CN111019911A - 葡萄脱落酸降解途径关键酶8’-羟化酶cyp707a蛋白、编码基因及其应用 - Google Patents
葡萄脱落酸降解途径关键酶8’-羟化酶cyp707a蛋白、编码基因及其应用 Download PDFInfo
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- CN111019911A CN111019911A CN201911058375.2A CN201911058375A CN111019911A CN 111019911 A CN111019911 A CN 111019911A CN 201911058375 A CN201911058375 A CN 201911058375A CN 111019911 A CN111019911 A CN 111019911A
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Abstract
一种葡萄脱落酸降解途径关键酶8’‑羟化酶CYP707A蛋白、编码基因及其应用,其特征在于:该蛋白的氨基酸序列如SEQ ID No.2所示,该编码基因的序列如SEQ ID No.1所示。能填补葡萄果实成熟过程中的调节因子,完善植物果实成熟的机制,填补脱落酸降解途径关键基因CYP707A家族成员的克隆、表达性分析及在新品种“鄞红”葡萄果实中CYP707A蛋白以及编码基因的空白。
Description
技术领域
本发明涉及“鄞红”葡萄中调节果实成熟的植物激素脱落酸(abscisic acid,ABA)降解途径中的关键酶与其编码基因,具体涉及葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白、编码基因及其应用,属于生物技术领域。
背景技术
植物中,果实不仅拥有植物的繁殖和传播的作用,而且也具有一定的经济和食用价值,又因其口味独特且营养价值高等特点,广受人们的喜爱。果实的生长发育是一个复杂而又循序渐进地过程,主要受外源因子、内源激素以及转录因子共同调控。内源基因控制多种内源激素的合成与降解,已知促进果实成熟相关植物激素主要有脱落酸 (Abscisicacid,ABA)、乙烯(Ethylene)、油菜素类脂(BRs),抑制果实成熟植物激素有生长素(IAA、NAA等)、细胞分裂素(CTK)、赤霉素(GAs)等。果实逐渐成熟的表现为:果实开始软化、可溶性固形物含量开始迅速上升、糖酸比升高、果面着色等。研究表明,ABA在调节非呼吸跃变型果实成熟的过程中起着重要的作用。
葡萄(Vitis vinifera)为葡萄科葡萄属木质藤本植物,具有很高的食用和经济价值,又因其口味独特,身受人们的喜爱。葡萄果实属于非呼吸跃变型果实,生长包括2 个相继的S周期,由形成期、停滞期和成熟期组成。脱落酸在葡萄果实的成熟过程中起着启动的作用,但具体的启动方式还有待继续研究。在高等植物中ABA的分解代谢可以通过氧化失活和结合失活两个步骤来完成。8’-羟化酶是氧化失活途径中的关键酶,由 CYP707A基因家族控制合成,CYP707A基因在高等植物中广泛存在,因而CYP707A基因是脱落酸氧化降解途径中关键限速酶基因。目前,该基因在拟南芥、番茄、烟草等植物中研究较为广泛,而在葡萄中的研究相对较少,且CYP707A基因的克隆、表达性分析都少见报道。
发明内容
本发明针对现有技术的上述不足,提供一种能填补葡萄果实成熟过程中的调节因子,完善植物果实成熟的机制,填补脱落酸降解途径关键基因CYP707A家族成员的克隆、表达性分析及在新品种“鄞红”葡萄果实中CYP707A蛋白以及编码基因的空白。
为了解决上述技术问题,本发明采用的技术方案为:一种葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白,该蛋白的氨基酸序列如SEQ ID No.2所示。
一种上述葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白的编码基因,该编码基因的序列如SEQ ID No.1所示。
优选的,该基因是从葡萄新品种“鄞红”葡萄的果实筛选获得。
一种含有所述葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白编码基因的表达盒、重组表达载体、转基因细胞系或重组菌也属于本发明的保护范围。
一种上述葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白、编码基因、表达盒、重组表达载体、转基因细胞系或重组菌在葡萄育种中的应用。
本发明的8’-羟化酶CYP707A基因通过葡萄果实CYP707A基因的克隆获得,具体的包括:(1)植物材料的获得,(2)RNA和DNA的提取,(3)基因的全长克隆。
本发明上述植物材料是采用鄞红”葡萄的果实。
