CN115011567A - (+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用 - Google Patents
(+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用 Download PDFInfo
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- CN115011567A CN115011567A CN202210726618.0A CN202210726618A CN115011567A CN 115011567 A CN115011567 A CN 115011567A CN 202210726618 A CN202210726618 A CN 202210726618A CN 115011567 A CN115011567 A CN 115011567A
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Abstract
本申请公开了(+)‑新薄荷醇合酶、合酶基因及其在植物抗病中的应用。该(+)‑新薄荷醇合酶是氨基酸序列如SEQ ID NO.1所示且具有(+)‑新薄荷醇合成酶活性的蛋白。本申请公开了一种编码(+)‑新薄荷醇合酶以及具备(+)‑新薄荷醇合酶的蛋白的基因CsSDR1，其序列如SEQ ID NO.2所示。本申请建立了生物合成(+)‑新薄荷醇的方法，可应用于(+)‑新薄荷醇的生物合成。本申请中，(+)‑新薄荷醇合酶、合酶基因可应用于提高植物尤其是茶树的抗病性。
Description
技术领域
本申请属于植物分子生物学领域，尤其涉及(+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用。
背景技术
植物生活在一个暴露于各种形式的生物胁迫(病原体和食草动物)的环境中。植物通过合成各种次生代谢产物，介导防御信号或生成直接防御武器来阻止病原体的入侵和昆虫的取食，大多数次生代谢物都有直接的抗菌活性。解析参与植物防御相关的代谢物为培育抗病植物提供了广泛而稳定的资源。
在这些次生代谢物中，萜类化合物是最大和最多样化的一类，对各种草食动物和病原体具有活性的天然产物。萜类化合物是由各种TPSs以GPP、FPP及GGPP等重要前体为底物生成以异戊二烯为基本结构单元的萜烯类物质及由各种修饰酶(如短链脱氢酶/还原酶)对基本萜烯类物质进行修饰形成的各种衍生物一起组合而成。许多研究报道了萜类化合物含量与抗病性之间的存在正相关关系。单萜生物合成的主要代谢途径的最终产物之一(+)-新薄荷醇，可以作为食品添加剂和香气单体用于食品和香水为其提供独特的风味和清凉感。
茶是最受欢迎的不含酒精的饮料。茶树种植是产茶国的重要经济来源。由病原真菌引起的茶树病害已严重影响茶叶产量和品质。然而，现阶段茶树病害的防控仍以化学药剂的后期治疗为主。长期和过度使用化学杀菌剂可能导致加工后的茶叶中出现农药残留，对环境和人体产生不良影响。也可能使得病原体对化学杀菌剂产生耐药。
因此，剖析萜类化合物在茶树抗病性中的功能及分子机制，不仅为茶树抗病品种的选育提供理论依据，同时对绿色、高效、安全的治理茶树病害具有重要意义。
发明内容
有鉴于此，本申请的目的在于公开了(+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用，以解决或缓解部分上述技术问题。
第一方面，本申请实施例提供了一种酶，催化(-)-薄荷酮合成(+)-新薄荷醇，其中所述酶为A～C中的至少一项：
A.其氨基酸序列如SED ID NO.1所示的蛋白质；
B.具有与A中所述蛋白质至少90％同源性氨基酸序列且具有和A中所述蛋白质相同酶活性的蛋白质；
C.由A或B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/或添加且具有和A或B所述蛋白质相同酶活性的衍生的蛋白质。
第二方面，本申请实施例公开了一种基因，所述基因用于编码第一方面所述酶，其中所述基因至少为(1)～(3)中至少一项：
(1)核苷酸序列如SED ID NO.2所示的DNA分子；
(2)与(1)所述DNA分子杂交且编码具有第一方面酶活性的蛋白的DNA分子；
(3)与(1)或(2)所述DNA分子具有至少65％、至少70％、至少75％、至少80％、至少85％、至少90％、至少95％或至少99％同源性且编码具有第一方面酶活性的蛋白的DNA分子。
第三方面，本申请实施例公开了含有第二方面所述基因的表达载体或表达盒。
第四方面，本申请实施例公开了含有第三方面所述表达载体或表达盒的重组菌或工程化细胞。
