CN108440661B - 吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用 - Google Patents
吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用 Download PDFInfo
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- CN108440661B CN108440661B CN201810636598.1A CN201810636598A CN108440661B CN 108440661 B CN108440661 B CN 108440661B CN 201810636598 A CN201810636598 A CN 201810636598A CN 108440661 B CN108440661 B CN 108440661B
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Abstract
本发明公开了一种吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用。本发明从吊兰根部中克隆了一个硝酸盐转运蛋白的编码基因CcNPF8.3.1,并将该基因导入△ynt‑Leu双突变多形汉逊酵母中,得到转CcNPF8.3.1酵母。通过试验证明:CcNPF8.3.1蛋白具有硝酸盐转运蛋白的功能,能提高植物的氮素利用效率,使植物快速生长。
Description
技术领域
本发明涉及一种吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用。
背景技术
氮素是植物生长发育所必须的基本营养元素,在植物生长发育和形态建成中起着重要作用,其对作物的生命活动和产量形成也具有重要意义。硝酸盐(NO3 -)是大多数作物最主要的氮源,硝酸盐供应量不足会严重抑制作物的生长发育。生理学研究表明,植物从土壤中吸收NO3 -需有一整套高-和低-亲和力NO3-转运系统参加,NO3 -的进入由跨质膜的H+梯度驱动。有些NO3 -转运系统是组成型表达,有的则受NO3 -诱导,且随着NO3 -的同化呈负反馈调控。
吊兰属百合科多年生常绿草本植物。其根系在水中蔓延生长,生长迅速,根系发达,茂密浓郁。在适宜的环境条件下快速蓬勃生长。仅仅用两天能够长出一个侧根。
在过去几十年中,多形汉逊酵母(Hansenula polymorpha,又称Pichia angusta)已经成为一种公认的模式生物。广泛应用于研究甲醇代谢、硝酸盐吸收机制等。在多形汉逊酵母中,所有与硝酸盐代谢相关的基因紧密地排列成基因簇,总长1040bp的DNA片段中大约92%为编码DNA。多形汉逊酵母可以同化硝酸盐并把硝酸盐作为唯一氮源,它只有一个高亲和硝酸盐转运蛋白(nitrate transporters,Ynt 1),可以转运硝酸盐但是不能转运氯酸盐。YNT1结构上属于NNP(nitrate-nitrite porter)家族并且受到硝酸盐和亚硝酸盐的诱导。汉逊酵母基因YNT 1,YNR 1和YNI1分别编码硝酸盐转运蛋白、硝酸盐还原酶(nitratereductase),亚硝酸盐还原酶(nitrite reductase),它们的表达受硝酸盐和亚硝酸盐诱导并受按盐和谷氨酸抑致。YNT 1缺失突变(ynt1)可以导致菌株在硝酸盐浓度低于500μM的条件下不能转运或生长。
发明内容
本发明的目的是提供一种吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用。
本发明提供了一种蛋白质,克隆自吊兰,命名为CcNPF8.3.1,是如下(a1)或(a2):
(a1)由序列表中序列5所示的氨基酸序列组成的蛋白质;
(a2)将序列5的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有硝酸盐转运功能的由序列5衍生的蛋白质。
为了使(a1)中的CcNPF8.3.1蛋白便于纯化和检测,可在由序列表中序列5所示的氨基酸序列组成的蛋白质的氨基末端或羧基末端连接上如表1所示的标签。
表1标签的序列
标签 | 残基 | 序列 |