本发明上述基因的全长克隆过程中使用了简并引物,该引物序列如下: F-1-CYP707A:ATGGGTTGGCCTTAYATAGGAGAAAC(SEQ ID No.3);R-4-CYP707A:ACTTTCCACCCTTTTGGT ATAAGATA(SEQ ID No.4);使用了PCR扩增引物,该引物序列如下:3'-C-5:CAAGCTGTCACGGAAGAGCAAGAGTCCATAA(SEQ ID No.5);使用了葡萄全基因组所需引物,该引物的序列如下:F-CYP707A-D2:TTTCTTCTCCACTCCTCACACCCATCCT(SEQ ID No.6);R-CYP707A-D2:GCCTTTGTGACCAGCACCAGCTTTG(SEQ ID No.7)。
本发明的这种8’-羟化酶CYP707A蛋白以及编码基因获得之后,对ABA脱落酸代谢途径进行了完善,有助于了解葡萄果实成熟机制,为下游实验的进行提供基础,以及为葡萄新品种的产生提供技术支持。
附图说明
图1 VvCYP707A基因的全基因组分析图。
图2 VvCYP707A基因开放阅读框预测图。
注:图中阴影部分为最大开放阅读框
图3“鄞红”葡萄CYP707A蛋白亲疏水性分布图。
注:横坐标为序列位置,纵坐标为氨基酸的标度值。Hphob.Kyte&Doolittle标度(默认值)定义疏水性氨基酸较高打分值(>0表示疏水性,<0表示亲水性)。
图4 CYP707A蛋白跨膜区预测分析图。
图5“鄞红”葡萄CYP707A蛋白二级结构预测结果图。
图6“鄞红”葡萄CYP707A蛋白三级结构预测结果图。
图7亚细胞定位结果。
图8 VvCYP707A蛋白基因家族与其他物种的同源性分析图。
图9 CYP707A基因家族的系统进化树分析图。
具体实施方式
下面通过具体实施例进一步详细描述本发明,但是本发明不仅仅局限于以下实施例。
实施例1、葡萄果实CYP707A基因的克隆
1.植物材料的获得
本实验所用的植物材料为吴月燕科研团队培育的葡萄新品种-“鄞红”葡萄的果实(原名:甬优1号,市售常熟品种;宁波东钱湖旅游度假区野马湾葡萄场,浙江万里学院,宁波市鄞州区林业技术管理服务站作为选育单位,该品种为欧美杂种。树势强,叶色浓,叶面光滑无毛。一般穗重500-700克。果实椭圆形,平均粒重12克左右。果皮厚韧,紫红色,果粉中等。果肉硬,风味甜,可溶性固形物17%以上。宁波地区大棚保护地栽培之成熟期一般为7月底至8月中旬。多年田间调查表明,该品种抗病性达到与巨峰和藤稔相等水平)。实验材料采摘自宁波市镇海滴翠园农场。在自然条件下,采摘葡萄不同发育时期(幼果期、膨大期、转色期、成熟期和过熟期)的果实提取RNA和DNA。
2.RNA和DNA的提取
采用total RNA提取试剂盒(RNAprep Pure Plant Kit,Polysaccharides&Polyphenolics-rich)提取RNA(购自天根生物科技公司)。变性后用1%琼脂糖凝胶电泳鉴定RNA的完整性,在分光光度计(Thermo Scientific NanoDrop2000)上测定RNA的纯度和浓度。采用HP Plant DNA Kit试剂盒提取总DNA(OMEGA Bio-tek(美国)公司)。
3.基因的全长克隆
基于Genebank中CYP707A基因的保守序列,用软件MEGA7.0进行序列比对,找出保守性高的区域设计简并引物,将提取的RNA进行逆转录(2XEs Taq MasterMix(Dye) 购自康伟生物有限公司),以第一链cDNA为模板,利用简并引物:F-1-CYP707A:ATGGGTTGGCCTTAYATAGGAGAAAC;R-4-CYP707A:ACTTTCCACCCTTTTGGT ATAAGATA进行PCR 扩增,PCR扩增反应程序:95℃预热5min;95℃变性30s,50~60℃退火30s,72℃延伸 2min;变性、退火、延伸共35个循环,最后72℃延伸10min。扩增得到的预期的目的条带后回收并连接到pMD-18T(宝日医生物技术(北京)有限公司)载体上,转化大肠杆菌DH5α,挑取白色单菌落进行菌液PCR鉴定,筛选阳性克隆,送至生工生物工程(上海)股份有限公司进行测序,得到DNA序列。
根据测序结果分析得到一段DNA序列,根据得到的序列设计一对引物:将提取的RNA进行逆转录(5’/3’Kit:宝日医生物技术(北京)有限公司)合成3’和 5’cDNA,以合成的cDNA为模板,利用引物3'-C-5: CAAGCTGTCACGGAAGAGCAAGAGTCCATAA进行PCR扩增,PCR扩增反应程序:94℃30s, 72℃3min,5个循环;94℃30s,70℃30s,72℃3min,5个循环;94℃30s,68℃30s, 72℃3min,25个循环。扩增得到的目的条带转化大肠杆菌DH5α,培养12~16h,挑取白色单菌落经过菌液PCR验证,筛选阳性克隆;送至生工生物工程(上海)股份有限公司进行测序,得到3’序列;将得到的序列进行拼接,得到基因部分全长序列;根据葡萄全基因组设计引物F-CYP707A-D2:TTTCTTCTCCACTCCTCACACCCATCCT;R-CYP707A-D2:GCCTTTGTGACCAGCACCAGCTTTG,并进行PCR扩增,PCR扩增反应程序:95℃预热5min; 95℃变性30s,50~60℃退火30s,72℃延伸2min;变性、退火、延伸共35个循环,最后72℃延伸10min。扩增得到的目的条带转化大肠杆菌DH5α,培养12~16h,挑取白色单菌落经过菌液PCR验证,筛选阳性克隆,送至生工生物工程(上海)股份有限公司进行测序,测序结果序列分析以及拼接,得到基因全长序列。
