CN1206358C - 羟苯丙酮酸双加氧酶基因的dna序列以及含有羟苯丙酮酸双加氧酶基因的抗特定除草剂植物的获得 - Google Patents

羟苯丙酮酸双加氧酶基因的dna序列以及含有羟苯丙酮酸双加氧酶基因的抗特定除草剂植物的获得 Download PDF

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CN1206358C
CN1206358C CNB96195857XA CN96195857A CN1206358C CN 1206358 C CN1206358 C CN 1206358C CN B96195857X A CNB96195857X A CN B96195857XA CN 96195857 A CN96195857 A CN 96195857A CN 1206358 C CN1206358 C CN 1206358C
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阿兰·赛兰德
安妮·罗兰
米歇尔·马特林格
肯·帕利特
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Abstract

羟苯丙酮酸双加氧酶基因的DNA序列以及含有羟苯丙酮酸双加氧酶基因的耐除草剂植物的生产。羟苯丙酮酸双加氧酶基因的DNA序列；自细菌或植物分离；用于获得耐除草剂植物。

Description

羟苯丙酮酸双加氧酶基因的DNA序列 以及含有羟苯丙酮酸双加氧酶 基因的抗特定除草剂 植物的获得
本发明涉及一个羟苯丙酮酸双加氧酶(HPPD)基因，一个含有此作为编码序列的此基因的嵌合基因及把它用于获得抗特定除草剂的植物。
特定的除草剂已经公开，例如特别是描述于法国专利申请95 06800和95 13570中的异噁唑，并且尤其是氟异噁唑(isoxaflutole)，一种选择性玉米除草剂，例如那些描述于欧洲专利申请0 496 630，0 496 631中的二酮腈(diketonitriles)，特别是2-氰基-3-环丙基-1-(2-SO2CH3-4-CF3-苯基)丙烷-1，3-二酮[2-cyano-3-cyclopropyl-1-(2-SO2CH3-4-CF3-phenyl)propane-1，3-dione]和2-氰基-3-环丙基-1-(2-SO2CH3-4-2，3-Cl2苯基)丙烷-1，3-二酮[2-cyano-3-cyclopropyl-1-(2-SO2CH3-4-2，3-Cl2 phenyl)propane-1，3-dione]，描述于欧洲专利申请0 625 505和0 625 508中的三酮，特别是硫三酮(sulcotrione)。但是，并没有记述对这些除草剂的耐受性基因。
羟基苯丙酮酸双加氧酶是一种催化对羟基苯丙酮酸到尿黑酸的转化反应的酶。
此外，描述了来源于假单胞菌菌株PJ.874(Pseudomonas sp.P.J.874)的羟基苯丙酮酸双加氧酶的氨基酸序列，但并未描述它在植物对除草剂耐受性方面的作用(Ruetschi等，“欧洲生物化学杂志”(Eur.J.Biochem.)205，459-466，1992)。本文没有给出编码此蛋白的基因的描述。
现在已经发现这种基因的序列，并且一旦***到植物细胞中，此类序列便能在植物中产生过量表达或HPPD的激活，给后者以很好的对特定的新除草剂，例如那些异噁唑族或三酮类除草剂的耐受性。
本发明的一个目的是一种分离的非人类来源的和非海洋细菌来源的基因的，或者一种植物基因的DNA序列，或者一种能与此分离的序列杂交的序列，特征在于它表达羟基苯丙酮酸双加氧酶(HPPD)。
更特殊地，此序列可以是细菌来源，例如特别是假单胞菌属或者是植物来源，例如特别是单子叶或双子叶植物，特别是拟南芥属(Arabidopsis)或者伞形科，例如，举例为胡萝卜(野胡萝卜(Daucus carotta))。它可以是天然的或野生型的或可能是突变的但同时基本保持对HPPD抑制剂的耐除草剂性，例如异噁唑族或三酮类除草剂。
发明也涉及分离上述基因的流程，其特征在于：
-从HPPD氨基酸序列中选择一些寡聚核苷酸作为引物
-从这些引物出发，通过PCR合成扩增片段
-通过构建并筛选基因组库分离该基因以及
-克隆该基因
优选地使用来自假单胞菌属细菌HPPD序列的引物。特别优选地，它们来源于荧光假单胞菌(PSeudomonas fluorescens)。
本发明也涉及编码HPPD的基因在转化植物的技术流程中的应用，可以用作标记基因或用作编码序列提供给植物对特定除草剂的耐受性。它也可以与其它标记基因和/或编码序列联合，用作一种农学性质。
编码基因可以是任何来源，天然的或野生型的或可能是突变的，但同时基本保留对HPPD抑制剂的除草剂耐受性的属性，例如异噁唑家族或三酮类除草剂。可以用作编码序列，特别是可以使用那些依据本发明的例如上述的编码基因。
植物细胞的转化可通过任何合适的已知方法完成。一序列的方法在于用偶合有DNA序列的微粒轰击细胞或原生质体。
另一序列的方法在于使用***到根癌土壤杆菌的Ti质粒(Agrobacteriumtumefaciens)或发根病土壤杆菌(Agrobacterium rhizogenes)的Ri质粒中的嵌合基因作为转移到植物中的工具。
本发明的另一个目的物也是嵌合基因，它在转录方向上包括至少一个启动子调节序列，一个表达羟基苯丙酮酸双加氧酶的异源编码序列以及至少一个终止子或聚腺苷酸化调节序列。
使用的启动子调节序列可以是在植物中天然表达基因的任何启动子序列，特别是来源于细菌，病毒或植物的启动子，诸如例如来自核酮糖-1，5-二磷酸羧化酶-加氧酶(RuBisCO)小亚单位基因或α-微管蛋白基因(欧洲专利申请EP No.0 652 286)，或者来源于植物病毒基因，诸如例为来自花椰菜花叶病毒(CAMV 19S或35S)，但任何合适的启动子均可使用。优选地借助于促进编码序列过量表达的启动子调节序列，诸如例为包括至少一种例如描述于欧洲专利申请EP 0507698中的组蛋白启动子。
