CN101123870A - 用于产生具有增加的角质层水透过性的植物的分离多肽以及编码该多肽的多核苷酸 - Google Patents
用于产生具有增加的角质层水透过性的植物的分离多肽以及编码该多肽的多核苷酸 Download PDFInfo
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Abstract
本发明提供一种分离的多核苷酸。所述分离的多核苷酸包含编码多肽的核酸序列,所述多肽具有与SEQ ID NO:22至少88%同源的氨基酸序列,所述多肽能够增加表达其的植物的角质层水透过性。本发明还提供产生表达所述多肽的植物的方法,所述多肽可用于产生脱水植物或其角质层覆盖部分。
Description
发明领域和背景
本发明涉及多核苷酸和多肽,所述多肽用于增加表达其的植物的角质层水透过性。更具体地说,本发明涉及遗传修饰的植物,其能够产生脱水果实,例如番茄。
高等植物的气生部分被角质层的连续胞外层覆盖。角质层是一种聚合物基质,主要由角质单体(主要是短链羟化脂肪酸)和各种量的角质层蜡质(溶剂可溶的脂质)组成。不同植物物种和植物器官间的角质和蜡质组分的量和成分都极为不同(Holloway,1982)。尽管已广泛研究了植物角质层的组成和结构以及其生物合成的生物和遗传调节(Kolattukudy,1980;Koornneef等,1989;Blee和Schuber,1993;Arts等,1996;Negruk等,1996;Millar等,1997;Todd等,1999;Yaphremov等,1999;Flebig等,2000;Pruitt等,2000;Wellesen等,2001;Hooker等,2002;Chen等,2003;Kuns和Samuels,2003;Kurata等,2003;Aharoni等,2004;Schnurr等,2004;),但控制角质层分化和/或功能的机制仍有待表征。
番茄果实角质层是具有4-10μm厚度的薄层,有两种特有的结构特性(Wilson和Sterling,1976)。首先,角质沉积物以层状结构排列-各层与表皮细胞平行地聚集。番茄果实角质层的第二种特性在于其不含任何气孔、孔或通道。因此,番茄皮的水透过性很低,完全成熟的番茄果实保持其含水量。许多没有气孔(无气孔)的其它角质层的水透过性和穿过其的水运输机制已成为众多研究的主题(Sch_nherr,1976a;Sch_nherr和Schmidt,1979;Riederer和Schreiber,2001)。显然,角质和蜡质组分这二者对抵抗器官失水具有集成作用。在某些情况下,角质层厚度与穿过角质膜的水扩散成反比(Lownds等,1993)。但是,在影响植物水透过性方面角质层蜡质组分常常占主导地位。例如,通过有机溶剂去除番茄果实角质层中角质层之上的蜡质层将其水透过性增加了300-500倍,以快速的植物脱水为证(Sch_nherr,1976b)。角质层蜡质在番茄果实中起蒸腾屏障作用的另一个证据是最近报道的编码酶超长链脂肪酸(VLFA)β-酮脂酰-CoA合酶的基因(LeCER6,Vogg等,2004)。该基因在果实角质层蜡质主要成分VLFA的合成中起重要作用。该基因中功能突变的缺失导致叶和果实蜡质中链长超过C30的正-烷烃和醛的减少。具有LeCER6突变的番茄果实的特征在于水透过性增加4倍。影响番茄果实角质层水透过性的另一个因素是在角质层表面上存在裂化现象。果实裂化已得到了更多的研究关注(Corner等,1969;Voisey等,1970;peet,1992;peet和willits,1995)。番茄果实受到3种主要类型的裂化作用侵袭:i)同心裂缝(粗裂);ii)径向裂缝(开裂);和iii)角质层裂缝(褐斑)(Bakker,1988)。前两类裂缝深,延伸的裂隙穿过果实果皮,除了失水外,还可今病原体进入和果实腐败。
与径向或同心裂缝不同,角质层裂缝是角质层的表面微裂隙,一般只限于角质层,不穿透入果皮细胞。导致番茄果实裂化的原因和环境大部分不清楚,可能与角质层厚度(Emmons和Scott,1998)、下面表皮细胞的形状(Conter等,1969;Emmons和Scott,1998)、果实形状(Considine和Brown,1981)、果实大小(Koske等,1980;Emmons和Scott,1997)、果实周围的相对湿度(Young,1947;Tukey,1959)、肥叶修剪(Ehret等,1993)以及表皮的抗张强度和延展性(Bakker,1988)有关。