实施例2.葡萄果实CYP707A基因的序列信息
得到的CYP707A基因经过ORF预测,序列中共有14条开放阅读框,最长的一条始于97终于1509碱基位置,共长1413bp和470AA。根据葡萄全基因组比对,可知CYP707A 基因序列位于葡萄2号染色体上,含有7个外显子,6个内含子,可编码653个氨基酸,且经过分析可知属于CYP707A基因家族中CYP707A1。
实施例3.葡萄果实CYP707A蛋白的理化性质及功能分析
根据Expasy的子程序分析可知:VvCYP707A编码的蛋白质分子式C3421H5336N874O931S25,相对分子质量为74406.80,氨基酸残基数为654,理论等电点为9.49,负电荷氨基酸残基总数(Asp+Glu)为54,正电荷氨基酸残基总数(Arg+Lys) 为82,脂肪系数(AI)87.40,总平均亲水性(GRAVY)为-0.141,不稳定系数(Ⅱ)为 42.85,显示该蛋白较为稳定(稳定蛋白标准在40左右)。
进行亲疏水性和跨膜结构域分析可知,CYP707A蛋白属于疏水性和跨膜蛋白(具体见附图4)。利用PredictProtein(https://www.predictprotein.org/)进行VvCYP707A蛋白二级结构的预测(具体见附图5),结果分析可知,VvCYP707A蛋白的二级组成中螺旋(helix,H)和环(loop,L)所占的比例最大,分别为H:44.95%和L:45.57%,而折叠 (sheet,E)非主要二级结构,仅占总蛋白的9.48%;蛋白质溶剂可及性(核心/表面比率)表明,暴露于膜表面超过16%的残留氨基酸占总蛋白的41.74%,其余残基所占比率为58.26%,且在29~54位置之间有一个跨膜螺旋区。用SWISS-MODEL (https://swissmodel.expasy.org/ interactive/Jyxvzw/models/)进行VvCYP707A蛋白质三级结构的预测和ProtComp(http://www.softberry.com/berry.phtml?topic=protcomppl&group=programs& subgroup=p roloc)亚细胞定位(具体见附图6),三级结构的预测结果与二级结构预测结果进行分析可知基本一致,亚细胞定位分析结果(具体见附图7)表明,预测结果是该蛋白位于内质网的积分值为7.40,位于质膜的积分值为2.55,属于过氧化物酶体的积分值为0.04,线粒体的分值为0.02,其余部位的得分值低于0.01,甚至为0.00,从而可以推断 VvCYP707A蛋白定位于内质网膜上的可能性很大。
实施例4.葡萄果实CYP707A蛋白的同源性分析
利用NCBI在线查找其他物种的CYP707A蛋白质序列,使用DNAMAN 8.0和MEGA 7.0进行序列比对和进化树的构建(具体见附图8-9),进行序列比对和进化树构建表明克隆得到的CYP707A基因属于该家族中的成员,与其它物种具有54.21%的相似度,以及与Camellia sinensis(MF765771.1)有较近亲缘关系,为CYP707A基因家族在葡萄中的研究提供了理论基础。
以上都是对本发明的具体实施方式进行了描述。需理解的是本发明并不是局限于上述特定实施方式,本领域技术人员可以在权利要求的范围内做出各种变性或修改,这并不影响本发明的实质内容。
SEQUENCE LISTING
<110> 浙江万里学院
<120> 葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白、编码基因及其应用
<130> 2019
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ttcttctcca ctcctcacac ccatcctccc cctcctctct tcttcactct ctccaccatc 60
atcaacccac aactactcaa accactacta cttctaatgg actcctcctt caccaccata 120
tgcttctcca ttgctctcct cctctttctc atccactttc tcctaaagct cttctccttc 180
ttctctgctt ccactgcccg gaaactccct ctcccacctg gctctctggg ctggccctac 240
ataggagaaa cctttcagct ttactcccag aaccccaatg tcttctttgc ctccaagcaa 300
gagaggtatg gaaccatatt caaaacccgc atattggggt gcccatgtgt gatgatttca 360
agtcctgaag ctgcaaagct ggtgctggtc acaaaggctc ctctctttaa gcctacattt 420