依据本发明，同样可能与启动子调节序列联合使用其它位于启动子和编码序列之间的调节序列，例如“增强子”转录激活物，诸如例为描述于申请WO 87/07644中的烟草蚀纹病毒(TEV)翻译激活物，或者来源于单或双转运肽，并且在这一情况下可能被一中间序列隔开，也就是说，在转录方向上包括编码质体定位酶的某植物基因的编码转运肽的序列，编码质体定位酶的某植物基因的N-末端成熟部分的部分序列，然后是编码质体定位酶的某植物基因的编码第二个转运肽的序列，由编码质体定位酶的某植物基因N-末端成熟部分的一部分序列形成，例如欧洲专利申请No.0 508 909中所述。
可能使用任何相应的来源于细菌(诸如举例为根癌土壤杆菌的nos终止子)或甚至来源于植物(诸如举例为描述于欧洲专利申请EP No.0 633 317中的组蛋白终止子)的序列作为终止子或聚腺苷酸化调节序列。
本发明的又一目的物单子叶或双子叶植物的植物细胞，尤其是农作物细胞，它们按照上述的一种流程转化并在其基因组中含有有效量的表达羟基苯丙酮酸双加氧酶(HPPD)的基因。已经观察到用这种方法转化的植物对特定的新除草剂有显著耐受性，这些新除草剂如特别是描述于法国专利申请9506800和95 13570的异噁唑类以及特别地属于4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑{4-[4-CF3-2-(methyl sulphonyl)benzoly]-5-cyclopropylisoxazole}的异噁唑类，以及特别地是氟异噁唑，一种选择性玉米除草剂，双酮腈类(diketonitriles)，例如描述于欧洲专利申请0496 630，0 496 631中的那些，特别是2-氰基-3-环丙基-1-(2-SO2CH3-4-CF3-苯基)丙烷-1，3-二酮[2-cyano-3-cyclopropyl-1-(2-SO2CH3-4-CF3-phenyl)propane-1，3-dione]和2-氰基-3-环丙基-1-(2-SO2CH3-4-2，3-Cl2苯基)丙烷-1，3-二酮[2-cyano-3-cyclopropyl-1-(2-SO2CH3-4-2，3-Cl2-phenyl)propane-1，3-dione]，描述于欧洲专利申请0 625 505和0 625 508中的三酮类，特别是硫三酮。
最后，本发明的目的是借助此类型除草剂来清除植物，特别是农作物杂草的一种方法，特征在于在农作物播种前，萌发前以及萌发后都把除草剂施用于按照发明转化的植物。
本发明还有一个目的是把HPPD基因作为标记基因用于植物物种“转化-再生”周期过程中并在上述除草剂基础上筛选。
借助在下面的实验实施例，发明的各方面特征将得以更好的理解。
实施例1：荧光假单胞菌A32(P.fluore scens A32)的HPPD基因的分离
从假单胞菌菌株PJ.874的HPPD氨基酸序列出发(RuetschiU.等发表，1992，“欧洲生物化学杂志”(Eur.J.Biochem.)205：459-466，推导出不同的寡聚核苷酸序列以通过PCR扩增荧光假单胞菌A32(P.fluorescens A32)(由McKellar，R.C.分离，1982，“应用细菌学杂志”(J.Appl.Bacteriol.)，53：305-316)的HPPD编码序列的一部分。已应用此HPPD基因的一个扩增片段来筛选荧光假单胞菌A32的部分基因组库并从而分离编码此酶的基因。
A)荧光假单胞菌A32基因组DNA的制备
细菌在40毫升M63基本培养基(KH2PO4 13.6克/升，(NH4)2SO4 2克/升，MgSO4 0.2克/升，FeSO4 0.005克/升，pH7，添加10mM L-酪氨酸作为单一碳源)中于28℃下培养48小时。
洗涤后，细胞溶解于1毫升裂解缓冲液(100mM tris HCl，pH8.3，1.4MNaCl和10mM乙二胺四乙酸钠(EDTA))并在65℃下温育10分钟。
酚/氯仿(24∶1)处理和氯仿处理后，通过加入一体积异丙醇沉淀核酸，然后溶解于300微升无菌水并用终浓度10微克/毫升的RNA酶处理。DNA重新用酚/氯仿，氯仿处理并通过加入1/10体积3M醋酸钠，pH5和2体积乙醇再沉淀。DNA然后溶解于无菌水并检测。
B)寡聚核苷酸的选择以及合成
从假单胞菌菌株PJ.874的HPPD氨基酸序列出发，选择五段寡聚核苷酸，其中两段定向为蛋白质NH2末端指向蛋白质COOH末端的方向，另三段定向相反方向(参看图1)。选择按下列标准决定：
-寡聚核苷酸有稳定3′末端，也就是说至少两个碱基没有歧义。
一可能的简并性最小
选择的寡聚核苷酸具有下面的序列：
P1：5′TA(C/T)GA(G/A)AA(C/T)CCTATGGG3′
P2：5′GA(G/A)ACIGGICCIATGGA3′
P3：5′AA(C/T)TGCATIA(G/A)(G/A)AA(C/T)TC(C/T)TC3′
P4：5′AAIGCIAC(G/A)TG(C/T)TG(T/G/A)ATICC3′
P5：5′GC(C/T)TT(A/G)AA(A/G)TTICC(C/T)TCIC3′
它们在MILLIPORE制造的Cyclone plus DNA合成仪中合成。
使用这五种寡聚核苷酸，通过起始于序列1的PCR，理论上应该得到的扩增片段具有下列大小：
使用引物P1和P3  ----------→
大约690碱基对(bp)
使用引物P1和P4  ----------→
大约720bp
使用引物P1和P5  ----------→
大约1000bp
使用引物P2和P3  ----------→
大约390bp
使用引物P2和P4  ----------→
大约420bp
使用引物P2和P5  ----------→
大约700bp
C)荧光假单胞菌A32的HPPD某编码部分的扩增