第五方面，本申请实施例公开了第一方面所述酶、第二面所述基因、第三方面所述表达载体或表达盒、第四方面所述重组菌或工程化细胞在合成(+)-新薄荷醇中的应用。
第六方面，本申请实施例公开了第一方面所述酶或第二方面所述基因在茶树抗病中的应用。
与现有技术相比，本申请至少具有以下有益效果：
本申请涉及(+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用。所述酶能够催化(-)-薄荷酮合成(+)-新薄荷醇，本申请还公开了编码所述酶的基因CsSDR1，所述酶和编码所述酶的基因CsSDR1能应用于茶树抗病，通过研究所述酶或所述基因的功能机制，不仅为茶树抗病品种的选育提供理论依据，同时对绿色、高效、安全的治理茶树病害具有重要意义。
附图说明
图1为本申请实施例提供的茶树CsSDR1基因的RCR扩增图。
图2为本申请实施例提供的重组质粒pGX4T1-CsSDR1的PCR鉴定图。
图3为本申请实施例提供的纯化的pGX4T1-CsSDR1蛋白SDS电泳图。
图4为本申请实施例提供的重组蛋白体外酶活的产物GC-MS分析。
图5为本申请实施例提供的茶树受致病菌侵染后的CsSDR1的定量分析。
图6为本申请实施例提供的茶树沉默CsSDR1表达后受致病菌侵染的病斑状况。
具体实施方式
为了使本申请的目的、技术方案及优点更加清楚明白，以下结合实施例对本申请进行进一步详细说明。应当理解，此处所描述的具体实施例仅仅用以解释本申请，并不用于限定本申请。
本申请实施例中所使用的实验方法如无特殊说明，均为常规方法。
本申请实施例中所用的材料、试剂等，如无特殊说明，均可从商业途径得到。
本申请实施例中，山茶炭疽菌(Colletotrichum camelliae)购买于宁波明舟生物科技有限公司，产品编号：BMZ098286；其它试剂中，总RNA提取试剂盒、质粒提取试剂盒、内切酶、高保真super酶购于南京诺唯赞科技公司，反转录试剂盒购于上海翊圣生物科技有限公司，pGX4T1载体、大肠杆菌菌株DH5α和BL21购于上海唯地生物技术有限公司，底物(-)-薄荷酮等其他化学试剂均购于sigma公司，引物和测序均有安徽通用生物公司完成。
(+)-新薄荷醇合酶基因的克隆
1、茶树总RNA的提取和cDNA文库的构建
用液氮迅速将茶叶组织样品研磨成粉状，称取0.05～0.1g样品用于总RNA的提取。使用RNA提取试剂盒提取样品总RNA，通过琼脂糖电泳凝胶检测其完整性，并测定RNA浓度。RNA保存于-80℃。使用RNA反转录试剂盒反转录所获得的RNA构建茶叶cDNA文库，方法参照试剂盒的说明手册。
2、引物对设计与基因克隆
引物对序列：
CsSDR1-F：如SEQ ID No.3所示；
CsSDR1-F：如SEQ ID No.4所示。
以茶树cDNA为模板扩增获得PCR产物(图1所示)，并用凝胶提取试剂盒纯化。
将纯化后的产物连接到经酶BamHI和酶SalI酶切线性化后的PGEX-4T1载体上，再转入感受态细胞DH5α。
将菌液均匀涂布到LB培养基(氨苄青霉素Amp：25μg/mL)上，37℃倒置过夜培养10h～12h。挑取单菌落于300μL液体LB培养基(Amp：25μg/mL)中，37℃、200rpm震荡培养2h，取1μL进行菌落PCR检测。菌落PCR产物用琼脂糖凝胶进行检测(如图2所示)，选择符合目的基因长度的菌液送安徽通用生物有限公司进行测序。
结果：寻找测序后序列的ORF框，获得该基因的全长序列见SEQ ID No.2所示，将基因命名为CsSDR1。该基因编码的CsSDR1蛋白为一种(+)-新薄荷醇合酶，其氨基酸序列见序列如SEQ ID No.1所示，共编码311个氨基酸。
CsSDR1蛋白在合成萜烯类化合物(+)-新薄荷醇中的应用
1、CsSDR1外源蛋白的表达和纯化
重组质粒和阴性对照转入BL21(DE3)pLysS感受态细胞，均匀涂布在LB培养基上(Amp：25μg/mL，氯霉素Cm：50μg/mL)，37℃倒置过夜培养10～12h。
挑取单克隆于相同抗性的液体LB培养基中，37℃、200rpm震荡培养至OD600在0.6～0.8之间。待培养基冷却到16℃后，加入终浓度为1mM异丙基-β-D-硫代半乳糖苷(IPTG)，16℃、150rpm震荡培养22h后离心收集菌落，超声破碎，使用GST结合树脂纯化，使用考马斯亮蓝法测定纯化后的蛋白浓度。
结果：通过十二烷基硫酸钠-聚丙烯酰胺凝胶电泳(SDS-PAGE)验证蛋白的大小，如图3所示。含有GST标签的重组CsSDR1蛋白是一种溶液中的单体酶，分子量约为60kD(标签大小约为24kD)
2、酶活性检测
使用如下反应体系对酶的活性进行初步检测。