ccggctagca aagagaggat gttggggaaa caagcaattt tctttcacca aggtgactac 480
catgccaagt tgaggaagct tgttctccga gccttcatgc caggagccat caaaaacatc 540
gtctctaaca tcgactccat cgccacccag accctgcagt cctgggaagg ccggtcgatc 600
aatactttcc aagaaatgaa gacatacact ttcaatgttg cactgctgtc catatttgga 660
aaagatgaaa ttctgtacag agaggaactc aagagatgct actacattct tgagaagggg 720
tataattcaa tgcccataaa cattcctgga acgctcttca acaaatcaat gaaagcaagg 780
aaggaattgg ctcagatctt ggctaaaatc ctctcaacaa ggaggcaggc gaacggtgat 840
cacaacgact tgttgggatc tttcatggga gacaaagaag ggctcacaga tgaacaaatt 900
gcagacaaca tcataggcgt catcttcgcg gcccgagaca ctactgccag tgtgctgaca 960
tggatcctca aatatctggg cgaaaaccca agtgttctgc aagctgtcac ggaagagcaa 1020
gagtccataa tgaggagtaa agaagacaag tgtggtgaag agaagtctct gacttgggaa 1080
gatgccaaga agatgccaat aacttcaagg gtgattcaag aaacacttag aattgcttca 1140
attttatctt tcacctttag agaggctgtt gaagatgttg aatttgaagg gtacctgatt 1200
ccaaaagggt ggaaagtttt accacttttc agaaacattc accacagtcc agacatatat 1260
ccagagccag agaagtttga tccctcaaga tttgaggttg ctccaaagcc caatacattt 1320
atgccatttg gaaatggggt ccactcatgc cctgggaatg agttggccaa actagagatt 1380
ttggtgcttc tacatcatct gactacaaag tacaggtggt ctatggtggg cacacagaat 1440
ggtattcagt atggcccttt tgctcttccc cagaatggtt tgcccatcac aatctccctc 1500
aagtcataga aacaaatctc tactaaatct actctcagtg ccaccaatca acttcactcc 1560
catattccta ttcaaagttt caaacaagaa acaaagaaac aatatgaaga gatctccctt 1620
gtttcctgcc aatctagaga cacccacatg aaggaatttt gatgatcaat actcaaaaag 1680
ggaaccaaaa aataactgta cagcaagagt cctcatacca caactaaaat tttgaaggga 1740
acaaccagaa agattgcaag ctgaatcaag ttcatttttc caggaattta gtcttaggga 1800
tgtgataaat gtatttaggt gggattattc cagtctcacc atgttttttt ctttttttct 1860
cctctctttt ttggttatta cttataatta taaatggccg taaggcaaaa taaatgtata 1920
tttgatcatt ggaattttga ttctgcataa tcatatcttg gatgctactt tactaaaaaa 1980
aaaaaaaaaa aaaaa 1995
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Phe Phe Ser Thr Pro His Thr His Pro Pro Pro Pro Leu Phe Phe Thr
1 5 10 15
Leu Ser Thr Ile Ile Asn Pro Gln Leu Leu Lys Pro Leu Leu Leu Leu
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Met Asp Ser Ser Phe Thr Thr Ile Cys Phe Ser Ile Ala Leu Leu Leu
35 40 45
Phe Leu Ile His Phe Leu Leu Lys Leu Phe Ser Phe Phe Ser Ala Ser