Poly-Arg | 5-6(通常为5个) | RRRRR |
Poly-His | 2-10(通常为6个) | HHHHHH |
FLAG | 8 | DYKDDDDK |
Strep-tag II | 8 | WSHPQFEK |
c-myc | 10 | EQKLISEEDL |
上述(a2)中的CcNPF8.3.1蛋白可人工合成,也可先合成其编码基因,再进行生物表达得到。上述(a2)中的CcNPF8.3.1蛋白的编码基因可通过将序列表中序列4所示的DNA序列中缺失一个或几个氨基酸残基的密码子,和/或进行一个或几个碱基对的错义突变,和/或在其5′端和/或3′端连上表1所示的标签的编码序列得到。
编码所述CcNPF8.3.1蛋白的基因(CcNPF8.3.1基因)也属于本发明的保护范围。
所述基因为如下(b1)-(b3)中任一所述的DNA分子:
(b1)序列表中序列4所示的DNA分子;
(b2)在严格条件下与(b1)限定的DNA序列杂交且编码具有硝酸盐转运功能的蛋白的DNA分子;
(b3)与(b1)或(b2)限定的DNA序列具有90%以上同源性且编码具有硝酸盐转运功能的蛋白的DNA分子。
上述严格条件可为用0.1×SSPE(或0.1×SSC),0.1%SDS的溶液,在DNA或者RNA杂交实验中65℃下杂交并洗膜。
含有CcNPF8.3.1基因的重组表达载体、表达盒、转基因细胞系或重组菌也属于本发明的保护范围。
所述重组表达载体具体可为将载体pYNR-LEU2的多克隆位点(具体可为salⅠ和speⅠ酶切位点)之间的片段替换为序列表的序列4所示的双链DNA分子得到的表达载体。
本发明还保护CcNPF8.3.1蛋白在转运硝酸盐中的应用。
本发明还保护CcNPF8.3.1蛋白或CcNPF8.3.1基因在调控生物硝酸盐吸收速率中的应用。
本发明还保护CcNPF8.3.1蛋白或CcNPF8.3.1基因在调控生物硝酸盐转运功能中的应用。
本发明还保护一种培育转基因生物的方法,是将CcNPF8.3.1基因导入目的生物中,得到转基因生物;所述转基因生物硝酸盐转运功能高于所述目的生物。
本发明还保护一种提高生物硝酸盐转运功能的方法,是增加目的生物中CcNPF8.3.1蛋白的表达量和/或活性,提高生物硝酸盐转运功能。
本发明还保护CcNPF8.3.1蛋白,或,CcNPF8.3.1基因,或,以上任一所述的方法,在生物育种中的应用。
所述生物育种的目的是为了选育硝酸盐转运功能高的生物。
以上任一所述生物具体可为植物,更具体可为吊兰。
以上任一所述生物具体可为酵母,更具体可为△ynt-Leu双突变多形汉逊酵母。
本发明从吊兰根部中克隆了一个硝酸盐转运蛋白的编码基因CcNPF8.3.1,并将该基因导入△ynt-Leu双突变多形汉逊酵母中,得到转CcNPF8.3.1酵母。通过试验证明:CcNPF8.3.1蛋白具有硝酸盐转运蛋白的功能,能提高植物的氮素利用效率,使植物快速生长。
附图说明
图1为硝酸盐转运蛋白基因鉴定图。
图2为3’RACE鉴定图。
图3为5’RACE鉴定图。
图4为转基因酵母△ynt-CcNPF8.3.1的鉴定图。
图5为CcNPF8.3.1蛋白功能验证。
图6为CcNPF8.3.1的硝酸盐吸收速率图。
具体实施方式
以下的实施例便于更好地理解本发明,但并不限定本发明。下述实施例中的实验方法,如无特殊说明,均为常规方法。下述实施例中所用的试验材料,如无特殊说明,均为自常规生化试剂商店购买得到的。以下实施例中的定量试验,均设置三次重复实验,结果取平均值。
吊兰:购自花鸟鱼市场。
△ynt-Leu双突变多形汉逊酵母:参考文献:Tobacco Nia2 cDNA functionallycomplements a Hansenulapolymorphayeast mutant lacking nitrate reductase.A newexpression system for thestudy of plant proteins involved in nitrateassimilation.;公众可以从中国农业科学院生物技术研究所获得。