反应体系为：30μg纯酶，1mL测定缓冲液(50mMhEPES，pH 7.5，100mM NaCl和5mMβ-巯基乙醇)，1mM薄荷酮和500μM NADPH。31℃下孵育12h。之后用0.5mL正己烷提取单萜产物。
反应产物用GC-MS进行检测，使用Chameleon软件，将采集到的数据在NIST标准谱库中进行检索分析。
结果：通过与标准品比较GC保留时间和质谱图进行鉴定分析，发现CsSDR1蛋白以NADPH为辅因子，以(-)-薄荷酮为底物，在pH 7.5条件下，生成以(+)-新薄荷醇为主的产物，其中(+)-新薄荷醇约占产物总峰面积的93％，也有少量的(-)-薄荷醇生成，约占产物峰面积的7％。质谱总离子流图如图4所示。
CsSDR1基因在植物抗病中的应用
1.CsSDR1基因表达受病原菌的侵染诱导
对致病菌侵染后的茶树中CsSDR1基因进行实时荧光定量PCR(RT-qPCR)检测，其中致病菌为山茶炭疽菌(Colletotrichum camelliae)。
引物对序列：
qSDR1-1F：如SEQ ID No.5所示：
qSDR1-1R：如SEQ ID No.6所示。
发现该基因受菌侵染的而大量诱导(如图5所示)，表参与茶树对病原菌的抵抗
以上实例中的致病菌为山茶炭疽菌(Colletotrichum camelliae)。
以上实例中定量引物为：
qSDR1-1F：5’-CCTGGTGCTGTTGACTGCTA-3’(SEQ ID No.5)
qSDR1-1R：5’-CATCCCAATCCGTCACGACT-3’(SEQ ID No.6)
RT-qPCR的扩增程序为：
预变性95℃，5min，变性95℃，10s，退火/延伸60℃，30s，再返回变性、退火/延伸，循环40次。
实时荧光定量PCR中，以甘油醛-3-磷酸脱氢酶(GAPDH)为内参基因相对定量，使用2-ΔCT法计算各基因的相对表达量。
结果：如图5所示，不同的小写字母表示各处理间经Duncan′s多重比较，在p<0.5水平上差异显著。从图5中可以看出，茶树受菌侵染后而诱导了CsSDR1基因的大量产生，表明CsSDR1基因参与茶树对病原菌的抵抗。
2、抑制该基因表达后的茶树更易感病
利用反义寡核苷酸(AsODN)技术对将该基因瞬时沉默，定量方法及定量程序同实例三中1一致使用反义寡核苷酸(AsODN)处理茶树24h能显著降低茶树体内CsSDR1的表达量(如图6所示)；使用致病菌侵染沉默后的茶树4天，所述致病菌为山茶炭疽菌(Colletotrichumcamelliae)，结果发现沉默组茶苗病斑面积显著高于对照组(如图6所示)，结果表明茶树中CsSDR1基因的表达量与其抗病性呈正相关。其中，图6的‘***’表示经t检验，p值<0.001。
综上所述，本申请提供了一种(+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用。所述酶能够催化(-)-薄荷酮合成(+)-新薄荷醇，本申请还提供了编码所述酶的基因CsSDR1，所述酶和编码所述酶的基因CsSDR1能应用于茶树抗病。
以上结合附图详细描述了本申请的优选实施方式，但是，本申请并不限于上述实施方式中的具体细节，在不脱离本申请的精神和范围内，都可做各种的改动与修饰，因此本申请的保护范围应该以权利要求书所界定的为准。
以上所述，仅为本申请较佳的具体实施方式，但本申请的保护范围并不局限于此，任何熟悉本技术领域的技术人员在本申请揭露的技术范围内，可轻易想到的变化或替换，都应涵盖在本申请的保护范围之内。
序列表
<110> 安徽农业大学
<120> (+)-新薄荷醇合酶、合酶基因及其在茶树抗病中的应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 300
<212> PRT
<213> 茶树(Camellia sinensis+-新薄荷醇合酶的蛋白序列)
<400> 1
Met Ala Glu Lys Ile Ser Thr Thr Lys Arg Tyr Ala Val Val Thr Gly
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Ala Asn Lys Gly Leu Gly Phe Glu Ile Cys Arg Gln Leu Ala Ser Lys
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Gly Ile Leu Val Leu Leu Thr Ala Arg Asp Glu Lys Lys Gly Leu Glu
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Ala Leu Glu Lys Leu Lys Ser Asp Gly Leu Phe Asp His Val Val Phe
