CN110423765A - 一种番茄红素ε-环化酶基因及其编码蛋白和应用 - Google Patents
一种番茄红素ε-环化酶基因及其编码蛋白和应用 Download PDFInfo
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Abstract
一种番茄红素ε‑环化酶基因及其编码蛋白和应用，它涉及一种PyLCYE蛋白及其编码基因在紫菜类胡萝卜素代谢中的应用。本发明的基因的核苷酸序列如序列表Seq ID No：1所示或与序列表SEQ ID NO:1核苷酸序列具有80％以上同源性且编码相同功能蛋白质的多核苷酸序列。它的氨基酸序列如序列表Seq ID No：2所示或由序列表Seq ID No：2所述的氨基酸序列经过一个或者几个氨基酸残基的取代、缺失或者添加且具有与序列表Seq ID No：2所述的氨基酸残基序列相同活性的氨基酸序列。它用于紫菜类胡萝卜素代谢基因工程中。
Description
技术领域
本发明属于基因工程技术领域，具体涉及一种条斑紫菜番茄红素ε-环化酶PyLCYE蛋白及其编码基因在紫菜类胡萝卜素和叶绿素代谢中的应用。
背景技术
紫菜是世界上重要的栽培海藻之一，主要产于中国、日本和韩国，据统计，2010-2011年度三个国家紫菜总产值约88亿元(Blouin et al.2011)。紫菜产业已成为沿海地区经济创收、出口创汇的重要支柱产业，也是吸引、转移传统捕捞行业从业人员最多的产业，对沿海地区渔民增收、经济发展和社会稳定起着不可或缺的作用。不仅如此，紫菜还是研究植物生理、生化以及早期进化的理想材料(Waaland et al.2004；Lin et al.2009；Su etal.2010)。
紫菜生长在潮间带，必须耐受由潮汐带来的日周期性的极端逆境(如高光、高盐、失水等)，长期的进化过程使其产生了在逆境下对光合***的保护机制。失水、高光和低温等逆境的最终特征都是在植物细胞内产生活性氧质(ROS，reactive oxygen species)，对生物造成损伤(Jithesh et al.2006)。研究表明，环化类胡萝卜素(cyclic carotenoids)是高等植物防御ROS损伤的第一道防线。研究表明，有氧光合生物中合成环化类胡萝卜素受阻可能是致死的，表明了环化类胡萝卜素在植物应对环境变化中的重要作用(Britton etal.1979)。
类胡萝卜素(carotenoids)在植物体内通常积累在花、果实和根部，鲜艳的颜色可以吸引昆虫或动物作为传粉或者散播种子的媒介。此外，类胡萝卜素作为维生素A的前体，对人类和动物有很大的营养价值(Cuningham and Gantt 1998；Shumskaya and Wurtzel2013)。而类胡萝卜素中的叶黄素(lutein)和玉米黄素(zeaxanthin)存在于人类和动物眼中的黄斑区域，对人类视觉感受光线变化有十分重要的作用(Handelman et al.1998)。根据其结构特征，类胡萝卜素可以分为线性类胡萝卜素和环化类胡萝卜素两大类。针对高等植物的研究表明，类胡萝卜素的代谢起始于八氢番茄红素(phytoene)的合成，经过相关代谢酶的一系列脱氢和异构，形成线性的全反式番茄红素(lycopene)。以番茄红素为底物，番茄红素环化酶(lycopene cyclase)负责类胡萝卜素的环化，并且不同的环化酶在线性的番茄红素两端产生不同的环，从而在类胡萝卜素代谢途径中首次出现代谢分支。番茄红素β-环化酶可以在线性的番茄红素两个末端催化形成两个β-环，生成β-胡萝卜素(β-carotene)。经过番茄红素β-环化酶和番茄红素ε-环化酶的依次作用，分别在线性番茄红素的末端形成一个β-环和一个ε-环，生成α-胡萝卜素(α-carotene)。在少数植物种类中(如莴苣、生菜)，其番茄红素ε-环化酶具有可以在线性番茄红素每个末端形成一个ε-环，生成ε-胡萝卜素(ε-carotene)。大多数植物的番茄红素ε-环化酶仅可以可以在线性番茄红素一端形成ε-环，生成δ-胡萝卜素。研究表明，α-和β-类胡萝卜素同时参与植物的逆境适应，并且在应对环境变化中有着截然不同却又相互补充的作用(Dall'Osto et al.2007)。因此，精确调控α-/β-类胡萝卜素的比例是植物应对不断变化环境重要适性反应(Krause etal.2004)。因此，在类胡萝卜素的代谢中，不同番茄红素环化酶的功能活性及其表达的变化可以控制类胡萝卜素代谢流向不同分支的分配，从而使植物适应不同的环境胁迫。
