCGTCATGCACACCAACGATATCGTGCCGAAGATCAAGGCTATGGTAGAGCAGATGAACCACGACGGCAGTTTGCCTCCAGGAGTTCATATTGAGCCTTACTACGATCGTTTAAATCTTGTGTCTACTACCACACATACCGTTTTCCATAATCTGATTTTCGGATGTCTGCTTGTTTTCGCAATTCAATGGATTTTTTTGGGAGATTTGAAGAGCGCTATCATTGTTGGTATTAACATTCCGTTTGCGCTCCTTTTCAGTATTATCATACTGGTGCTTCAAGGAGAGGATGCCAACCTTCTATCCATTAGAGCTGTTGACTTCGGTATCATCATCGATTCGGCTGTCATCTTGGTCGAGAATATATTCAGGAATCTACAAGCCAAGAAAGAAGAACGTCAGCGCCTGCTGAATGACCTTTCCGAAGGCCAAATGGGGCATGACCCCACGACCGATCGGTCGGTTGATAGCAACGTAACATGGACTGACCAGTTGCGACTTATCTTCATCAGTTCTATGCAAATCAATCGTTCCGTTCTTTTTTCCGTGATGATTACAGTGGCTGCATTTGTCCCCCTGTTTACTATGCAGGGGGTTGAGGGTCAGATTTTTGGCCCCATGGCGCGGACCTATGGTTATGCCCTGGCTGGTGCCTTGATTTCGAGTTTTACCATCACACCCGTACTGTCATCCTTTCTGTTGCGCGGCAATATTCAGGAAGTTGAAACATTTCTGGTGCGACATCTACATAACTTGTACATTCCGATTCTGCGCTGGTCCATTGTGAACCGTAAGGGGGTGGTACTGTCGGGACTGCTGTTTCTCATTCTGTCGGGCATTCTGGGGACGCGCTTGGGTTCCGAGTTTCTTCCTGCCTTGGAAGAGGGAAATTTCTGGATTCGCGCTTCAATGCCGCTCACAATGGGGCTGGATGCGGGCACAGAAGCCACGCGTAAAATGCGCGAAATTCTTCTCCGTCACCCTGAAGTTATTACGGTGGTTTCACAGCATGGTCGCCCAGATAATGGTAGCGATGCTTCGGCATTTTCCAACGTGGAACTTTTTGTGCCACTCAAACCTTATGATCAATGGCCCAGTGGTTTGACCAAGGAAAAATTGACCCAGACACTGCAACAAGAATTTTCAGATGCGTTACCCGGAGTGGGGTTCAATTTTTCACAATATATCCAGGATAATGTTGAAGAAGCTTTGTCAGGGGTTAAAGGGGCCAATTCAGTCAAAATCATTGGCCATGACTTGCCAACTCTGGAAAGATTGGCAGGCCAGGTCATGGATCAAATGAATCATGTTGAGGGGGTTGCTGATCTTGGTATTTTCCGGGTCATGGGGCAACCCAACCTCAACATTACCATCGATCGTGATAAGGCAGCGCGTTACGGGCTCAATACGGGTGATATCAATACGGTTATTCAGGCCACCATGGGAGATGCTGTGGCAAGTACGGTTCTGGAAGGTGATCGGCAATTCAACCTGACGGTTCGATTGCCGTTACAGCAACGAGATAATGTTGAAGTCATCGGCAATATCAAGGTTGGATACACAACACCGACCGGCGGAACGGCTTTTATCCCACTGCGTGAACTGGCAACGATCACCTTGGACAGTGGAGCATCCTACATCTATCACGAAACCACCCAACGCTATATTCCCATCAAGTTCAGCGTACGTGGACGCGATCTGGGAAGCACGGTAAGCGAAGCACAGAAAAGAATCGCAGAAAACATACAACTACCGAATGGTTACCGCATCATCTGGGCAGGTGAGTTTGAGGATCTTGAAAAAGCCAAAAAACGCTTGGCTATCATTGTTCCAATCAGTCTTATGCTGATCGCTGTTCCTCTTTACGGATTATTTAATTCCGCGCGTGATAGTATGCTTGCCCTACTGGGTATTCCTTTTGCTGTTGGCGGGGGCATTCTTTCGCTCTTTATCACGGGTATTCCGTTCAGTGTGTCAGCGGCCATCGGGTTTATCTCTCTTCTTGGGGTATCTGTGATGGATCGTATCCTGATCATTACTTATTTCAATCAGTTACGTATGTCAGGCGTGTCACCGATCAAGGCATTGGTCGAAGCCTCAGAAAAACGTATGCGCCCCTTGCTGATGACTGCACTATCGGCTTGTATCGGTTTATTCCCTGCTGCAATTTCACATGGTATCGGTAGTCAAGTGCAGCGCCCATTGGCGACCGTCGTGGTCGGCGGCTTGGCTATTGGTTCCCTGCTACTATTGATTGTTGTTCCGGCTTTGCATGCCTTGCTGCTTCCCAAAGAGAAGCGTTCCTCCTTCGGAAAAGCC C A G A T G G A T G A G G A A G A TCAGTGA。
SEQUENCE LISTING
<110> 中山大学
<120> 一种重金属镉抗性相关蛋白DbsCzcA及其编码基因和应用
<130>
<160> 4
<170> PatentIn version 3.2
<210> 1
<211> 1079
<212> PRT
<213> 人工序列