扩增在PEPKIN ELMER 9600 PCR仪中进行并在标准条件下使用PERKIN ELMER的Taq聚合物酶及其缓冲液，也就是说对50微升反应混合液，含有200μMdNTP，20μM引物，2.5单位Taq聚合酶和2.5微克荧光假单胞菌A32的DNA。
使用的扩增程序是95℃5分钟，然后＜45秒95℃，45秒49℃，1分钟72℃＞进行35个循环，随后是72℃5分钟。
在这种条件下，所有获得的扩增片段大小与上述给出的理论大小相符，从而很好的表明扩增特异性。
用各对引物P1/P4，P1/P5和P2/P4获得的扩增片段，在用Eco RV消化质粒pBSII SK(-)并如HOLTONT.A.和GRAHAMM.W.1991，N.A.R.，Vol.19，No.5，p.1156所述在dd TTP存在下用末端转移酶处理后，连接到pBSII SK(-)中。
对三种类型每一种的一个克隆进行了部分测序；这使得可以证实在三种情况下荧光假单胞菌A32的HPPD部分编码区真正得到了扩增。P1/P4片段保留作为探针以筛选荧光假单胞菌A32部分基因组库并分离完整的HPPD基因。
D)基因的分离
通过Southern检测表明，用限制性酶Bam HI消化荧光假单胞菌A32DNA后的7Kbp片段与探针HPPD P1/P4杂交。400微克荧光假单胞菌A32DNA从而用限制性酶Bam HI消化并且出等于大约7Kbp的DNA片段在琼脂糖凝胶中纯化。
这些片段连接到pBSII SK(-)中，后者已用Bam HI消化并用碱性磷酸酶处理去磷酸化。转化到大肠杆菌DH 10b(E.coli DH10b)后，用探针HPPD P1/P4筛选部分基因组库。
筛选到一个阳性克隆并命名为pRP A。它的简化图谱示于图2。此图中标出了HPPD基因编码部分的位置。它由编码358个氨基酸的1077个核苷酸组成(参看序列1)。荧光假单胞菌A32的HPPD与假单胞菌菌株PJ.874具有很好的氨基酸同源性，实际上两蛋白92％一致(参见图3)。
实施例2：两个嵌合基因的构建
为使植物有对抑制HPPD的除草剂的耐受性，构建了两个嵌合基因：
第一个嵌合基因在于把荧光假单胞菌A32 HPPD基因的编码部分置于双组蛋白启动子(欧洲专利No.0 507 698)控制之下，其后是烟草蚀纹病毒翻译增强子(TEV)(pRTL-GUS(Carrington和Freed，1990；“病毒学杂志”(J.Virol)64：1590-1597))以及胭脂氨酸合成酶基因的终止子。HPPD然后将定位到细胞质。
第二个嵌合基因除了把最优化转运肽(OTP)***到TEV转录激活物和HPPD编码部分之间外(欧洲专利申请EP No.0508 909)，几乎与第一个相同。HPPD然后将定位于叶绿体。
A)载体pRPA-RD-153的构建
-pRPA-RD-11：含有胭脂氨酸合成酶聚腺苷酸化位点(NOS polyA)(欧洲专利申请EP No.0652 286)的pBSII SK(-)(Stratagene目录#212206)衍生物克隆到KpmI和SalI位点之间。KpnI位点在150μM脱氧核苷三磷酸存在下用T4DNA聚合酶I处理转化到NotI位点，然后用一个NotI接头连接(Stratagene目录#1029)。从而获得了一个NOS polyA克隆盒。
-pRPA-RD-127：一个pRPA-BL-466(欧洲专利申请EP No.0337 899)的衍生物克隆到pRPA-RD-11，产生oxy基因的表达盒并含有核酮糖-1，5-二磷酸羧化酶-加氧酶小亚单位启动子：
“启动子(SSU)-oxy基因-NOS poly A”
为产生此质粒，用XbaI和HindIII消化pRPA-BL-448以分离含有SSU启动子和oxy基因的1.9Kbp片段，它再连接到用兼容酶消化的质粒pRPA-RD-11中。
-pRPA-RD-132：这是克隆到pRPA-RD-127产生一个具有双组蛋白启动子的oxy基因表达盒pRPA-BL-488(欧洲专利申请EPNo.0507 698)的一个衍生物：
“双组蛋白启动子-oxy基因-NOS poly A”
为产生此质粒，用HindIII消化pRPA-BL-466，用Klenow酶处理然后用NcoI再消化。纯化的含有组蛋白双启动子H3A 748的1.35Kbp片段与用XbaI消化，Klenow处理NcoI再消化的质粒pRPA-RD-127相连接。
-pRPA-RD-153：这是一个含有烟草蚀纹病毒(TEV)的pRPA-RD-132衍生物。用NcoI和EcoRI消化pRTL-GUS(Carrington和Freed，1990；“病毒学杂志”(J.Virol.，64：1590-1597)并把150bp片段连接到相同酶消化的pRPA-RD-132中。从而产生了含有启动子“双组蛋白启动子-TEV-oxy基因-NOS poly A”的表达盒。
B)载体pRPA-RD-185的构建：
pUC19/GECA：含有多克隆位点的PUC19(Gibco目录#15364-011)的衍生物。用EcoRI消化pUC-19并与寡聚核苷酸接头1相连接：
接头1：AATTGGGCCA GTCAGGCCGT TTAAACCCTA
GGGGGCCCG CCCGGT CAGTCCGGCA AATTTGGGAT
CCCCCGGGC TTAA
选择的克隆含有EcoRI位点，其后是含有下列位点的多接头：EcoRI，ApaI，AvrII，PmeI，SfiI，SacI，KpnI，SmaI，BamHI，XbaI，SalI，PstI，SphI和HindIII。
pRPA-RD-185：这是一个含有修饰多接头的pUC19/GECA衍生物。用HindIII消化pUC19/GECA并与寡核者酸接头2相连接：
接头2：AGCTTTTAAT TAAGGCGCGC CCTCGAGCCT
GGTTCAGGG AAATTA ATTCCGCGCG GGAGCTCGGA
CCAAGTCCC TCGA