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Phe Gln Leu Asp Val Val Asp Pro Ser Ser Ile Ala Ser Leu Ala Asp
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Phe Ile Lys Ser Lys Phe Gly Arg Leu Asp Ile Leu Val Asn Asn Ala
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Gly Ile Ser Gly Val Val Thr Asp Trp Asp Ala Leu Lys Ser Ala Asp
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Ala Ala Asp Pro Gly Leu Leu Asn Lys Trp Lys Glu Val Met Thr Gln
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Asn Tyr Glu Leu Thr Asp Glu Cys Leu Gln Thr Asn Tyr Tyr Gly Ala
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Lys Arg Met Ile Glu Ala Leu Ile Pro Leu Leu Gln Leu Ser Asp Ser
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Pro Arg Ile Val Asn Val Ser Ser Gly Met Gly Lys Leu Lys Asn Ile
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Pro Ser Glu Trp Ala Lys Gly Ala Leu Asn Asp Ala Glu Ser Leu Thr
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Glu Glu Arg Ile Asp Glu Val Leu Asn Lys Phe Leu Lys Asp Phe Lys
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Glu Gly Ser Leu Glu Ser Lys Gly Trp Pro Thr Leu Leu Ser Ala Tyr
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Thr Leu Ser Lys Ala Ala Met Asn Ala Tyr Thr Arg Phe Leu Ala Lys
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Lys Tyr Pro Thr Phe Arg Ile Asn Cys Val Cys Pro Gly Tyr Val Lys
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Thr Asp Ile Asn Asn Asn Asn Gly Ile Leu Ser Thr Glu Gln Gly Ala
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Glu Cys Pro Val Lys Leu Ala Leu Leu Pro Asp Glu Gly Pro Ser Gly
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Leu Phe Phe Val Cys Asn Glu Leu Ser Ser Phe Glu
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<210> 2
<211> 936
<212> DNA
<213> 茶树(Camellia sinensis+-新薄荷醇合酶的DNA序列)
<400> 2
atggcagaga aaatcagtac aacaaagaga tacgcagtgg ttacaggggc aaataaagga 60
ctaggatttg aaatatgtag gcaattagct tctaagggaa tcctggtgct gttgactgct 120