50 55 60
Thr Ala Arg Lys Leu Pro Leu Pro Pro Gly Ser Leu Gly Trp Pro Tyr
65 70 75 80
Ile Gly Glu Thr Phe Gln Leu Tyr Ser Gln Asn Pro Asn Val Phe Phe
85 90 95
Ala Ser Lys Gln Glu Arg Tyr Gly Thr Ile Phe Lys Thr Arg Ile Leu
100 105 110
Gly Cys Pro Cys Val Met Ile Ser Ser Pro Glu Ala Ala Lys Leu Val
115 120 125
Leu Val Thr Lys Ala Pro Leu Phe Lys Pro Thr Phe Pro Ala Ser Lys
130 135 140
Glu Arg Met Leu Gly Lys Gln Ala Ile Phe Phe His Gln Gly Asp Tyr
145 150 155 160
His Ala Lys Leu Arg Lys Leu Val Leu Arg Ala Phe Met Pro Gly Ala
165 170 175
Ile Lys Asn Ile Val Ser Asn Ile Asp Ser Ile Ala Thr Gln Thr Leu
180 185 190
Gln Ser Trp Glu Gly Arg Ser Ile Asn Thr Phe Gln Glu Met Lys Thr
195 200 205
Tyr Thr Phe Asn Val Ala Leu Leu Ser Ile Phe Gly Lys Asp Glu Ile
210 215 220
Leu Tyr Arg Glu Glu Leu Lys Arg Cys Tyr Tyr Ile Leu Glu Lys Gly
225 230 235 240
Tyr Asn Ser Met Pro Ile Asn Ile Pro Gly Thr Leu Phe Asn Lys Ser
245 250 255
Met Lys Ala Arg Lys Glu Leu Ala Gln Ile Leu Ala Lys Ile Leu Ser
260 265 270
Thr Arg Arg Gln Ala Asn Gly Asp His Asn Asp Leu Leu Gly Ser Phe
275 280 285
Met Gly Asp Lys Glu Gly Leu Thr Asp Glu Gln Ile Ala Asp Asn Ile
290 295 300
Ile Gly Val Ile Phe Ala Ala Arg Asp Thr Thr Ala Ser Val Leu Thr
305 310 315 320
Trp Ile Leu Lys Tyr Leu Gly Glu Asn Pro Ser Val Leu Gln Ala Val
325 330 335
Thr Glu Glu Gln Glu Ser Ile Met Arg Ser Lys Glu Asp Lys Cys Gly
340 345 350
Glu Glu Lys Ser Leu Thr Trp Glu Asp Ala Lys Lys Met Pro Ile Thr
355 360 365
Ser Arg Val Ile Gln Glu Thr Leu Arg Ile Ala Ser Ile Leu Ser Phe
370 375 380
Thr Phe Arg Glu Ala Val Glu Asp Val Glu Phe Glu Gly Tyr Leu Ile
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Pro Lys Gly Trp Lys Val Leu Pro Leu Phe Arg Asn Ile His His Ser
405 410 415
Pro Asp Ile Tyr Pro Glu Pro Glu Lys Phe Asp Pro Ser Arg Phe Glu
420 425 430
Val Ala Pro Lys Pro Asn Thr Phe Met Pro Phe Gly Asn Gly Val His
435 440 445
Ser Cys Pro Gly Asn Glu Leu Ala Lys Leu Glu Ile Leu Val Leu Leu
450 455 460
His His Leu Thr Thr Lys Tyr Arg Trp Ser Met Val Gly Thr Gln Asn
465 470 475 480
Gly Ile Gln Tyr Gly Pro Phe Ala Leu Pro Gln Asn Gly Leu Pro Ile
485 490 495
Thr Ile Ser Leu Lys Ser Lys Gln Ile Ser Thr Lys Ser Thr Leu Ser
500 505 510