载体pYNR-LEU2:参考文献:Tobacco Nia2 cDNA functionally complements aHansenulapolymorphayeast mutant lacking nitrate reductase.A new expressionsystem for thestudy of plant proteins involved in nitrate assimilation.;公众可以从中国农业科学院生物技术研究所获得。
野生型汉逊酵母(WT):参考文献:Tobacco Nia2 cDNA functionallycomplements a Hansenulapolymorphayeast mutant lacking nitrate reductase.A newexpression system for thestudy of plant proteins involved in nitrateassimilation.;公众可以从中国农业科学院生物技术研究所获得。
实施例1、CcNPF8.3.1蛋白及其编码基因的获得
一、硝酸盐转运蛋白基因克隆
1、使用TIANGEN公司的RNAprep Pure Plant试剂盒,参照试剂盒说明书提取吊兰根部的总RNA,反转录获得cDNA。反转录体系如表2所示。将反转录体系65℃,10min;在冰上急降至室温,加入1μl Rnase,42℃延伸2h,70℃,15min终止反应。
表2反转录体系
2、通过比较小粒碗藓、野生稻、拟南芥、水稻等硝酸盐转运蛋白的氨基酸序列,根据保守区设计了一对简并引物,由引物L5和引物L3组成;
引物L5:5′-ACNGAYGTNGARGARGTL-3′;
引物L3:5′-CATNCCYTTNGGRCAYTC-3′。
3、以步骤1得到的cDNA为模板,采用引物L5和引物L3进行PCR扩增,得到PCR扩增产物。PCR反应体系如表3所示。PCR反应程序为:95℃预5min;95℃变性30s;62℃-43℃退火30s(每个循环降低1℃),72℃延伸30s,95℃变性30s,42℃退火,72℃延伸30s,20cycle;72℃,10min。
表3 PCR反应体系
将PCR扩增产物进行1%琼脂糖凝胶电泳检测,并将PCR产物扩增连接到T载体进行测序,结果表明:PCR扩增得到一条大小为586bp的条带(即硝酸盐转运蛋白基因电泳结果见图1),其核苷酸序列如序列表中序列1所示。
二、3’RACE的获得
1、使用TIANGEN公司的RNAprep Pure Plant试剂盒,参照试剂盒说明书提取矮珍珠根部的总RNA,反转录获得cDNA。反转录体系如表4所示。反转录程序为:42℃,60min;70℃,15min。
表4反转录体系
2、以步骤1得到的cDNA模板,采用引物Outer(5′-CCGAGATCAGTAATTGTAGCAGC-3′)和引物Inner(5′-GCTCTGCACAGTCACTCAAATCGAAG-3′)使用TaKaRa LA Taq(CodeNo.RR002A)进行PCR反应,实验操作具体如下:
(1)Outer PCR反应
Outer PCR反应体系如表5所示。Outer PCR反应程序为:95℃预3min;95℃变性30s;55℃退火30s;72℃延伸30s,20个循环;72℃,10min。
表5 Outer PCR反应体系
(2)Inner PCR反应
Inner PCR反应体系如表6所示。Inner PCR反应程序为:95℃预3min;95℃变性30s;61℃退火30s;72℃延伸1min,30个循环;72℃,10min。
表6 Inner PCR反应体系
将PCR扩增产物进行1%琼脂糖凝胶电泳检测,并将PCR产物扩增连接到T载体进行测序,结果表明:PCR扩增得到一条大小为1057bp的条带(即3’RACE,电泳结果见图2),其核苷酸序列如序列表中序列2所示。
三、5’RACE的获得
1、使用TIANGEN公司的RNAprep Pure Plant试剂盒,参照试剂盒说明书提取矮珍珠根部的总RNA。
2、配置表7中所示的溶液。
表7溶液体系
3、将1μl步骤1得到的总RNA加至步骤2得到的体系中,混匀离心(14000×g)后,72℃反应3min,42℃反应2min,得到反应液。
4、完成步骤3后,按照表8配置溶液,同时向体系中加入1μl SmarterⅡA oligo–nucleotide,混匀离心(14000×g)后,42℃反应90min,70℃反应10min,得到反应液。
表8溶液体系