<400> 1
Met Leu Lys Ala Ile Leu Ala Phe Val Leu Thr Arg Arg Pro Ile Ile
1 5 10 15
Leu Leu Gly Phe Ala Val Phe Ile Gly Ala Gly Leu Val Ala Val Asn
20 25 30
Lys Leu Asn Ile Glu Ala Tyr Pro Asn Pro Ala Pro Val Ile Leu Glu
35 40 45
Ile Thr Ala Gln Ala Pro Gly Leu Ser Ala Glu Glu Met Glu Arg Tyr
50 55 60
Tyr Thr Ile Pro Met Glu Ile Gly Leu Tyr Ser Ala Leu Gly Ile Asp
65 70 75 80
Asn Ile Arg Ser Thr Ser Phe Tyr Gly Leu Ser Phe Val Arg Val Thr
85 90 95
Phe Lys Tyr Gly Val Asp Phe His Phe Ala Tyr Glu Gln Ala Ala Leu
100 105 110
Ala Leu Gln Gln Asn Val Thr Leu Pro Gln Asn Gln Ile Pro Gln Ile
115 120 125
Gln Gln Ser Ser Leu Val Gly Glu Val Tyr Arg Tyr Gln Val Val Gly
130 135 140
Pro Pro His Phe Gly Leu Thr Asn Leu Arg Ser Val Gln Asp Trp Ile
145 150 155 160
Ile Leu Arg Arg Leu Ser Thr Ile Pro Gly Val Val Gln Val Asn Ser
165 170 175
Trp Gly Gly Thr Thr Lys Glu Phe Gln Val Glu Ala Asp Leu Asn Lys
180 185 190
Leu Gln Ala Tyr Asn Val Thr Val Gln Gln Leu Met Ala Ala Leu Gly
195 200 205
Asn Gly Asn Ile Asn Val Gly Gly Arg Glu Ile Thr Ile Gly Gln Gln
210 215 220
Ser Val Asn Ile Arg Gly Ile Gly Leu Ile Asp Asp Gly Gly Ala Asp
225 230 235 240
Asp Leu Thr Lys Gly Tyr His Val Asp Asp Ile Glu Lys Ile Val Leu
245 250 255
Ser Gln Thr Asn Gly Val Pro Val Arg Val Arg Asp Val Ala Lys Val
260 265 270
Arg Val Gly Phe Val Pro Arg Leu Gly Ile Ala Gly Arg Asp His Asp
275 280 285
Asp Asp Ile Ala Ala Ser Ile Val Val Met Ser Arg Val Met His Thr
290 295 300
Asn Asp Ile Val Pro Lys Ile Lys Ala Met Val Glu Gln Met Asn His
305 310 315 320
Asp Gly Ser Leu Pro Pro Gly Val His Ile Glu Pro Tyr Tyr Asp Arg
325 330 335
Leu Asn Leu Val Ser Thr Thr Thr His Thr Val Phe His Asn Leu Ile
340 345 350
Phe Gly Cys Leu Leu Val Phe Ala Ile Gln Trp Ile Phe Leu Gly Asp
355 360 365
Leu Lys Ser Ala Ile Ile Val Gly Ile Asn Ile Pro Phe Ala Leu Leu
370 375 380
Phe Ser Ile Ile Ile Leu Val Leu Gln Gly Glu Asp Ala Asn Leu Leu
385 390 395 400
Ser Ile Arg Ala Val Asp Phe Gly Ile Ile Ile Asp Ser Ala Val Ile
405 410 415
Leu Val Glu Asn Ile Phe Arg Asn Leu Gln Ala Lys Lys Glu Glu Arg
420 425 430
Gln Arg Leu Leu Asn Asp Leu Ser Glu Gly Gln Met Gly His Asp Pro
435 440 445
Thr Thr Asp Arg Ser Val Asp Ser Asn Val Thr Trp Thr Asp Gln Leu
450 455 460
Arg Leu Ile Phe Ile Ser Ser Met Gln Ile Asn Arg Ser Val Leu Phe
465 470 475 480
Ser Val Met Ile Thr Val Ala Ala Phe Val Pro Leu Phe Thr Met Gln
485 490 495
Gly Val Glu Gly Gln Ile Phe Gly Pro Met Ala Arg Thr Tyr Gly Tyr
500 505 510
Ala Leu Ala Gly Ala Leu Ile Ser Ser Phe Thr Ile Thr Pro Val Leu