申请者前期的研究表明，紫菜中可以代谢产生并积累两类环化类胡萝卜素：α-类胡萝卜素(α-caroteniods)和β-类胡萝卜素(β-carotenoids)，具体来说，主要包括α-胡萝卜素、叶黄素(lutein)、β-胡萝卜素和玉米黄素(zeaxanthin)及几种微量的代谢中间产物(Yang et al.2014)。然而，目前紫菜类胡萝卜素代谢途径的相关研究还比较少，仅有申请者对脐形紫菜中胡萝卜素羟化酶(PuCHY1)和牻牛儿基牻牛儿基二磷酸合酶(PuGGPS)的报道，尚未有任何关于番茄红素ε-环化酶的报道。对以条斑紫菜为代表的紫菜物种中PyLCYE的发现和功能鉴定有利于未来利用基因工程技术提高紫菜中类胡萝卜素等营养成分的含量来提高紫菜的营养价值，并且通过提高萜类物质的含量改善紫菜的口感和风味，从而整体上提高紫菜的经济价值。
发明内容
本发明提供了一种番茄红素ε-环化酶基因及其编码蛋白和应用，该基因转录组编码番茄红素ε-环化酶，从而使紫菜中能够合成并积累α-胡萝卜素和叶黄素等类胡萝卜素。
本发明的一种番茄红素ε-环化酶基因，它的核苷酸序列如序列表Seq ID No：1所示。
本发明的一种番茄红素ε-环化酶基因，它的核苷酸序列为与序列表SEQ ID NO:1核苷酸序列具有80％以上同源性且编码相同功能蛋白质的多核苷酸序列。
本发明的一种番茄红素ε-环化酶基因编码的蛋白，它的氨基酸序列如序列表SeqID No：2所示。
本发明的一种番茄红素ε-环化酶基因编码的蛋白，它的氨基酸序列为由序列表Seq ID No：2所述的氨基酸序列经过一个或者几个氨基酸残基的取代、缺失或者添加且具有与序列表Seq ID No：2所述的氨基酸残基序列相同活性的氨基酸序列。
本发明的番茄红素ε-环化酶基因来源于条斑紫菜(Pyropia yezoensis)，名称为PyLCYE，以下该基因简称为PyLCYE。
所述的番茄红素ε-环化酶来源条斑紫菜。
一种重组表达载体，它包括权利要求1或2所述的基因。
一种转化子，它包括含有权利要求6所述的重组表达载体的宿主细胞。
所述的一种转化子，它的宿主为微生物、植物或转基因细胞系。
本发明的一种番茄红素ε-环化酶基因的应用，它用于紫菜类胡萝卜素代谢基因工程中。
本发明的番茄红素ε-环化酶基因(PyLCYE)获取方式为：
一、总RNA的提取
选取条斑紫菜(Pyropia yezoensis)(来自江苏省海洋水产研究所国家级紫菜种质库)，采用RNAiso试剂盒(Takara公司)提取得到条斑紫菜总RNA，经电泳检测和紫外分光光度计检测确认RNA的完整性、纯度以及浓度后，于-80℃保存；
二、条斑紫菜番茄红素ε-环化酶基因(PyLCYE)的克隆
用1μg步骤一获得的总RNA作为反转录的模板，按照SMARTer RACE cDNA Kit(Clontech)的说明书操作，使用SSRT-II反转录酶，经反转录合成cDNA第一链后，用Touchdown PCR程序进行巢式PCR扩增，以获得PyLCYE ORF全长。
其中
5'-RACE所用引物如下：
PyLCY2-ER1：ACCCGCAGGTAGGGGCGGTCGAGCACAA
PyLCY2-ER2：CGCGGTCACGTTTGCCCGTTCAATGACCAC
3'-RACE所用引物如下：
PyLCY2-HF1：CCTCCAGGCCTCTAACCGCATCCGGGGG
PyLCY2-HF2：TGATCCGCTCGGTGCTGCCCCTGTGGCTG
将PCR产物连接至pMD19-T(Takara公司)载体，测序后经拼接获得具有完整编码区的条斑紫菜番茄红素ε-环化酶基因PyLCYE的ORF序列Seq ID No：1。
三、大肠杆菌表达载体的构建
用TargetP和ChloroP等软件推测PyLCYE蛋白中信号肽的位点，然后用带有EcoR I酶切位点的引物扩增PyLCYE截除信号肽后部分序列并克隆至pMAL-C5X载体。
带有EcoR I酶切位点的引物序列如下：
PyLCY2-pMAL-HF：