5、采用引物Outer(5′-CTTCGATTTGAGTGACTGTGCAGAGC-3′)使用TaKaRa LA Taq(Code No.RR002A)进行Outer PCR反应,反应体系见见表9。Outer PCR反应程序为:第一轮:95℃预3min;94℃变性30s;70℃退火30s;72℃延伸1min,5个循环;94℃变性30s;68℃退火30s;72℃延伸1min,5个循环;94℃变性30s;66℃退火30s;72℃延伸1min,5个循环;94℃变性30s;64℃退火30s;72℃延伸1min,20个循环;72℃,10min。第二轮:95℃预3min;94℃变性30s;68℃退火30s;72℃延伸1min,30个循环,72℃,10min。
表9 Outer PCR反应反应体系
将PCR扩增产物进行1%琼脂糖凝胶电泳检测,并将PCR产物扩增连接到T载体进行测序,结果表明:PCR扩增得到一条大小为850bp的条带(即5’RACE,电泳结果见图3),其核苷酸序列如序列表中序列3所示。
四、全基因克隆
根据上述步骤二和步骤三获得的3’RACE和5’RACE的核苷酸序列,设计了引物F8-3a-5(5′-gaattcATGGGCTCTCTCGGT-3′)和引物F8-3a-3(5′-tctagaGTAGGCTCTCTTGCTTTTAT-3′),以步骤一的1获得的cDNA为模板进行PCR扩增,得到PCR扩增产物。将其连接到pEasy-T1,挑单克隆,测序。
测序结果表明:PCR扩增得到了大小为1749bp的条带,其核苷酸序列如序列表中序列4所示,将序列4所示的基因命名为CcNPF8.3.1基因,CcNPF8.3.1基因编码的蛋白(命名为CcNPF8.3.1蛋白)的氨基酸序列如序列表中序列5所示。
实施例2、CcNPF8.3.1蛋白及其编码基因的功能验证
本实施例利用△ynt-Leu双突变多形汉逊酵母来验证CcNPF8.3.1蛋白及其编码基因的功能。
一、转基因酵母的构建
1、将载体pYNR-LEU2的salⅠ和speⅠ酶切位点之间的片段替换为序列表的序列4所示的双链DNA分子,得到穿梭质粒pYNR-CcNPF8.3.1。
2、将步骤1得到的穿梭质粒pYNR-CcNPF8.3.1用Bstp1酶切线性化,1%琼脂糖凝胶电泳,回收纯化目标产物,得到线性化片段。
3、制备转基因△ynt-Leu双突变多形汉逊酵母,具体步骤如下:
(1)挑取△ynt-Leu双突变多形汉逊酵母单菌落,接种至5mLYGNH液体培养基中,37℃、200r/min培养过夜。
YGNH液体培养基:0.17%(质量百分比)YNB,2%(质量百分比)葡萄糖,5mmol/LNH4Cl,溶剂为水。
(2)将100μL步骤(1)的培养物转接至100mL YGNH液体培养基中,37℃、200r/min培养14-16h,使菌液浓度达到OD600nm=1.3~1.5。
(3)将步骤(2)得到的菌液转接至预冷的50mL无菌用离心管中,4℃、1200r/min离心5min,收集菌体。
(4)将步骤(3)收集的菌体分别用50mL和25mL预冷的无菌水重悬各清洗一次。
(5)采用2mL冰预冷的1mol/L山梨醇溶液重悬步骤(4)得到的菌体,得到菌悬液(△ynt-LeuΔynt双突变多形汉逊酵母感受态细胞)。
(6)将50mg步骤2得到的线性化片段加至80μL步骤(5)得到的菌悬液中,混匀,冰浴5min。
(7)将步骤(6)得到的冰浴处理后的菌悬液150v、130Ω电击后加入600μl冰预冷的1mol/L山梨醇溶液将菌体混匀,37℃静置培养1h。
(8)完成步骤(7)后,将菌液涂布于含100mg/mL氨苄抗性的YNGH固体培养基上,37℃培养3~4天至有单克隆。
YNGH固体培养基:0.17%(质量百分比)YNB,2%(质量百分比)葡萄糖,NO3 -(0.1011g/200mL),溶剂为水。
(9)挑单菌落,以F8-1-5和F8-1-3为引物利用PCR筛选酵母阳性克隆。PCR反应程序为:95℃预15min;95℃变性30s;56℃退火30s;72℃延伸1min,30个循环;72℃,10min。PCR反应结束后,取5~10μl的PCR反应液进行1%的琼脂糖凝胶电泳检测,PCR扩增得到1749bp的阳性条带(电泳结果见图4,经测序,如序列表的序列4所示),结果表明转CcNPF8.3.1基因的酵母阳性株,将这类阳性菌株统一命名为△ynt-CcNPF8.3.1。