选择的克隆在多接头中间含有一个HindIII位点，多接头现在含有下列位点：EcoRI，ApaI，AvrII，PmeI，SfiI，SacI，KpnI，SmaI，BamHI，XbaI，SalI，PstI，SphI，HindIII，PacI，AscI，XhoI和EcoNI。
C)载体pRP T的构建：
-pRP O：一个含有HPPD表达盒，双组蛋白启动子-TEV-HPPD基因-终止子Nos的pRPA-RD-153衍生物。为了产生pRP O，用HindIII消化pRPA-RD-153，Klenow处理，然后用NcoI再消化以去除oxy基因，并使用来自以BstEII消化，Klenow处理以及NcoI再消化的pRPA质粒的HPPD基因来代替它。
-pRP R：为获得此质粒，用PvuII和SacI消化pRP O，纯化嵌合基因并连接到pRPA-RD-185中，且后者已用PvuII和SacI消化。
-pRP T：通过把来源于SacI和HindIII消化的pRP R的嵌合基因连接到用相同酶消化的质粒pRPA-BL 150 alpha2(欧洲专利申请EP No.0508 909)中获得。
pRP T载体的嵌合基因从而具有下列结构：
双组蛋白启动子 TEV  HPPD编码区 nos终止子
D)pRP V载体的构建：
-pRP P：这是含有最优化的转运肽且其后是HPPD基因pRPA-RD-7(欧洲专利申请EP No.0 652 286)的衍生物。它通过连接来自用BstEII和NcoI消化，Klenow处理的pRPA和来自质粒pRPA-RD-7的HPPD编码部分获得，后者用SphI和AccI消化并用DNA酶聚合酶T4处理。
-pRP Q：含有HPPD表达盒，双组蛋白启动子-TEV-OTP-HPPD基因-Nos终止子的pRPA-RD-153的衍生物。为构建它，用SalI消化质粒pRPA-RD-153，Klenow处理然后用NcoI再消化去除oxy基因并用由BstEII消化，Klenow处理并用NcoI再消化的DRP P质粒释放的HPPD基因代替它。
-pRP S：为获得它，有PvuII和SacI消化质粒pRP Q以释放嵌合基因并连接到pRPA-RD-185，后者已用PvuII和SacI消化。
-pRP V：它通过把SacI和HindIII消化后从pRP S释放的嵌合基因连接到质粒pRPA-BL 150alpha2(欧洲专利申请EP No.0508 909)中获得。pRP V载体的嵌合基因从而含有以下结构：
双组蛋白启动子 TEV  OTP  HPPD编码区 nos终止子
实施例3：工业用烟草PBD6的转化
为了检测这两种嵌合基因的效率，根据欧洲专利申请EP No.0 508 909中所述的转化和再生工艺把它们转移到工业用烟草PBD6中。
1)转化：
把载体引入到携带粘粒pTVK291(Komari等，1986)的非致癌菌株土壤杆菌EHA 101中(Hood等，1987)。转化技术基于Horsh R.等(1985)，“科学”(Science)，227，1229-1231中的流程。
烟草PBD6(来自SEITA，法国)从叶外植体的再生在含有30克/升蔗糖和200微克/毫升卡那霉素的Murashige和Skoog(MS)基础培养基中进行。叶外植体从温室或试管植物上选择并按照叶盘法(“科学”(Science)1985，Vol，227，P.1229-1231)通过三个连续步骤转化：第一步包括在加有30克/升蔗糖并含有0.05毫克/升萘乙酸(NAA)和2毫克/升苄基氨基嘌呤(BAP)的MS培养基诱导苗15夭。这一步形成的苗通过在加有30克/升蔗糖但不含任何激素的MS培养基中培养10天进行发育。选择发育的苗并在含有含量为一半的盐，维生素和糖并不含任何激素的MS生根培养基中培养。在接近15夭时，把生根的苗置于土壤。
实施例4：检测烟草对4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑的耐受性：萌发后处理。
在离开离体培养后，转化烟草小植株在温室中(60％相对湿度；温度：夜晚20℃，白天23℃)驯化五周，然后用4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑处理。
未转化的对照烟草用剂量范围50到400克/公顷(g/ha)的4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑处理，在大约72小时内发生黄化，在一周内增强至发展成显著的枯斑(将最后的叶覆盖大约80％)。
转化后过量表达荧光假单胞菌HPPD的同一种烟草被很好地保护起来，抗400克/公顷剂量的4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑处理。
如果过量表达的酶在细胞质中，也就是说如果转化通过载体pRP T携带的基因完成，那么植物显示全部位于中间叶的非常轻微的黄化。
如果过量表达的酶在叶绿体中，也就是说如果转化通过载体pRPV携带的基因完成，那么植物被非常好地保护起来并不显示任何症状。实施例5：检测烟草对4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑的耐受性：萌发前处理。
a)体外试验
在实施例1到3中所述的400克/公顷的剂量时，使用收获于来自“转化-再生”周期并对氟异噁唑叶处理有抗性的植株的烟草种子。
把这些种子播种到有10克/升的植物琼脂以及从0到1毫克/升范围内的不同浓度氟异噁唑的盒子中。发芽在25℃下12小时光照/12小时黑暗的光周期中进行。
根据这一方案，萌发野生型烟草种子以及两种转基因烟草的种子，也就是说HPPD定位于细胞质的CY烟草和HPPD定位于叶绿体的CO烟草的种子。
播种2和3周后以目测进行抗性测定。
结果记录于下表中。
  氟异唑的浓度 野生型烟草     CY烟草   CO烟草
  0mg/l 100％种子萌发，无症状°     100％种子萌发，无症状°   100％种子萌发，无症状
  0.05mg/l 20％种子萌发*，表现出症状°     75％种子萌发*，无症状°   75％种子萌发*，无症状°