515 520 525
Ser Ser Phe Leu Leu Arg Gly Asn Ile Gln Glu Val Glu Thr Phe Leu
530 535 540
Val Arg His Leu His Asn Leu Tyr Ile Pro Ile Leu Arg Trp Ser Ile
545 550 555 560
Val Asn Arg Lys Gly Val Val Leu Ser Gly Leu Leu Phe Leu Ile Leu
565 570 575
Ser Gly Ile Leu Gly Thr Arg Leu Gly Ser Glu Phe Leu Pro Ala Leu
580 585 590
Glu Glu Gly Asn Phe Trp Ile Arg Ala Ser Met Pro Leu Thr Met Gly
595 600 605
Leu Asp Ala Gly Thr Glu Ala Thr Arg Lys Met Arg Glu Ile Leu Leu
610 615 620
Arg His Pro Glu Val Ile Thr Val Val Ser Gln His Gly Arg Pro Asp
625 630 635 640
Asn Gly Ser Asp Ala Ser Ala Phe Ser Asn Val Glu Leu Phe Val Pro
645 650 655
Leu Lys Pro Tyr Asp Gln Trp Pro Ser Gly Leu Thr Lys Glu Lys Leu
660 665 670
Thr Gln Thr Leu Gln Gln Glu Phe Ser Asp Ala Leu Pro Gly Val Gly
675 680 685
Phe Asn Phe Ser Gln Tyr Ile Gln Asp Asn Val Glu Glu Ala Leu Ser
690 695 700
Gly Val Lys Gly Ala Asn Ser Val Lys Ile Ile Gly His Asp Leu Pro
705 710 715 720
Thr Leu Glu Arg Leu Ala Gly Gln Val Met Asp Gln Met Asn His Val
725 730 735
Glu Gly Val Ala Asp Leu Gly Ile Phe Arg Val Met Gly Gln Pro Asn
740 745 750
Leu Asn Ile Thr Ile Asp Arg Asp Lys Ala Ala Arg Tyr Gly Leu Asn
755 760 765
Thr Gly Asp Ile Asn Thr Val Ile Gln Ala Thr Met Gly Asp Ala Val
770 775 780
Ala Ser Thr Val Leu Glu Gly Asp Arg Gln Phe Asn Leu Thr Val Arg
785 790 795 800
Leu Pro Leu Gln Gln Arg Asp Asn Val Glu Val Ile Gly Asn Ile Lys
805 810 815
Val Gly Tyr Thr Thr Pro Thr Gly Gly Thr Ala Phe Ile Pro Leu Arg
820 825 830
Glu Leu Ala Thr Ile Thr Leu Asp Ser Gly Ala Ser Tyr Ile Tyr His
835 840 845
Glu Thr Thr Gln Arg Tyr Ile Pro Ile Lys Phe Ser Val Arg Gly Arg
850 855 860
Asp Leu Gly Ser Thr Val Ser Glu Ala Gln Lys Arg Ile Ala Glu Asn
865 870 875 880
Ile Gln Leu Pro Asn Gly Tyr Arg Ile Ile Trp Ala Gly Glu Phe Glu
885 890 895
Asp Leu Glu Lys Ala Lys Lys Arg Leu Ala Ile Ile Val Pro Ile Ser
900 905 910
Leu Met Leu Ile Ala Val Pro Leu Tyr Gly Leu Phe Asn Ser Ala Arg
915 920 925
Asp Ser Met Leu Ala Leu Leu Gly Ile Pro Phe Ala Val Gly Gly Gly
930 935 940
Ile Leu Ser Leu Phe Ile Thr Gly Ile Pro Phe Ser Val Ser Ala Ala
945 950 955 960
Ile Gly Phe Ile Ser Leu Leu Gly Val Ser Val Met Asp Arg Ile Leu
965 970 975
Ile Ile Thr Tyr Phe Asn Gln Leu Arg Met Ser Gly Val Ser Pro Ile
980 985 990
Lys Ala Leu Val Glu Ala Ser Glu Lys Arg Met Arg Pro Leu Leu Met
995 1000 1005
Thr Ala Leu Ser Ala Cys Ile Gly Leu Phe Pro Ala Ala Ile Ser
1010 1015 1020
His Gly Ile Gly Ser Gln Val Gln Arg Pro Leu Ala Thr Val Val
1025 1030 1035
Val Gly Gly Leu Ala Ile Gly Ser Leu Leu Leu Leu Ile Val Val