番茄果实中出现裂缝还具有显著的遗传成分,主要由野生种番茄(Lycopersicon)在基因转移时显现。Fulton等(2000)描述了一种性状“表皮网纹”(Er),并使用高代回交QTL分析策略(采用野生型小花番茄(L.parviflorum))报告了4个影响其的QTL。在源自普通番茄(L.esculentum)和其它野生种如多毛番茄(L.hirsutum)(WO 0113708)和潘那利番茄(L.penellii)(Monforte等,2001)杂交的番茄果实中也已报告了角质层裂缝。
由于沿着裂隙出现纤维质化表层(Monforte等,2001)而产生的果实角质层中的裂缝,特别是以表皮网纹为证的目视可见的大裂缝,一般被看作是无益现象,原因是降低果实价值的美感损害(Tukey,1959)以及果实含水量的损失。但是,脱水但完整并仍连接在藤上的果实的经济潜力高。脱水番茄产品构成了番茄产业的一个重要部分。番茄糊(tomato paste)、蕃茄酱(ketchup)和其它加工番茄产品的生产依赖于需要能量的脱水步骤。另外,以干燥工艺制备“晒干的”番茄果实,所述干燥工艺由在太阳下或在烘箱中脱水已切开的番茄果实组成。晒干和烘箱干燥都可导致食物品质损失。例如,番茄在人类膳食中的其中一种主要营养成分抗坏血酸的水平在晒干和烘箱干燥作用下都显著下降(Ojimelukwe,1994)。而且,在干燥工艺前必须将番茄果实切成两半使得不能生产完整的干番茄果实。
本发明的发明人先前已描述了使用经典遗传育种技术获得的、具有降低的水含量的脱水番茄(WO 01/13708)。要认识到的是,经典遗传育种技术限于种内或同一属的密切相关种之间的基因转移。另外,经典育种的特征在于相对大的基因渗入,其包含与目标基因紧密连锁的其它不需要基因。
Eshed和Zamir(1985)先前已描述了基因渗入的栽培番茄植物,其具有可能与不需要的性状相关的遗传背景(基因渗入系IL4-4,即产生于由端粒标记TG464延伸至着丝粒标记CT50的基因渗入;约20cM)。同样,Monforte等(2001)已描述了来源于多毛番茄(L.hirsutum)的具有相似遗传背景的番茄植物[次近基因渗入系TA1468、TA1469、TA1476,其覆盖并包括TG464至CT173(约10cM)],其表现出众多作用,包括不需要的作用。
因此,获得具有增加的角质层水透过性、避免以上限制的遗传修饰植物存在广泛认可的需求,应当是高度有利的。
发明概述
按照本发明的一个方面,提供一种包含编码多肽的核酸序列的分离多核苷酸,所述多肽具有的氨基酸序列与SEQ ID NO:22至少88%同源,所述多肽能够增加表达其的植物的角质层水透过性。
按照下文描述的本发明优选实施方案的其它特征,所述核酸序列如SEQ ID NO:21或23所述。
按照所述优选实施方案的再其它特征,所述氨基酸序列如SEQ IDNO:22所述。
按照本发明的另一方面,核酸构建物包含所述分离的多核苷酸。
按照所述优选实施方案的再其它特征,所述核酸构建物还包含有效连接至所述核酸序列的启动子。
按照本发明的另一方面,宿主细胞包含所述核酸构建物。
按照本发明的另一方面,遗传修饰的植物含所述分离的多核苷酸。
按照本发明的另一方面,寡核苷酸能够与所述分离的多核苷酸特异性杂交。
按照本发明的另一方面,提供一种分离的多肽,其包含的氨基酸序列与SEQ ID NO:22至少88%同源,所述多肽能够增加表达其的植物的角质层水透过性。
按照本发明的再另一方面,提供能够特异性识别所述多肽的抗体。
按照本发明的再另一方面,提供一种栽培的番茄植物,其具有的基因组包含来源于野生番茄(Lycopersicon spp.)的基因渗入,所述基因渗入包含番茄(Lycopersicon spp.)染色体4的一部分,小于由端粒标记TG464延伸至着丝粒标记CT173的染色体片断,所述基因渗入能够增加栽培番茄植物的角质层水透过性。
按照本发明的再另一方面,提供一种生产作物的脱水果实的方法,该方法包括遗传修饰植物,以表达具有与SEQ ID NO:22至少30%同源的氨基酸序列的多肽,所述多肽能够增加表达其的植物的角质层水透过性。
按照所述优选实施方案的再其它特征,所述方法还包括:
使果实在植物上脱水;随后
收集脱水果实。
按照所述优选实施方案的再其它特征,所述方法还包括:
在果实脱水前由作物上取下果实;随后令果实脱水。