  0.1mg/l 不萌发     75％种子萌发*，无症状°   75％种子萌发*，无症状°
  0.5mg/l 不萌发     75％种子萌发*，无症状°   75％种子萌发*，无症状°
  1mg/l 不萌发     75％种子萌发*，有轻微症状°   75％种子萌发*，无症状°
°小植物在发芽过程中表现的症状是或多或少的子叶的显著畸变，并首先是正常进行光合作用并从而是绿色的组织变白。
*75％的种子发芽是因为种子来自“转化-再生”周期并从而只在一条染色体上携带耐受性基因的单基因座植物的自体受精。
用同样的方法以下面的产品进行测定；
使用美国专利4 780 127 No.51产品在0毫克/升和0.1毫克/升浓度下处理野生型烟草和CO烟草获得相同的结果。
b)温室试验：
除处理要在播种前24小时，萌发前进行外，检测如实施例4中所述进行。田间播种按常规进行。在这些条件下观察到，对于未处理的对照播种，在至少等于10克/公顷的任何浓度除草剂处理后未出现发芽。与此相反，CY烟草在高至并包括100克/公顷时不表现如段落a)中定义的任何症状。相似地，CO烟草在高至并包括200克/公顷时不表现如段落a)中定义的任何症状。
这些结果清楚地表明荧光假单胞菌HPPD基因使烟草有对氟异噁唑萌发前处理的耐受性。如果蛋白定位到叶绿体而不是细胞质时，耐受性更好。
实施例6：
为了研究是否荧光假单胞菌HPPD基因在植物物种“转化-再生”周期过程中可用作标记基因，用HPPD基因转化烟草并在氟异  唑筛选后获得转化的植株。
材料、方法和结果
按照欧洲专利申请EP No.0 508 909中所述的转化和再生方法，把描述于下面的嵌合基因pRP V转移到工业用烟草pBD6中。
载体pRP V的嵌合基因有下列结构：
双组蛋白启动子 TEV  OTP  HPPD编码区 nos终止子
1)转化：
把载体引入到携带粘粒pTVK 291(Komari等，1986)的土壤杆菌EHA101非致癌菌株(Hood等，1987)中。转化技术以Horsh等的方法(1985)为基础。
2)再生：
烟草PBD6(来自SEITA，法国)从叶外植体的再生在含有30克/升蔗糖以及350毫克/升氨噻肟酯头孢菌素(cefotaxime)和1毫克/升氟异噁唑的Murashige和Skoog(MS)基础培养基中进行。叶外植体选自温室或试管植物并按照叶盘法(“科学”(Science)1985，Vol.227，P.1229-1231)通过三个连续步骤转化：第一步包括在添加30克/升蔗糖且包含有0.05毫克/升萘乙酸(NAA)和2毫克/升苄基氨基嘌呤(BAP)及1毫升/升氟异噁唑的MS培养基中诱导苗生长15天。这一步骤中形成的绿苗然后通过在添加30克/升蔗糖和1毫克/升氟异噁唑但不含激素的MS培养基中培养10天进行发育。选择发育的苗并在含量为一半的盐，维生素和糖和1毫克/升氟异噁唑并不含任何激素的MS生根培养基中培养。大约15夭后，把生根的苗移至土壤。
使用荧光假单胞菌HPPD的特异引物通过PCR检测了根据这种方法获得的所有小植物。PCR检测使得可以证实这样获得的所有小植物整合了HPPD基因。
总之，此检测证实HPPD基因可用作标记基因并且与此基因相联系，氟异噁唑可作为良好的选择剂。
实施例7和8：分离拟南芥(Arabidopsis thaliana)的HPPD基因和胡萝卜(野胡萝卜(Daucus carotta))的HPPD基因
a)cDNA库构建
从拟南芥幼小植株提取的mRNAs和从培养的胡萝卜细胞提取的mRNAs通过从Stratagen公司购买的载体Uni ZapTMXR；按照该公司推荐的方法用于构建两个cDNA库。
b)筛选cDNA库
使用通过拟南芥生物资源中心(Arabidopsis Biological ResourceCenter)(俄亥俄州，美国)获得的并检索为：EST clone No.91B 13T17的部分长度的拟南芥cDNA相对应的探针筛选这两个库。这一克隆由大约500碱基对组成，其中只有228碱基对已由MSU-DOE植物研究实验室(MSU-DOEPlant Research Laboratory)测序于拟南芥cDNA随机测序序列中。通过裂解区域杂交的经典技术，我们在使用这个克隆筛选拟南芥和胡萝卜cNDA库之前，完全测定了500碱基对序列。
c)获得了一段对应1338碱基对的拟南芥cDNA(序列2)。此cDNA在25位有一个翻译起始密码子，在1336位有一个翻译终止密码子。此cDNA从而是完全的并编码一种445氨基酸的蛋白质。
d)获得了一段对应1329碱基对的胡萝卜(野胡萝卜(Daucus carotta))cDNA(序列3)。此cDNA在1位有一个翻译起始密码子，在1329位有一个翻译终止密码子。此cDNA从而是完全的并编码一个442氨基酸的蛋白质。
图示的序列如下：
序列1
荧光假单胞菌A32HPPD基因序列
序列2
拟南芥HPPD的cDNA序列
序列3
Daucus carotta HPPD的cDNA序列通过标示给出下列图形以说明本发明。
图1表示假单胞菌PJ.874 HPPD的蛋白质序列以及相应编码部分的理论核苷酸序列；五个箭头表示选择的用于扩增部分此编码区的五段寡聚核苷酸。
图2表示具有包含荧光假单胞菌A32HPPD基因的7kb基因组DNA片段的质粒图谱。
图3给出了荧光假单胞菌A32HPPD氨基酸序列与假单胞菌P.J.874HPPD氨基酸序列的比较(只标出了两个序列相异的氨基酸)以及共有序列。
                              序列表
(1)一般资料
(i)申请人：Sailland，Alain
   Rolland，Anne
   Matringe，Michel
   Palltt，Kenneth E
(ii)发明名称：
羟苯丙酮酸双加氧酶的DNA序列以及含有这个羟苯丙酮酸双加氧酶基因且对特定除草剂有抗性的植物的获得
(iii)序列数目：3
(iv)通讯地址：