ccgcgatatcgtcgacggatccGCCACCAGCCGGTCGGCGGGGGC
PyLCY2-pMAL-HF：
tacctgcagggaattcggatccCTACTGCTGGTTCTCCCCCCGCTG
其中，小写字母代表的同源臂序列，大写字母是作为引物的基因序列，下划线部分为引入的酶切位点。
四、酶活鉴定
将步骤三获得的表达载体pMAL-PyLCYE与携带有一系列噬夏孢欧文氏菌类胡萝卜素合成相关基因CrtE(GGPS)、CrtB(PSY)和CrtI(PDS/ZDS)的载体pAC-LYC共转入大肠杆菌。载体pAC-LYC上携带的基因在有番茄红素ε-环化酶活性的情况下，可以使大肠杆菌的细胞积累ε-胡萝卜素或其中间产物δ-胡萝卜素(单ε-环化)，从而使大肠杆菌菌落或菌液呈橙黄色；若没有番茄红素ε-环化酶活性，大肠杆菌为橙红色。空载体pMAL-C5X与pAC-LYC共转作为负对照。HPLC分析各个样品的产物。
本发明的PyLCYE基因来源于脐形紫菜，其基因工程受体植物适用于水稻等生物。
利用本法发明的PyLCYE基因作为目的基因构建植物表达载体，可以利用花椰菜花叶病毒CAMV35S启动子、乙醇诱导启动子等，必要时可以包括增强子。为了简化转化植株的鉴定，可使用选择性标记(如抗生素酶)。所用的表达载体可使用Ti质粒、Ri质粒以及植物病毒载体等。转化方法可用农杆菌介导法或其他方法转化植物。
本发明首次发现紫菜属物种里参与类胡萝卜素合成代谢途径里的一个酶即番茄红素ε-环化酶，并证实该酶参与紫菜中类胡萝卜素的代谢。
本发明获得编码该酶的基因序列，为利用基因工程技术改进紫菜里类胡萝卜素代谢途径，以提高紫菜里类胡萝卜素和萜类物质的含量提供了基础。
通过本发明提供的PyLCYE基因和蛋白，对紫菜属物种进行基因工程改进，其产业价值着重于两个方面，一是通过提高紫菜中类胡萝卜素的含量从而提高紫菜的营养价值，从而提高紫菜的经济价值。
附图说明
图1为PyLCYE的核苷酸和推定氨基酸序列；其中，“*”代表终止密码子；
图2为不同物种番茄红素环化酶蛋白序列的多序列比对，标记区域为保守的罗斯曼折叠(1),两个环化酶结构域(2,3),两个跨膜螺旋结构域(4,6)和带点结构域(5)，方框内区域只存在于红藻番茄红素环化酶序列中；
图3为PyLCYE在大肠杆菌中实验的HPLC分析图；其中，A：不同类胡萝卜素的吸收光谱；B：pMAL-C5X和pAC-LYC共转进大肠杆菌作为负对照；C：pMAL-PyLCYE和pAC-LYC共转进大肠杆菌；
图4为不同物种中番茄红素环化酶的***发生分析图。
具体实施方式
本发明内容不仅限于上述各实施例的内容，含有本发明基因的重组表达载体、转基因细胞系及宿主菌均属本发明的保护范围。
实施例1
本实施例的PyLCYE的基因获得过程如下：
1)条斑紫菜PyLCYE转录本鉴定和编码区克隆
用拟南芥中编码LCYE基因的开放阅读框(ORF)，搜索条斑紫菜基因组中的同源序列，搜索方法为tblastx，截取e-value值为1×10-10已筛选可能的同源序列，只发现一个转录本contig_25259。根据该转录本基因序列设计RACE引物扩增PyLCYE的ORF(图1所示)。
选用RNAiso试剂(Takara公司)提取条斑紫菜叶状体总RNA，经电泳检测和紫外分光光度计检测确认RNA的完整性、纯度以及浓度，于-80℃保存。用1μg总RNA作为反转录的模板，经反转录合成cDNA第一链后，进行PCR扩增，反应结束后将PCR产物连接至pMAL-C5X载体，该基因全长1830bp，共编码含有609个氨基酸残基的蛋白序列和一个终止密码子，经分析，在该蛋白的N端有一段叶绿体定位的信号肽。将所推测的蛋白序列与其他物种中的同源番茄红素环化酶多序列对比(图2所示)，说明所获得的PyLCYE与其他物种中的番茄红素环化酶同源。
2)条斑紫菜PyLCYE表达载体的构建
用1μg步骤1)获得的总RNA作为反转录模板，使用Takara公司的“PrimeScriptTM1st Strand cDNA Synthesis Kit”试剂盒反转合成cDNA第一链。
根据RACE获得的PyLCYE的全长编码序列，用ChloroP和TargetP推测其编码蛋白的信号肽序列，设计两端引物并引入EcoR I限制性内切酶位点，扩增PyLCYE截除信号肽后的部分序列(引物由Genscript公司合成)。