4、采用载体pYNR-LEU2替代穿梭质粒pYNR-CcNPF8.3.1,按照步骤2和步骤3进行操作,得到转空载体酵母。
二、功能验证
待测菌株:△ynt-Leu双突变多形汉逊酵母、野生型汉逊酵母(WT)、△ynt-CcNPF8.3.1和转空载体酵母。
挑取待测菌株单菌落,接种到10mL YNGH液体培养基中,37℃、200r/min培养12h,利用分光光度计测量OD600nm的吸光值。
结果如图5所示。结果表明,△ynt-Leu双突变多形汉逊酵母和转空载体酵母在YNGL培养基中不能生长,而野生型和转基因酵母△ynt-CcNPF8.3.1在YNGL培养基中可以正常生长,这表明CcNPF8.3.1基因使△ynt-Leu双突变多形汉逊酵母恢复生长,具有硝酸盐转运蛋白的功能。
三、吸收速率测定
硝酸根离子在紫外区有强烈的吸收,利用它在220nm波长处的吸光度可定量测定硝酸盐的浓度。虽然溶解在溶液中的有机物在220nm处也会有吸收,但硝酸根离子在275nm处没有吸收。因此,在275nm处作另一次测量,以校正硝酸盐氮值。A校=A220nm-A275nm。根据文献报道,通过测定硝酸盐转运蛋白的硝酸盐吸收效率Km来判定该蛋白是高亲和或低亲和转运蛋白。当Km<1000μM为高亲和转运蛋白;当Km>1000μM为低亲和转运蛋白。
(1)挑取△ynt-CcNPF8.3.1单菌落,接种至5mL YGNH液体培养基中,37℃、200r/min培养12h。
(2)将100μl步骤(1)得到的培养物接种至100mL YGNH液体培养基中,37℃、200r/min培养14-16h,使菌液浓度达到OD600nm=1.3~1.5,4℃、1200r/min离心5min,收集菌体。
(3)取6个装有10mL YG液体培养基的50mL的离心管,每管中分别加入100mg步骤(2)得到的菌体,37℃、200r/min震荡培养2h,以使每管中的酵母处于相同的生长状态。
(4)完成步骤(3)后,4℃、1200r/min离心5min,弃上清,将各管的酵母菌体沉淀分别用50mL冰冷的含有50μM、100μM、300μM、500μM、1000μM和1500μM硝酸盐的YG液体培养基重悬,冲洗一遍。
(5)重复步骤(4)两次。
(6)完成步骤(5)后,分别向各管加入10mL含有相应硝酸盐浓度(50μM、100μM、300μM、500μM、1000μM和1500μM)的YG液体培养基,37℃、200r/min振荡培养30分钟后测定吸光度,绘制吸收速率曲线,计算Km值。
结果如图6所示。通过计算,CcNPF8.3.1蛋白的的Km值为1100μM,大于1000μM,表明CcNPF8.3.1蛋白是一个低亲和的硝酸盐转运蛋白。
序列表
<110> 中国农业科学院生物技术研究所
<120> 吊兰根系硝酸盐转运蛋白CcNPF8.3.1及其编码基因与应用
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gcttctggcg ggattgagtg ccctgaacct tatggtttac gtctgttgtg ctatgaggta 720
taaaagcaag agagcctact aattgtaagt agtattgtat ggtttatgaa gttgtatgtg 780
gttacgtttt gaaagttatg gaattcgtat tcaaagttca tttcaaaaaa aaaaacctat 840
agtgaaatca ctagtggagg atccgcgaaa gggcaattcc agcacactgg cggccgttac 900
tagtggatcc gagctcggta ccaagcttgg cgtaatcatg gtcatagctg tttcctgtgt 960
gaaattgtta tccgctcaca attccacaca acatacgagc cggaagcata aagtgtaagc 1020
ctgggggtgc ctaatgagtg aagctaactc acattaa 1057
<210> 3
<211> 850
<212> DNA
<213> 吊兰(Chlorophytum comosum(Thunb.)Baker.)