agagatgaga agaagggtct tgaagctctt gagaaactca aaagtgatgg cctctttgat 180
catgtggtgt tctttcagct tgatgttgtg gacccatcta gtattgcttc ccttgccgat 240
ttcatcaagt ctaaatttgg aaggcttgat atcttggtga acaatgctgg gattagtgga 300
gtcgtgacgg attgggatgc tttaaagtca gcagatgctg ctgatcctgg attgcttaac 360
aaatggaaag aagtaatgac tcaaaactat gagttgactg atgaatgctt gcaaacaaac 420
tactatggag caaaaaggat gattgaagca ctcattcccc tcctccaact ctcagattca 480
ccaaggatag taaatgtctc ttctggcatg gggaagttaa agaacatacc cagtgaatgg 540
gctaaaggag tgttgaatga tgccgaaagc cttacagaag agagaataga cgaggtacta 600
aacaagtttc taaaagattt caaagaaggt tcgttagaat ccaaaggctg gcctacgttg 660
ttgtctgcct atacattgtc gaaagcggct atgaacgcgt acacaagatt tctggccaaa 720
aagtatccca cattccgcat caattgtgtc tgccccggtt atgttaaaac agatataaac 780
aacaacaacg gcatactaag cacagaacaa ggtgctgaat gtcctgtgaa gctagcccta 840
cttcctgatg aaggaccttc tggcctcttt tttgtttgca atgaactgtc tcttttgagt 900
gacccgggtc gactcgagcg gccgcatcgt gactga 936
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<212> DNA
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<212> DNA
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Claims (6)
1.一种酶，其为A～C中的至少一项：
A.其氨基酸序列如SED ID NO.1所示的蛋白质；
B.具有与A中所述蛋白质具有至少90％同源性的氨基酸序列且具有和A中所述蛋白质相同酶活性的蛋白质；
C.由A或B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/或添加且具有和A或B所述蛋白质相同酶活性的衍生的蛋白质。
2.一种基因，其为(1)～(3)中至少一项：
(1)核苷酸序列如SED ID NO.2所示的DNA分子；
(2)与(1)所述DNA分子杂交且编码具有权利要求1酶活性的蛋白的DNA分子；
(3)与(1)或(2)所述DNA分子具有至少65％、至少70％、至少75％、至少80％、至少85％、至少90％、至少95％或至少99％同源性且编码具有权利要求1酶活性的蛋白的DNA分子。
3.含有权利要求2所述基因的表达载体或表达盒。
4.含有权利要求3所述表达载体或表达盒的重组菌或工程化细胞。
5.权利要求1所述酶、权利要求2所述基因、权利要求3所述表达载体或表达盒、权利要求4所述重组菌或工程化细胞在合成(+)-新薄荷醇中的应用。
6.权利要求1所述酶、权利要求2所述基因在茶树抗病中的应用。
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| YING ZHOU ET AL.: "Molecular Cloning and Characterization of a Short-Chain Dehydrogenase Showing Activity with Volatile Compounds Isolated from Camellia sinensis", 《PLANT MOL BIOL REP 》, vol. 33, pages 253 - 263, XP035461545, DOI: 10.1007/s11105-014-0751-z * |
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