用1μg步骤2)获得的总RNA作为反转录的模板，经反转录合成cDNA第一链后，进行PCR扩增，PCR程序如下：94℃预变性2min；94℃变性30s，67℃退火1min，72℃延伸1min，35个循环后，72℃10min。反应结束后将PCR产物连接至pMAL-C5X载体，连接产物转化大肠杆菌BL21(DE3)感受态细胞后筛选阳性克隆，得到携带PyLCYE的重组载体，命名为pMAL-PyLCYE。
3)PyLCYE的异源表达和功能鉴定
用步骤2)构建好的含有PyLCYE的载体转化入大肠杆菌BL21(DE3)细胞中，在含有羧苄霉素(50μg·mL-1)的LB-Agar培养基上培养过夜筛选，挑单菌落转入含有相同浓度羧苄霉素的LB液体培养基上37℃，200rpm振荡培养至OD600为0.4～0.6，加入异丙基-β-D-硫代半乳糖苷(IPTG，0.5mmol·L-1)诱导表达3～4h。表达的蛋白经SDS-聚丙烯酰胺凝胶电泳(PAGE，10％)分析。
实施例2
PyLCYE的功能验证如下：
为了鉴定PyLCYE的功能，我们采用的是大肠杆菌体内酶活鉴定***，用颜色互补的方法结合HPLC分析酶的活性。该***用到质粒pAC-LYC，该质粒携带噬夏孢欧文氏菌类胡萝卜素合成相关基因牻牛儿基牻牛儿基二磷酸合酶(CrtE，GGPS)、八氢番茄红素合酶(CrtB，PSY)、八氢番茄红素去饱和酶(CrtI，PDS/ZDS)，可以在大肠杆菌细胞里积累番茄红素(橙红色)，作为番茄红素环化酶的底物。在有番茄红素ε-环化酶的情况下可以在大肠杆菌细胞中合成并积累ε-胡萝卜素或其中间产物δ-胡萝卜素，使大肠杆菌呈现橙黄色；而没有番茄红素ε-环化酶的情况下大肠杆菌仍然是橙红色，可以很容易分辨是否有番茄红素ε-环化酶活存在。
4)将可以正常表达的pMAL-PyLCYE与pAC-LYC用热激法共转进大肠杆菌BL21(DE3)菌株里。以空载体pMAL-C5X与pAC-LYC共转作为负对照。共转的菌株在含氯霉素(50μg·mL-1)和羧苄霉素(50μg·mL-1)的LB培养基中培养。挑选黄色菌落接种至5mL带有氯霉素和羧苄青霉素的液体LB培养基，37℃、200rpm振荡培养过夜，取200μL接种至20mL抗性LB培养基中200rpm振荡培养3d，离心收集菌体，发现在含有条斑紫菜PyLCYE的样品中，大肠杆菌的颜色都呈橙黄色，而含有空载体pMAL-C5X的负对照大肠杆菌呈橙红色，说明PyLCYE和具有催化番茄红素ε-环化的活性，从而使大肠杆菌细胞积累δ-胡萝卜素。
5)色素提取及HPLC分析
将含有双载体pAC-LYC和pMAL-C5X/pMAL-PyLCYE的大肠杆菌菌体经10,000g离心1min收集后超声破碎。向超声破碎的材料中加入400μL 80％的丙酮，剧烈振荡30min以彻底提取材料中所含色素，然后依次加入250μL乙酸乙酯和ddH2O，分别剧烈振荡15sec，静置5min后，10,000g、4℃离心5min。吸取上清，用氮气吹干后溶于100μL乙酸乙酯。所提取的色素用反相高效液相色谱(reverse-phase HPLC)方法，在Spherisorb ODS2 C18色谱柱(Waters)上分离，色谱柱为4.6×250mm，流动相为在乙腈:水:三乙胺(9:1:0.01)中线性展开乙酸乙酯(0-100％)。展开时间为45min，流速为1mL·min-1，柱温50℃。检测器扫描波长为300～800nm。选择296nm和440nm的扫描结果。类胡萝卜素的鉴定根据标准品或已报道的保留时间和吸收谱来确定。所有的化学试剂均为色谱纯。分析结构表明样品中积累了δ-胡萝卜素，而负对照中没有δ-胡萝卜素的积累(图3)，进一步证明了PyLCYE可以催化番茄红素的ε-环化生成δ-胡萝卜素。
6)***学分析
为了确定所克隆基因PyLCYE所属的类型，，我们检索了GenBank里PyLCYE的同源序列，并根据前人报道取不同植物的番茄红素环化酶序列并对其进行***发生分析。所有序列用ClustalX比对，用MEGA5.1(Tamura et al.,2011)中的邻接法构建***树。自举检验重复1,000次用于分析每个节点的可靠性。分析结果显示PyLCYE确实属于植物中的番茄红素ε-环化酶(图4所示)。