1040 1045 1050
Pro Ala Leu His Ala Leu Leu Leu Pro Lys Glu Lys Arg Ser Ser
1055 1060 1065
Phe Gly Lys Ala Gln Met Asp Glu Glu Asp Gln
1070 1075
<210> 2
<211> 3240
<212> DNA
<213> 人工序列
<400> 2
atgctcaaag ccattttagc cttcgtgctg actcgacgac cgattatctt actggggttt 60
gctgtgttta tcggggctgg ccttgttgct gtgaacaaac tcaacatcga agcctatccc 120
aacccggcac ctgttatttt ggaaattacc gctcaggctc cgggtctctc ggcagaagaa 180
atggagcgtt actatacgat tcctatggaa atcggcctgt attcggcgtt gggtatcgac 240
aatattcgct cgacttcttt ttatggtttg tcgtttgttc gtgtgacgtt caaatatggg 300
gttgattttc attttgccta cgagcaagct gctttggcgc tccagcaaaa tgtcaccttg 360
ccgcaaaatc agattcctca gattcagcag tccagtctgg ttggtgaggt atatcgttat 420
caagtggtag ggccaccgca ctttggctta acaaatcttc gctcagtaca agactggatt 480
atcctgcgtc gcttatccac catccccgga gtggttcaag tcaacagttg gggggggaca 540
accaaggaat tccaagttga ggcagatctc aacaaactac aagcctacaa tgtcaccgta 600
caacagttga tggcggcgct gggtaacggc aatatcaacg tgggtggtcg cgaaatcacc 660
ataggccaac agtcggtcaa cattcgcggc atcggcttga tcgatgatgg aggagcggat 720
gacctgacca aaggctacca tgttgatgac atcgaaaaga ttgtcctgtc acaaaccaat 780
ggggttccag ttcgagttcg ggatgtcgct aaagtcaggg ttgggtttgt accccgactg 840
ggaattgcag gtcgtgatca tgacgatgac attgccgcat ccatcgtggt catgagccgc 900
gtcatgcaca ccaacgatat cgtgccgaag atcaaggcta tggtagagca gatgaaccac 960
gacggcagtt tgcctccagg agttcatatt gagccttact acgatcgttt aaatcttgtg 1020
tctactacca cacataccgt tttccataat ctgattttcg gatgtctgct tgttttcgca 1080
attcaatgga tttttttggg agatttgaag agcgctatca ttgttggtat taacattccg 1140
tttgcgctcc ttttcagtat tatcatactg gtgcttcaag gagaggatgc caaccttcta 1200
tccattagag ctgttgactt cggtatcatc atcgattcgg ctgtcatctt ggtcgagaat 1260
atattcagga atctacaagc caagaaagaa gaacgtcagc gcctgctgaa tgacctttcc 1320
gaaggccaaa tggggcatga ccccacgacc gatcggtcgg ttgatagcaa cgtaacatgg 1380
actgaccagt tgcgacttat cttcatcagt tctatgcaaa tcaatcgttc cgttcttttt 1440
tccgtgatga ttacagtggc tgcatttgtc cccctgttta ctatgcaggg ggttgagggt 1500
cagatttttg gccccatggc gcggacctat ggttatgccc tggctggtgc cttgatttcg 1560
agttttacca tcacacccgt actgtcatcc tttctgttgc gcggcaatat tcaggaagtt 1620
gaaacatttc tggtgcgaca tctacataac ttgtacattc cgattctgcg ctggtccatt 1680
gtgaaccgta agggggtggt actgtcggga ctgctgtttc tcattctgtc gggcattctg 1740
gggacgcgct tgggttccga gtttcttcct gccttggaag agggaaattt ctggattcgc 1800
gcttcaatgc cgctcacaat ggggctggat gcgggcacag aagccacgcg taaaatgcgc 1860
gaaattcttc tccgtcaccc tgaagttatt acggtggttt cacagcatgg tcgcccagat 1920