按照本发明的另一方面,提供一种遗传修饰的种子,其包含的分离的多核苷酸含有编码多肽的核酸序列,所述多肽具有的氨基酸序列与SEQ ID NO:22至少30%同源,所述多肽能够增加表达其的植物的角质层水透过性。
按照本发明的再另一方面,提供一种遗传修饰的果实,其包含的分离的多核苷酸含有编码多肽的核酸序列,所述多肽具有的氨基酸序列与SEQ ID NO:22至少30%同源,所述多肽能够增加表达其的植物的角质层水透过性。
按照所述优选实施方案的再其它特征,所述核酸序列如SEQ IDNO:21、23、24、26、28、30、32、34、36、38、40、42、44、46、48、50、52、54或56所述。
按照所述优选实施方案的再其它特征,所述氨基酸序列如SEQ IDNO:22、25、27、29、31、33、35、37、39、41、43、45、47、49、51、53、55或57所述。
按照本发明的再另一方面,提供一种遗传修饰的植物,其表达的多肽具有与SEQ ID NO:22至少30%同源的氨基酸序列,所述多肽能够增加植物的角质层水透过性。
本发明通过提供能够增加表达其的植物的角质层水透过性的多核苷酸和多肽,并通过提供用于生产脱水果实的遗传修饰植物,成功地解决了目前已知形态的缺点。
除非另有定义,否则本文使用的所有技术和科学术语都具有和本发明所属领域一般技术人员的通常理解相同的含义。尽管可使用与本文所述类似或等同的方法和材料实施或检验本发明,但下文描述了合适的方法和材料。在有抵触的情况下,以包含定义的专利说明书为准。另外,所述材料、方法和实施例仅为阐述性的,没有限制意图。
附图简述
仅通过示例的方式,本文参考附图描述本发明。现在具体地讨论附图,要强调指出的是,显示的详解仅作为实例,并仅用于本发明优选实施方案的说明性论述目的,提出这些详解是为了提供一般认为对本发明的原则和概念方面最有用的和最容易理解的描述。就此而言,未尝试更详细地表明超过基本理解本发明所需的本发明结构性细节,描述连带附图使本领域技术人员更清楚如何具体地实施本发明的几种形式。
在附图中:
图1a-b图示了2148群(图1a)和2149群(图1b)中cwp(PUT)基因型对脱水率的作用。在2148群中,脱水性状表现为完全显性性状,而在2149中,其表现为部分显性的性状。果实在红熟时采摘,并令其在室温脱水,以约每日一次的间隔称重。数据表示为Log%重量。上标HH、HE和EE表示分离植株的基因型。
图2a-c显示了CWP基因的精细图谱。图2a-TG464分子标记的CAPS标记分析。基因组DNA由20株根据脱水率分离的F2个体提取。使用适合于表现出两个亲本物种间多态性的TG464标记的引物进行PCR分析。PCR产物用HinF1内切核酸酶限制性位点酶切割,并在2%琼脂糖凝胶上电泳。+或-信号指示存在或没有微裂隙和脱水状况。E-普通番茄(L.esculentum.),H-多毛番茄(L.hirsutum.),M-HindIII/EcorIλ标记(Fermentas目录号SM0191)。图2b-CWP染色体区的遗传连锁图(以cM表示),相对于着丝点位置定向。标示了潘那利番茄(L.penellii)基因渗入系IL4.3和IL4.4(Eshed和Zamir,1995)。阴影条代表在该分析中用作脱水供体亲本的近等基因系中的多毛番茄(L.hirsutum.)节段。图2c-Cwp基因侧翼染色体节段的放大。
图3a-b表示CWP基因的物理定位。图3a-在CT61和TG464 CAPS标记之间建桥的遗传有序的连续BAC,以及根据测序BAC末端的多态性的重组体表型分析和重组体特征。每个重组体基因型都由分为阴影(多毛番茄(L.hirsutum)基因型)和空(普通番茄(L.esculentum)基因型)节段的条代表。图3b-BAC 37B8中3个交换事件的放大。3个交换事件是图3a中列出的前3个重组体。
图4图示了来自多毛番茄(L.hirsutum)的15kb基因渗入,其包含Cwp基因。使用NCBI程序TBLAST分析序列的同源可读框。鉴别出3个同源序列,各个可读框的方向由箭头指示。
图5a-b图示了发育中的番茄子房和小果中PUT(图5a)和DBP(图5b)基因的表达分析。如在方法章节中的描述使用在线定量PCR检测提取的cDNA的表达,表示为相对于各个样品中肌动蛋白基因的表达。Ov,子房;开花后5-15天;IG,生绿期,MG,熟绿期;B,花端着色期。阴影条是CwpHH基因型,实心条是CwpEE基因型。