序列表
<110> 南京大学
<120> 一种番茄红素ε-环化酶基因及其编码蛋白和应用
<160> 8
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1830
<212> DNA
<213> 条斑紫菜(Pyropia yezoensis)
<400> 1
atgcccccct ccgcggcctt cctgcccgtc cccccgcggg cgcccgccgc gggcgctgcc 60
gccgccgcgg gcacccgggc gccccggcgg ggctgccgac tacgcccgcg ggtgtcggtg 120
gtcgccacca gccggtcggc gggggcggac gcggtccggc cggcgcgggg ggtgtggcca 180
ccgcgcggcg cggccgccgc atcacccccc acccccaccc ccacccccac tcccgtggtc 240
accgtgtcag cggcagctgc gacagcgtcc tcgacggcgg cgccgaccgc ggcggtgccg 300
gtagcgccag cgacggcacc gtcggtggcc acggatccgg ctgctccccc gctccgcacc 360
tatgacgtga cggtcattgg gggtggaccg gcgggcctct ccctggccgc cgcccttggc 420
gccacgggcc tgagtgtggc cgccctcgac gctgccattg atgtgccgtg gcccaaccac 480
tatggtgtgt gggcagatga gtttgagccc ctcggcctgg ccgactgcgc cacctccacc 540
tacccgacca ccaccatcta tgtgccgtct gccgcaggtg ggggagcggg ggcggcgcct 600
gttgtgctcg accgccccta cctgcgggtg gaccgggtga agctcaaggc gcggctgctg 660
gagcggtgcg cagcgggagg ggtggtcatt gaacgggcaa acgtgaccgc ggtggatcac 720
agtagcgata ctcactcgac ggtgacattt gtggagggtc acccgcgccc atccacatcg 780
aggggggacg gcgcgggctc accaccaacc ccaccgtcgg cgcctgcacc acgacagctg 840
cggacgcggc tggttgtgga cgccactggt cacgcgctgc gatttgtaca gaccgcccca 900
ggctacagcc ccggcttcca ggccgcttat ggcattgagg cggttgttga gggccacccc 960
tttcctctcg acgagatgct gctgatggac tttcgggacg accacatgca gggggatgcg 1020
gccgatgtgg ctgcctcgtc ggcggcgccc accttcctct acgtctttcc cacagatagc 1080
aaccgggtct ttctggagga aacgtcgatc attttgcccg aagccatgcc atttgaaaca 1140
ctgcgggagc ggctgaacaa gcggcttgcc cacctgggcg tcaccgtcaa gttggtggtc 1200
gaggaggagt actcgctgat ccccctgggc gggtcggtgc cgctgctcgg ccagcgggtt 1260
gttggatttg gggggtctgc ctgcctcgtc cacccggcca cgggctacat ggttgcccgc 1320
acactgtcac tcgccgagtc gctcgcgggg gagattgcca ctgccttgcg gactcccgct 1380
gcagggacgt ctcgtcgggg atggggggcg aatgccgccg ccgccgccga cgcaccgcca 1440