aatggtagcg atgcttcggc attttccaac gtggaacttt ttgtgccact caaaccttat 1980
gatcaatggc ccagtggttt gaccaaggaa aaattgaccc agacactgca acaagaattt 2040
tcagatgcgt tacccggagt ggggttcaat ttttcacaat atatccagga taatgttgaa 2100
gaagctttgt caggggttaa aggggccaat tcagtcaaaa tcattggcca tgacttgcca 2160
actctggaaa gattggcagg ccaggtcatg gatcaaatga atcatgttga gggggttgct 2220
gatcttggta ttttccgggt catggggcaa cccaacctca acattaccat cgatcgtgat 2280
aaggcagcgc gttacgggct caatacgggt gatatcaata cggttattca ggccaccatg 2340
ggagatgctg tggcaagtac ggttctggaa ggtgatcggc aattcaacct gacggttcga 2400
ttgccgttac agcaacgaga taatgttgaa gtcatcggca atatcaaggt tggatacaca 2460
acaccgaccg gcggaacggc ttttatccca ctgcgtgaac tggcaacgat caccttggac 2520
agtggagcat cctacatcta tcacgaaacc acccaacgct atattcccat caagttcagc 2580
gtacgtggac gcgatctggg aagcacggta agcgaagcac agaaaagaat cgcagaaaac 2640
atacaactac cgaatggtta ccgcatcatc tgggcaggtg agtttgagga tcttgaaaaa 2700
gccaaaaaac gcttggctat cattgttcca atcagtctta tgctgatcgc tgttcctctt 2760
tacggattat ttaattccgc gcgtgatagt atgcttgccc tactgggtat tccttttgct 2820
gttggcgggg gcattctttc gctctttatc acgggtattc cgttcagtgt gtcagcggcc 2880
atcgggttta tctctcttct tggggtatct gtgatggatc gtatcctgat cattacttat 2940
ttcaatcagt tacgtatgtc aggcgtgtca ccgatcaagg cattggtcga agcctcagaa 3000
aaacgtatgc gccccttgct gatgactgca ctatcggctt gtatcggttt attccctgct 3060
gcaatttcac atggtatcgg tagtcaagtg cagcgcccat tggcgaccgt cgtggtcggc 3120
ggcttggcta ttggttccct gctactattg attgttgttc cggctttgca tgccttgctg 3180
cttcccaaag agaagcgttc ctccttcgga aaagcccaga tggatgagga agatcagtga 3240
<210> 3
<211> 31
<212> DNA
<213> 人工序列
<400> 3
cgcggatcca tgctcaaagc cattttagcc t 31
<210> 4
<211> 31
<212> DNA
<213> 人工序列
<400> 4
ccgctcgagc tgatcttcct catccatctg g 31
Claims (5)
1.一种重金属镉抗性相关蛋白DbsCzcA,其特征在于氨基酸序列如SEQ ID NO:1所示。
2.权利要求1所述重金属镉抗性相关蛋白DbsCzcA的编码基因,其特征在于核苷酸序列如SEQ ID NO:2所示。
3.含有权利要求2所述重金属镉抗性相关蛋白DbsCzcA的编码基因的表达载体。
4.根据权利要求3所述载体,其特征在于,所述表达载体为pET28a-DbsCzcA。
5.一种基因工程菌,其特征在于是由权利要求4所述的表达载体转染大肠杆菌得到的。
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CN101153272A (zh) * | 2007-09-26 | 2008-04-02 | 南京农业大学 | 一种铅镉抗性细菌及其用于土壤重金属污染的植物修复方法 |
CN102020706A (zh) * | 2009-09-23 | 2011-04-20 | 中国科学院植物研究所 | 一种生物镉抗性相关蛋白及其编码基因 |
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CN1433676A (zh) * | 2002-01-21 | 2003-08-06 | 中山大学 | 总dna导入培育富集重金属转基因植物的方法 |
CN101153272A (zh) * | 2007-09-26 | 2008-04-02 | 南京农业大学 | 一种铅镉抗性细菌及其用于土壤重金属污染的植物修复方法 |
CN102020706A (zh) * | 2009-09-23 | 2011-04-20 | 中国科学院植物研究所 | 一种生物镉抗性相关蛋白及其编码基因 |
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