    (A)收信人：Francois Chretien
    (B)街道：14-20 rue Pierre BAIZET
    (C)城市：Lyon Cedex 09
    (E)国家：法国
    (F)邮编：89263
(v)计算机可读形式：
    (A)媒介类型：软盘
    (B)计算机：IBM PC兼容
    (C)操作***：PC-DOS/MS-DOS
    (D)软件：PatentIn Release#1.0，#1.25版本
  (vi)现在申请数据：
    (A)申请号：FR PH95033
    (B)填表日期：1995年6月2日
    (C)分类：
  (viii)律师/代理人资料：
    (A)姓名：Chretien，Francois
  (ix)通讯资料
    (A)电话：72-29-26-42
    (B)传真：72-29-28-43
(2)序列1的资料：
  (i)序列特征：
    (A)长度：1077碱基对
    (B)类型：核酸
    (C)链型：双链
    (D)拓朴结构：线性
  (ii)分子类型：DNA(基因组)
  (iii)假设：无
  (iv)反义：无
  (vi)来源
    (A)生物：荧光假单胞菌(Pseudomonas fluorescens)
  (xi)序列描述：序列1：
ATGGCAGATC TATACGAAAA CCCAATGGGC CTGATGGGCT TTGAATTCAT CCAATTAGCG    60
TCGCCGACGC CCGCTACCCT GGAGCCGATC TTCGAGATCA TGGGCTTCAC CAAAGTCGCG   120
ACCCACCGTT CCAAGAACGT GCACCTGTAC CGCCAGGGCG AGATCAACCT GATCCTCAAC   180
AACGAGCCCA ACAGCATCGC CTCCTACTTT GCGGCCGAAC ACGGCCCGTC GGTGTGCGGC   240
ATGGCGTTCC GCGTGAAGGA CTCGCAAAAG GCCTACAACC GCGCCCTGGA ACTCGGCGCC   300
CAGCCGATCC ATATTGACAC CGGGCCGATG GAATTGAACC TGCCGGCGAT CAAGGGCATC   360
GGCGGCGCGC CGTTGTACCT GATCGACCGT TTCGGCGAAG GCAGCTCGAT CTACGACATC   420
GACTTCGTGT ACCTCGAAGG TGTGGAGCGC AATCCGGTCG GTGCAGGTCT CAAAGTCATC   480
GACCACCTGA CCCACAACGT CTATCGCGGC CGCATGGTCT ACTGGGCCAA CTTCTACGAG   540
AAATTGTTCA ACTTCCGTGA AGCGCGTTAC TTCGATATCA AGGGCGAGTA CACCGGCCTG    600
ACTTCCAAGG CCATGAGTGC GCCGGACGGC ATGATCCGCA TCCCGCTGAA CGAAGAGTCG    660
TCCAAGGGCG CGGGGCAGAT CGAAGAGTTC CTGATGCAGT TCAACGGCGA AGGCATCCAG    720
CACGTGGCGT TCCTCACCGA CGACCTGGTC AAGACCTGGG ACGCGTTGAA GAAAATCGGC    780
ATGCGCTTCA TGACCGCGCC GCCAGACACT TATTACGAAA TGCTCGAAGG CCGCCTGCCT    840
GACCACGGCG AGCCGGTGGA TCAACTGCAG GCACGCGGTA TCCTGCTGGA CGGATCTTCC    900
GTGGAAGGCG ACAAACGCCT GCTGCTGCAG ATCTTCTCGG AAACCCTGAT GGGCCCGGTG    960
TTCTTCGAAT TCATCCAGCG CAAGGGCGAC GATGGGTTTG GCGAGGGCAA CTTCAAGGCG   1020
CTGTTCGAGT CCATCGAACG TGACCAGGTG CGTCGTGGTG TATTGACCGC CGATTAA      1077
(2)序列2的资料：
(i)序列特征：
(A)长度：1338碱基对
(B)类型：核酸
(C)链型：双链
(D)拓朴结构：线性
(ii)分子类型：cDNA
(iii)假设：无
(iv)反义：无
(vi)来源
(A)生物：拟南芥(Arabidopsis thaliana)
(B)品系：Columbia
(D)发育阶段：初生绿色植株
(vii)直接来源：
(A)库：Uni zap XR STRATAGENE
(xi)序列描述：序列2：
ATGGGCCACC AAAACGCCGC CGTTTCAGAG AATCAAAACC ATGATGACGG CGCTGCGTCG    60
TCGCCGGGAT TCAAGCTCGT CGGATTTTCC AAGTTCGTAA GAAAGAATCC AAAGTCTGAT   120
AAATTCAAGG TTAAGCGCTT CCATCACATC GAGTTCTGGT GCGGCGACGC AACCAACGTC   180
GCTCGTCGCT TCTCCTGGGG TCTGGGGATG AGATTCTCCG CCAAATCCGA TCTTTCCACC   240
GGAAACATGG TTCACGCCTC TTACCTACTC ACCTCCGGTG ACCTCCGATT CCTTTTCACT   300
GCTCCTTACT CTCCGTCTCT CTCCGCCGGA GAGATTAAAC CGACAACCAC AGCTTCTATC   360