ccgcccctgg cgcccgcgga cacggtggcg gccgccgcat gggcgcacct gtggtctgaa 1500
tcccgaatcc gcgagcggga ctttctcctc tttggcgccg acctcttggc gggcctggac 1560
ctgggggcga tgaaggagtt cttcgccgcc tttttccgcc tgccccaacc actgtgggcg 1620
ggcttcctca gctttcggct ggagcggccg agcgagcggg cgacgtttgc gtttgtcttt 1680
ttcctccagg cctctaaccg catccggggg gggctgctgc gggggattgt gacgacgggg 1740
cggtggaaac tgatccgctc ggtgctgccc ctgtggctgg cacggctggg cgactcttct 1800
cgcgaacagc ggggggagaa ccagcagtag 1830
<210> 2
<211> 609
<212> PRT
<213> 条斑紫菜(Pyropia yezoensis)
<400> 2
Met Pro Pro Ser Ala Ala Phe Leu Pro Val Pro Pro Arg Ala Pro Ala
1 5 10 15
Ala Gly Ala Ala Ala Ala Ala Gly Thr Arg Ala Pro Arg Arg Gly Cys
20 25 30
Arg Leu Arg Pro Arg Val Ser Val Val Ala Thr Ser Arg Ser Ala Gly
35 40 45
Ala Asp Ala Val Arg Pro Ala Arg Gly Val Trp Pro Pro Arg Gly Ala
50 55 60
Ala Ala Ala Ser Pro Pro Thr Pro Thr Pro Thr Pro Thr Pro Val Val
65 70 75 80
Thr Val Ser Ala Ala Ala Ala Thr Ala Ser Ser Thr Ala Ala Pro Thr
85 90 95
Ala Ala Val Pro Val Ala Pro Ala Thr Ala Pro Ser Val Ala Thr Asp
100 105 110
Pro Ala Ala Pro Pro Leu Arg Thr Tyr Asp Val Thr Val Ile Gly Gly
115 120 125
Gly Pro Ala Gly Leu Ser Leu Ala Ala Ala Leu Gly Ala Thr Gly Leu
130 135 140
Ser Val Ala Ala Leu Asp Ala Ala Ile Asp Val Pro Trp Pro Asn His
145 150 155 160
Tyr Gly Val Trp Ala Asp Glu Phe Glu Pro Leu Gly Leu Ala Asp Cys
165 170 175
Ala Thr Ser Thr Tyr Pro Thr Thr Thr Ile Tyr Val Pro Ser Ala Ala
180 185 190
Gly Gly Gly Ala Gly Ala Ala Pro Val Val Leu Asp Arg Pro Tyr Leu
195 200 205
Arg Val Asp Arg Val Lys Leu Lys Ala Arg Leu Leu Glu Arg Cys Ala
210 215 220
Ala Gly Gly Val Val Ile Glu Arg Ala Asn Val Thr Ala Val Asp His
225 230 235 240
Ser Ser Asp Thr His Ser Thr Val Thr Phe Val Glu Gly His Pro Arg
245 250 255
Pro Ser Thr Ser Arg Gly Asp Gly Ala Gly Ser Pro Pro Thr Pro Pro