<400> 3
ctaatacgac tcactatagg gcaagcagtg gtatcaacgc agagtacatg ggggagggaa 60
cccatctcgt tgccaagtca aacaacgacc catgggctct ctcggtgagg aacaggatgg 120
gcgtgcgctc atggaagaag cgcttttgga gaacaaaagt ggaggaccat ataccggtga 180
tgggtctgtc gactaccatg gaaatccagt tttgaagatt cacagtggca actggaaagc 240
ttgtgcattc attctaggta ccgaatgctg cgaacgtttg gcctattttg gaatagcaac 300
aaatcttgtt tcttatctca ctaaagagtt gcacaaagga aatgttgaag ctgcaagaac 360
tgttacgact tggtccggaa cttgctattt gacaactctc attggagctg ttattgcgga 420
tgcatcttgg ggaagatatt ggaccattgc agttttttca acaatatacc tcattggaat 480
gagtgcatta actctttcgg cttcctttcc agctctccag cctccaaaat gtgcaggatc 540
tatttgccca gaagcaagcc cagttcaaca ttgggcactg tatattggcc tgtatctcgt 600
tgccttagga acaggtggaa tcaaaccatg tgttccctcc tttggggccg atcagtttga 660
tgacacagat gcatcagaga gagtgagaaa gggatcattt tttaactggt tcttcatgtg 720
catatacatt ggttccttcg tttctagtag ttttgttgtg tgggtgcaag acaattgcgg 780
ttggggtata ggattcggaa taccaacatt ttttatggct ctcgccattg gaagcttctt 840
tgctgggacc 850
<210> 4
<211> 1749
<212> DNA
<213> 吊兰(Chlorophytum comosum(Thunb.)Baker.)
<400> 4
atgggctctc tcggtgagga acaggatggg cgtgcgctca tggaagaagc gcttttggag 60
aacaaaagtg gaggaccata taccggtgat gggtctgtcg actaccatgg aaatccagtt 120
ttgaagattc acagtggcaa ctggaaagct tgtgcattca ttctaggtac cgaatgctgc 180
gaacgtttgg cctattttgg aatagcaaca aatcttgttt cttatctcac taaagagttg 240
cacaaaggaa atgttgaagc tgcaagaact gttacgactt ggtccggaac ttgctatttg 300
acaactctca ttggagctgt tattgcggat gcatcttggg gaaaatattg gaccattgca 360
gttttttcaa caatatacct cattggaatg agtgcattaa ctctttcggc ttcctttcca 420
gctctccagc ctccaaaatg tgcaggatct atttgcccag aagcaagccc agttcaacat 480
tgggcactgt atattggcct gtatctcgtt gccttaggaa caggtggaat caaaccatgt 540
gttccctcct ttggggccga tcagtttgat gacacagatg catcagagag agtgagaaag 600
ggatcatttt ttaactggtt cttcatgtgc atatacattg gttccttcgt ttctagtagt 660
tttgttgtgt gggtgcaaga caattgcggt tggggtatag gattcggaat accaacattt 720
tttatggctc tcgccattgg aagcttcttc gctgggaccc cgctttatag gtttcagaag 780
cctgggggaa gccctgttac ccgagtttgc caggttgttg ttgcatcatt tcgaaagtgg 840
aaggtgtgtg tccctgctga cagttctcta ctatatgaac ttcctggcaa tggttcagct 900