CCAAGTTTCG ATCACGGCTC TTGTCGTTCC TTCTTCTCTT CACATGGTCT CGGTGTTAGA   420
GCCGTTGCGA TTGAAGTAGA AGACGCAGAG TCAGCTTTCT CCATCAGTGT AGCTAATGGC   480
GCTATTCCTT CGTCGCCTCC TATCGTCCTC AATGAAGCAG TTACGATCGC TGAGGTTAAA    540
CTATACGGCG ATGTTGTTCT CCGATATGTT AGTTACAAAG CAGAAGATAC CGAAAAATCC    600
GAATTCTTGC CAGGGTTCGA GCGTGTAGAG GATGCGTCGT CGTTCCCATT GGATTATGGT    660
ATCCGGCGGC TTGACCACGC CGTGGGAAAC GTTCCTGAGC TTGGTCCGGC TTTAACTTAT    720
GTAGCGGGGT TCACTGGTTT TCACCAATTC GCAGAGTTCA CAGCAGACGA CGTTGGAACC    780
GCCGAGAGCG GTTTAAATTC AGCGGTCCTG GCTAGCAATG ATGAAATGGT TCTTCTACCG    840
ATTAACGAGC CAGTGCACGG AACAAAGAGG AAGAGTCAGA TTCAGACGTA TTTGGAACAT    900
AACGAAGGCG CAGGGCTACA ACATCTGGCT CTGATGAGTG AAGACATATT CAGGACCCTG    960
AGAGAGATGA GGAAGAGGAG CAGTATTGGA GGATTCGACT TCATGCCTTC TCCTCCGCCT   1020
ACTTACTACC AGAATCTCAA GAAACGGGTC GGCGACGTGC TCAGCGATGA TCAGATCAAG   1080
GAGTGTGAGG AATTAGGGAT TCTTGTAGAC AGAGATGATC AAGGGACGTT GCTTCAAATC   1140
TTCACAAAAC CACTAGGTGA CAGGCCGACG ATATTTATAG AGATAATCCA GAGAGTAGGA   1200
TGCATGATGA AAGATGAGGA AGGGAAGGCT TACCAGAGTG GAGGATGTGG TGGTTTTGGC   1260
AAAGGCAATT TCTCTGAGCT CTTCAAGTCC ATTGAAGAAT ACGAAAAGAC TCTTGAAGCC   1320
AAACAGTTAG TGGGATGA                                                 1338
(2)序列3的资料
  (i)序列特征：
    (A)长度：1329碱基对
    (B)类型：核酸
    (C)链型：双链
    (D)拓朴结构：线性
  (ii)分子类型：cDNA
  (iii)假设：无
  (iv)反义：无
  (vi)来源
    (A)生物：野胡萝卜(Daucus carota)
    (D)发育阶段：悬浮细胞
  (vii)直接来源：
    (A)库：Uni zap XR STRATAGENE
  (xi)序列描述：序列3
ATGGGGAAAA AACAATCGGA AGCTGAAATT CTCTCAAGCA ATTCATCAAA CACCTCTCCT    60
GCAACATTCA AGCTGGTCGG TTTCAACAAC TTCGTCCGCG CCAACCCCAA GTCCGATCAC    120
TTCGCCGTGA AGCGGTTCCA CCACATTGAG TTCTGGTGCG GCGACGCCAC CAACACGTCG    180
CGGCGGTTCT CGTGGGGCCT CGGCATGCCT TTGGTGGCGA AATCGGATCT CTCTACTGGA    240
AACTCTGTTC ACGCTTCTTA TCTTGTTCGC TCGGCGAATC TCAGTTTCGT CTTCACCGCT    300
CCTTACTCTC CGTCCACGAC CACTTCCTCT GGTTCAGCTG CCATCCCGTC TTTTTCGGCA    360
TCGGGTTTTC ACTCTTTTGC GGCCAAACAC GGCCTTGCTG TTCGGGCTAT TGCTCTTGAA    420
GTTGCTGACG TGGCTGCTGC GTTTGAGGCC AGTGTTGCGC GTGGGGCCAG GCCGGCTTCG    480
GCTCCTGTTG AATTGGACGA CCAGGCGTGG TTGGCTGAGG TGGAGTTGTA CGGAGATGTG    540
GTCTTGAGGT TTGTTAGTTT TGGGAGGGAG GAGGGTTTGT TTTTGCCTGG ATTCGAGGCG    600
GTGGAGGGGA CGGCGTCGTT TCCGGATTTG GATTATGGAA TTAGAAGACT TGATCATGCG    660
GTGGGGAATG TTACCGAGTT GGGGCCTGTG GTGGAGTATA TTAAAGGGTT TACGGGGTTT    720
CATGAATTTG CGGAGTTTAC AGCGGAGGAT GTGGGGACTT TGGAGAGTGG GTTGAATTCG    780
GTGGTGTTGG CGAATAATGA GGAGATGGTT CTGTTGCCCT TGAATGAGCC TGTGTATGGG    840
ACCAAGAGGA AGAGTCAGAT ACAGACTTAC TTGGAGCACA ATGAAGGGGC TGGAGTGCAG    900