260 265 270
Ser Ala Pro Ala Pro Arg Gln Leu Arg Thr Arg Leu Val Val Asp Ala
275 280 285
Thr Gly His Ala Leu Arg Phe Val Gln Thr Ala Pro Gly Tyr Ser Pro
290 295 300
Gly Phe Gln Ala Ala Tyr Gly Ile Glu Ala Val Val Glu Gly His Pro
305 310 315 320
Phe Pro Leu Asp Glu Met Leu Leu Met Asp Phe Arg Asp Asp His Met
325 330 335
Gln Gly Asp Ala Ala Asp Val Ala Ala Ser Ser Ala Ala Pro Thr Phe
340 345 350
Leu Tyr Val Phe Pro Thr Asp Ser Asn Arg Val Phe Leu Glu Glu Thr
355 360 365
Ser Ile Ile Leu Pro Glu Ala Met Pro Phe Glu Thr Leu Arg Glu Arg
370 375 380
Leu Asn Lys Arg Leu Ala His Leu Gly Val Thr Val Lys Leu Val Val
385 390 395 400
Glu Glu Glu Tyr Ser Leu Ile Pro Leu Gly Gly Ser Val Pro Leu Leu
405 410 415
Gly Gln Arg Val Val Gly Phe Gly Gly Ser Ala Cys Leu Val His Pro
420 425 430
Ala Thr Gly Tyr Met Val Ala Arg Thr Leu Ser Leu Ala Glu Ser Leu
435 440 445
Ala Gly Glu Ile Ala Thr Ala Leu Arg Thr Pro Ala Ala Gly Thr Ser
450 455 460
Arg Arg Gly Trp Gly Ala Asn Ala Ala Ala Ala Ala Asp Ala Pro Pro
465 470 475 480
Pro Pro Leu Ala Pro Ala Asp Thr Val Ala Ala Ala Ala Trp Ala His
485 490 495
Leu Trp Ser Glu Ser Arg Ile Arg Glu Arg Asp Phe Leu Leu Phe Gly
500 505 510
Ala Asp Leu Leu Ala Gly Leu Asp Leu Gly Ala Met Lys Glu Phe Phe
515 520 525
Ala Ala Phe Phe Arg Leu Pro Gln Pro Leu Trp Ala Gly Phe Leu Ser
530 535 540
Phe Arg Leu Glu Arg Pro Ser Glu Arg Ala Thr Phe Ala Phe Val Phe
545 550 555 560
Phe Leu Gln Ala Ser Asn Arg Ile Arg Gly Gly Leu Leu Arg Gly Ile
565 570 575
Val Thr Thr Gly Arg Trp Lys Leu Ile Arg Ser Val Leu Pro Leu Trp
580 585 590
Leu Ala Arg Leu Gly Asp Ser Ser Arg Glu Gln Arg Gly Glu Asn Gln
595 600 605
Gln
<210> 3
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
acccgcaggt aggggcggtc gagcacaa 28
<210> 4
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
cgcggtcacg tttgcccgtt caatgaccac 30