atcactggaa gtaggaaatt agtacacatc aatgacctca gattcttcga taaagctgct 960
acaattactg atctcgacga ggcaggaaat gtttccaacc cttggaagct ctgcacagtc 1020
actcaaatcg aagagctaaa gacactaata agaatgttcc ccgtctgggc caccatcatc 1080
atgtttgcag tcgtttacgc ccagatttca accacattcg tcgaacaagg catggtcctc 1140
aatacaaccg taggctcctt caccatcccc ccggcttctc tttcaacagt catcgttatc 1200
agtgtcattc tcccggttcc actgtacgat acatttttcg tgccaacagc agcaaggttc 1260
acaggcaatg aaagaggcat cagccaactc caaaggatag gcatcggcct gttcatatca 1320
gtactcgcca tggtagctgc agcattgctg gagaggaaga ggttgaagac ctcaaagtca 1380
aatgcaggcg atgtagacca gatgagtatc ttgtggcaat tcccacagta ttttctagtc 1440
ggagcagcag aggtttttac gttcattggc cagtcagagc ttttctatga gtatgctcca 1500
gatgccatga ggagcctctg cacttctctg gcgctgatca cgactggttt cggaaactac 1560
ctgagttcct tcatcttggc tgcagtgacg tacttgactg cacgtggagg agaacctgga 1620
tgggttcctg atgatctcaa caaagggcat ttagactatc tcttctggct tctggcggga 1680
ttgagtgccc tgaaccttat ggtttacgtc tgttgtgcta tgaggtataa aagcaagaga 1740
gcctactaa 1749
<210> 5
<211> 582
<212> PRT
<213> 吊兰(Chlorophytum comosum(Thunb.)Baker.)
<400> 5
Met Gly Ser Leu Gly Glu Glu Gln Asp Gly Arg Ala Leu Met Glu Glu
1 5 10 15
Ala Leu Leu Glu Asn Lys Ser Gly Gly Pro Tyr Thr Gly Asp Gly Ser
20 25 30
Val Asp Tyr His Gly Asn Pro Val Leu Lys Ile His Ser Gly Asn Trp
35 40 45
Lys Ala Cys Ala Phe Ile Leu Gly Thr Glu Cys Cys Glu Arg Leu Ala
50 55 60
Tyr Phe Gly Ile Ala Thr Asn Leu Val Ser Tyr Leu Thr Lys Glu Leu
65 70 75 80
His Lys Gly Asn Val Glu Ala Ala Arg Thr Val Thr Thr Trp Ser Gly
85 90 95
Thr Cys Tyr Leu Thr Thr Leu Ile Gly Ala Val Ile Ala Asp Ala Ser
100 105 110
Trp Gly Lys Tyr Trp Thr Ile Ala Val Phe Ser Thr Ile Tyr Leu Ile
115 120 125
Gly Met Ser Ala Leu Thr Leu Ser Ala Ser Phe Pro Ala Leu Gln Pro
130 135 140
Pro Lys Cys Ala Gly Ser Ile Cys Pro Glu Ala Ser Pro Val Gln His
145 150 155 160
Trp Ala Leu Tyr Ile Gly Leu Tyr Leu Val Ala Leu Gly Thr Gly Gly
165 170 175
Ile Lys Pro Cys Val Pro Ser Phe Gly Ala Asp Gln Phe Asp Asp Thr
180 185 190
Asp Ala Ser Glu Arg Val Arg Lys Gly Ser Phe Phe Asn Trp Phe Phe
195 200 205
Met Cys Ile Tyr Ile Gly Ser Phe Val Ser Ser Ser Phe Val Val Trp
210 215 220
Val Gln Asp Asn Cys Gly Trp Gly Ile Gly Phe Gly Ile Pro Thr Phe