CATTTGGCTT TAGTGAGTGA GGATATTTTT AGGACTTTAA GGGAGATGAG GAAGAGGAGT    960
TGCCTTGGTG GTTTTGAGTT TATGCCTTCG CCACCGCCTA CGTATTACAA GAATTTGAAG   1020
AATAGGGTCG GGGATGTGTT GAGTGATGAA CAGATCAAGG AGTGTGAAGA TTTGGGGATT   1080
TTGGTGGATA GGGATGATCA GGGTACATTG CTTCAAATCT TTACCAAGCC TGTAGGTGAC   1140
AGGCCTACCT TATTCATAGA GATCATTCAG AGGGTAGGGT GCATGCTCAA GGACGATGCA   1200
GGGCAGATGT ACCAGAAGGG CGGGTGCGGA GGATTTGGGA AGGGGAACTT CTCAGAGCTG   1260
TTCAAGTCCA TCGAAGAATA TGAAAAAACA CTTGAAGCTA AACAAATCAC TGGATCTGCT   1320
GCTGCATGA                                                           1329

Claims (20)

1.从荧光假单胞菌分离到的序列1所示的基因序列，其特征是表达羟苯丙酮酸双加氧酶。
2.用于植物遗传转化的嵌合基因，在转录方向上包括：
-至少一个来自在植物中天然表达自身的某基因的启动子调节序列，
-一个异源编码序列，
-至少一个聚腺苷酸化序列，
特征在于异源编码序列是一个表达羟苯丙酮酸双加氧酶基因的序列1。
3.根据权利要求2的嵌合基因，其特征是启动子调节序列有利于编码序列的过量表达。
4.根据权利要求3所述的嵌合基因，其特征是启动子调节序列至少包括一个组蛋白启动子。
5.根据权利要求2-4中任一项的嵌合基因，特征是在启动子调节序列和编码序列之间包含一个运转肽。
6.根据权利要求2的嵌合基因，特征是在启动子调节序列和编码序列之间包含一个最优化转运肽区，该最优化转运肽区在转录方向上包括编码质体定位酶的某植物基因的编码运转肽的序列，编码质体定位酶的某植物基因N末端成熟部分的序列的一部分，然后编码质体定位酶的某植物基因的编码第二个转运肽的序列。
7.根据权利要求2-4中任一项的一个嵌合基因，特征是它在启动子调节序列和编码序列之间包含一个增强子序列。
8.可用于植物遗传转化的载体，特征是它含有根据权利要求2-7任一所述的嵌合基因。
9.植物细胞，特征是它含有根据权利要求2-7任一所述的嵌合基因。
10.转化植物以使它们耐受羟苯丙酮酸双加氧酶抑制剂的方法，其特征是把一个表达外源羟苯丙酮酸双加氧酶的序列1所示的基因导入植物细胞。
11.根据权利要求10所述的方法，特征是使用根癌土壤杆菌或发根病土壤杆菌完成转移。
12.根据权利要求10所述的方法，特征是使用负载DNA的颗粒通过基因枪导入完成转移。
13.植物转化的方法，特征是把表达外源羟苯丙酮酸双加氧酶的序列1所示的基因作为选择标记导入植物细胞。
14.一种用植物选择性除草剂处理植物的方法，特征是把一个羟苯丙酮酸双加氧酶基因的抑制剂施加到从根据权利要求9所述的细胞再生的植株。
15.根据权利要求14所述的方法，特征是其中的植株属于双子叶类。
16.根据权利要求14的方法，特征是羟苯丙酮酸双加氧酶基因抑制剂是一种异噁唑。
17.根据权利要求16所述的方法，特征是异噁唑为4-[4-CF3-2-(甲磺酰)苯甲酰基]-5-环丙基异噁唑。
18.根据权利要求14的方法，特征是羟苯丙酮酸双加氧酶基因抑制剂是一种二酮腈。
19.根据权利要求14的方法，其特征是羟苯丙酮酸双加氧酶基因抑制剂是一种三酮。
20.根据权利要求14的方法，特征是羟苯丙酮酸双加氧酶基因抑制剂是硫三酮。
CNB96195857XA 1995-06-02 1996-06-03 羟苯丙酮酸双加氧酶基因的dna序列以及含有羟苯丙酮酸双加氧酶基因的抗特定除草剂植物的获得 Expired - Fee Related CN1206358C (zh)

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FR95/06800 1995-06-02
FR9506800A FR2734840B1 (fr) 1995-06-02 1995-06-02 Gene de l'hydroxy-phenyl pyruvate dioxygenase et obtention de plantes contenant ce gene resistantes aux herbicides
FR9513570A FR2734841B1 (fr) 1995-06-02 1995-11-10 Gene de l'hydroxy-phenyl pyruvate dioxygenase et obtention de plantes contenant ce gene resistantes aux herbicides
FR95/13570 1995-11-10
FR9605944A FR2734842B1 (fr) 1995-06-02 1996-05-07 Sequence adn d'un gene de l'hydroxy-phenyl pyruvate dioxygenase et obtention de plantes contenant un gene de l'hydroxy-phenyl pyruvate dioxygenase, tolerantes a certains herbicides
FR96/05944 1996-05-07

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