<210> 5
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
cctccaggcc tctaaccgca tccggggg 28
<210> 6
<211> 29
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
tgatccgctc ggtgctgccc ctgtggctg 29
<210> 7
<211> 45
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
ccgcgatatc gtcgacggat ccgccaccag ccggtcggcg ggggc 45
<210> 8
<211> 46
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
tacctgcagg gaattcggat ccctactgct ggttctcccc ccgctg 46
Claims (9)
1.一种番茄红素ε-环化酶基因，其特征在于它的核苷酸序列如序列表Seq ID No：1所示。
2.一种番茄红素ε-环化酶基因，其特征在于它的核苷酸序列为与序列表SEQ ID NO:1核苷酸序列具有80％以上同源性且编码相同功能蛋白质的多核苷酸序列。
3.一种番茄红素ε-环化酶基因编码的蛋白，其特征在于它的氨基酸序列如序列表SeqID No：2所示。
4.一种番茄红素ε-环化酶基因编码的蛋白，其特征在于它的氨基酸序列为由序列表Seq ID No：2所述的氨基酸序列经过一个或者几个氨基酸残基的取代、缺失或者添加且具有与序列表Seq ID No：2所述的氨基酸残基序列相同活性的氨基酸序列。
5.根据权利要求1、2、3或4所述的番茄红素ε-环化酶基因编码的蛋白，其特征在于番茄红素ε-环化酶来源条斑紫菜。
6.一种重组表达载体，其特征在于它包括权利要求1或2所述的基因。
7.一种转化子，其特征在于它包括含有权利要求6所述的重组表达载体的宿主细胞。
8.根据权利要求7所述的一种转化子，其特征在于它的宿主为微生物、植物或转基因细胞系。
9.一种番茄红素ε-环化酶基因的应用，其特征在于它用于紫菜类胡萝卜素代谢基因工程中。
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| CN111454976A (zh) * | 2020-03-17 | 2020-07-28 | 深圳大学 | 用于提高莱茵衣藻叶黄素含量的转基因衣藻及其构建方法、应用 |
| CN111518779A (zh) * | 2020-05-08 | 2020-08-11 | 沈阳药科大学 | 一种恶唑环化酶基因及其编码蛋白和应用 |
| CN113549639A (zh) * | 2021-07-21 | 2021-10-26 | 云南中烟工业有限责任公司 | 一种降低烟叶总蛋白及烟气苯酚含量的调控基因 |
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| CN111454976B (zh) * | 2020-03-17 | 2022-11-01 | 深圳大学 | 用于提高莱茵衣藻叶黄素含量的转基因衣藻及其构建方法、应用 |
| CN111518779A (zh) * | 2020-05-08 | 2020-08-11 | 沈阳药科大学 | 一种恶唑环化酶基因及其编码蛋白和应用 |
| CN111518779B (zh) * | 2020-05-08 | 2022-03-11 | 沈阳药科大学 | 一种恶唑环化酶基因及其编码蛋白和应用 |
| CN113549639A (zh) * | 2021-07-21 | 2021-10-26 | 云南中烟工业有限责任公司 | 一种降低烟叶总蛋白及烟气苯酚含量的调控基因 |
| CN113549639B (zh) * | 2021-07-21 | 2022-07-29 | 云南中烟工业有限责任公司 | 一种降低烟叶总蛋白及烟气苯酚含量的调控基因 |
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