Ala Thr Asn Gln Leu His Ser His Ile Pro Ile Gln Ser Phe Lys Gln
515 520 525
Glu Thr Lys Lys Gln Tyr Glu Glu Ile Ser Leu Val Ser Cys Gln Ser
530 535 540
Arg Asp Thr His Met Lys Glu Phe Ser Ile Leu Lys Lys Gly Thr Lys
545 550 555 560
Lys Leu Tyr Ser Lys Ser Pro His Thr Thr Thr Lys Ile Leu Lys Gly
565 570 575
Thr Thr Arg Lys Ile Ala Ser Ile Lys Phe Ile Phe Pro Gly Ile Ser
580 585 590
Gly Cys Asp Lys Cys Ile Val Gly Leu Phe Gln Ser His His Val Phe
595 600 605
Phe Phe Phe Ser Pro Leu Phe Phe Gly Tyr Tyr Leu Leu Met Ala Val
610 615 620
Arg Gln Asn Lys Cys Ile Phe Asp His Trp Asn Phe Asp Ser Ala Ser
625 630 635 640
Tyr Leu Gly Cys Tyr Phe Thr Lys Lys Lys Lys Lys Lys Lys
645 650
<210> 3
<211> 26
<212> DNA
<213> 人工序列
<400> 3
atgggttggc cttayatagg agaaac 26
<210> 4
<211> 26
<212> DNA
<213> 人工序列
<400> 4
actttccacc cttttggtat aagata 26
<210> 5
<211> 31
<212> DNA
<213> 人工序列
<400> 5
caagctgtca cggaagagca agagtccata a 31
<210> 6
<211> 28
<212> DNA
<213> 人工序列
<400> 6
tttcttctcc actcctcaca cccatcct 28
<210> 7
<211> 25
<212> DNA
<213> 人工序列
<400> 7
gcctttgtga ccagcaccag ctttg 25
Claims (8)
1.一种葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白,其特征在于:该蛋白的氨基酸序列如SEQ ID No.2所示。
2.一种权利要求1所述的葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白的编码基因,其特征在于:该编码基因的序列如SEQ ID No.1所示。
3.根据权利要求2所述的葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白的编码基因,其特征在于:该基因是从葡萄新品种“鄞红”葡萄的果实筛选获得。
4.一种含有权利要求2-3任一权利要求所述的葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白编码基因的表达盒、重组表达载体、转基因细胞系或重组菌。
5.一种葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A蛋白、编码基因、表达盒、重组表达载体、转基因细胞系或重组菌在葡萄育种中的应用。
6.一种葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A基因的制备方法,其特征在于:该方法通过葡萄果实CYP707A基因的克隆获得,具体的包括:(1)植物材料的获得,(2)RNA和DNA的提取,(3)基因的全长克隆。
7.根据权利要求6所述的葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A基因的制备方法,其特征在于:所述的植物材料是采用鄞红”葡萄的果实。
8.根据权利要求6所述的葡萄脱落酸降解途径关键酶8’-羟化酶CYP707A基因的制备方法,其特征在于:所述的基因的全长克隆过程中使用了简并引物,该引物序列如下:F-1-CYP707A:ATGGGTTGGCCTTAYATAGGAGAAAC;R-4-CYP707A:ACTTTCCACCCTTTTGGTATAAGATA;使用了PCR扩增引物,该引物序列如下:3'-C-5:CAAGCTGTCACGGAAGAGCAAGAGTCCATAA;使用了葡萄全基因组所需引物,该引物的序列如下:F-CYP707A-D2:TTTCTTCTCCACTCCTCACACCCATCCT;R-CYP707A-D2:GCCTTTGTGACCAGCACCAGCTTTG。
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