225 230 235 240
Phe Met Ala Leu Ala Ile Gly Ser Phe Phe Ala Gly Thr Pro Leu Tyr
245 250 255
Arg Phe Gln Lys Pro Gly Gly Gly Pro Val Thr Arg Val Cys Gln Val
260 265 270
Val Val Ala Ser Phe Arg Lys Trp Lys Val Cys Val Pro Ala Asp Ser
275 280 285
Ser Leu Leu Tyr Glu Leu Pro Gly Asn Gly Ser Ala Ile Thr Gly Ser
290 295 300
Arg Lys Leu Val His Ile Asn Asp Leu Arg Phe Phe Asp Lys Ala Ala
305 310 315 320
Thr Ile Thr Asp Leu Asp Glu Ala Glu Asn Val Ser Asn Pro Trp Lys
325 330 335
Leu Cys Thr Val Thr Gln Ile Glu Glu Leu Lys Thr Leu Ile Arg Met
340 345 350
Phe Pro Val Trp Ala Thr Ile Ile Met Phe Ala Val Val Tyr Ala Gln
355 360 365
Ile Ser Thr Thr Phe Val Glu Gln Gly Met Val Leu Asn Thr Thr Val
370 375 380
Gly Ser Phe Thr Ile Pro Pro Ala Ser Leu Ser Thr Val Ile Val Ile
385 390 395 400
Ser Val Ile Leu Leu Val Pro Leu Tyr Asp Thr Phe Phe Val Pro Thr
405 410 415
Ala Ala Arg Phe Thr Gly Asn Glu Arg Gly Ile Ser Gln Leu Gln Arg
420 425 430
Ile Gly Ile Gly Leu Phe Ile Ser Val Leu Ala Met Val Ser Ala Ala
435 440 445
Leu Leu Glu Arg Lys Arg Leu Lys Thr Ser Lys Ser Asn Ala Gly Asp
450 455 460
Val Asp Gln Met Ser Ile Leu Trp Gln Phe Pro Gln Tyr Phe Leu Val
465 470 475 480
Gly Ala Ala Glu Val Phe Thr Phe Ile Gly Gln Ser Glu Phe Phe Tyr
485 490 495
Glu His Ala Pro Asp Ala Met Arg Ser Leu Cys Thr Ser Leu Ala Leu
500 505 510
Ile Thr Thr Gly Phe Gly Asn Tyr Leu Ser Ser Phe Ile Leu Ala Ala
515 520 525
Val Thr Tyr Leu Thr Ala Arg Gly Gly Glu Pro Gly Trp Val Pro Asp
530 535 540
Asp Leu Asn Lys Gly His Leu Asp Tyr Leu Phe Trp Leu Leu Ala Gly
545 550 555 560
Leu Ser Ala Leu Asn Leu Met Val Tyr Val Cys Cys Ala Met Arg Tyr
565 570 575
Lys Ser Lys Arg Ala Tyr
580
Claims (10)
1.一种蛋白质,是由序列表中序列5所示的氨基酸序列组成的蛋白质。
2.编码权利要求1所述蛋白质的基因。
3.如权利要求2所述的基因,其特征在于:所述基因为序列表中序列4所示的DNA分子。
4.含有权利要求2或3所述基因的重组表达载体、表达盒或重组菌。
5.权利要求1所述蛋白质在酵母转运硝酸盐中的应用。
6.权利要求1所述蛋白质,或,权利要求2或3所述基因,在调控酵母硝酸盐吸收速率中的应用。
7.权利要求1所述蛋白质,或,权利要求2或3所述基因,在调控酵母硝酸盐转运功能中的应用。
8.一种培育转基因酵母的方法,是将权利要求2或3所述基因导入目的酵母中,得到转基因酵母;所述转基因酵母硝酸盐转运功能高于所述目的酵母。
9.一种提高酵母硝酸盐转运功能的方法,是增加目的酵母中权利要求1所述蛋白质的表达量,提高酵母硝酸盐转运功能。
10.权利要求1所述的蛋白质,或,权利要求2或3所示的基因,或,权利要求8或9所述的方法,在酵母育种中的应用。
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