图6图示了番茄基因型15天小果中PUT基因的表达分析。HH,CwpHH基因型;HE,杂合CwpHE基因型;EE,CwpEE基因型。3种基因型选自分离型杂合群。IL4.4代表潘那利番茄(L.penellii)基因渗入系IL4.4(Eshed和Zamir,1985),其包含PUT的潘那利番茄(L.penellii)同源物。M82是IL 4.4的回交普通番茄(L.esculentum)亲本。
图7a-b显示了在35S组成型启动子控制下表达野生番茄种Solatium habrochaites S.(先前称为多毛番茄(Lycopersicon hirsutumMill.))的PUT基因的转基因番茄植物(T0)。图7a显示了双筒显微照片,列出了野生型MP1番茄系(W.T.)果实以及微裂隙转基因果实(Mp1-4)的完整表面。图7b显示了野生型MP1番茄系(W.T.)和另一个独立转基因T0植物(MP1-1)之间的脱水率对比。果实于熟红发育期采摘,置于室温(15-25℃)。图片得自实验开始时(T0)和脱水7天后(T7)。
图8a-b显示了PUT转基因拷贝数对微裂隙严重性(评分在1-5之间,图8a)和果实失重百分率(在室温14天后,图8b)的作用。由两个独立的转基因(T1)分离群(每个群16个个体)收集检测结果。每张图显示了各个拷贝数组个体的平均数(在各个菱形中间的水平线)、95%置信限(菱形上下界线)和分散程度。当一组三角形的底与另一组的三角形底不一致时,组间差异是显著的。通过JMP程序进行统计。
图9a-b显示了类似于野生型的无拷贝表达的转基因番茄个体(T1代)和表达两个拷贝的野生番茄种Solatium habrochaites S.的PUT基因的转基因番茄个体(T1代)之间的对比。图9a-扫描电子显微照片,呈现了无PUT基因拷贝(0拷贝)的个体果实以及具有两个拷贝转基因(2拷贝)的个体微裂隙果实的完整表面。图9b-无PUT基因拷贝(0拷贝)的个体以及具有两个拷贝转基因(2拷贝)的个体之间的脱水率对比。果实于熟红发育期采摘,置于室温(15-25℃)。图片得自实验开始时(T0)和脱水7天后(T7)。
图10a-b是树状图,图示了CWP1和CWP2的保守性以及单子叶和双子叶物种的相关序列(SEQ ID NO.21、24、26、28、30、32、34、36、38、40、42、44、46、48、50、52、54和56)。这些序列基于与CWP1的序列同源性由EST TIGR数据库检索。上方标示了与CWP1的同源性百分率。图10a-在氨基酸水平上的保守性。图10b-在核酸水平上的保守性。
图11显示了利用EMBL-EBI的ClustalW软件产生的本发明CWP1家族不同蛋白成员之间的多重比对。
优选实施方案的描述
本发明为可用于增加植物的角质层水透过性的分离多核苷酸和多肽。具体地说,本发明可用于生产脱水果实,例如番茄果实。
参阅附图和所附说明书可更好地理解本发明的原则和实施。
在详细解释本发明的至少一个实施方案前,要理解本发明的应用不限于以下说明书陈述的或实施例例举的细节。本发明允许有其它实施方案,或者以各种方式实施或进行。另外,要理解本文使用的措辞和术语用于描述目的,不应被理解为限制作用。
能够天然脱水但仍连接在藤上、不伴随导致果实腐败的降解过程的番茄品种的开发对许多果实产业(如番茄产业)是高度有价值的。
Schaffer的PCT公开号WO 01/13708教导了使用经典遗传育种技术(WO 01/13708)产生角质层水含量降低的脱水番茄。要认识到的是,经典遗传育种技术限于种内或同一属的密切相关种之间的基因转移。另外,经典育种的特征在于相对大的基因渗入,其包含与目标基因紧密连锁的其它不需要基因。
Eshed和Zamir(1985)先前已描述了基因渗入的栽培番茄植物,其具有可能与不需要的性状相关的遗传背景(基因渗入系IL4-4,即产生于由端粒标记TG464延伸至着丝粒标记CT50的基因渗入;约20cM)。同样,Monforte等(2001)已描述了来源于多毛番茄(L.hirsutum)的具有相似遗传背景的番茄植物[次近基因渗入系(NIL),其覆盖TG464至CT173(>10cM)]。在后一研究中,相对大的基因渗入伴随不需要的园艺性状,包括白利糖度产量、总产量和果实重量的性状。
在简化本发明以实施的同时,本发明的发明人揭示了单个基因cwp1(也称为put,在本文可交互使用),其能够增加表达其的植物的角质层水透过性。
如下文和以下的实施例章节所阐述的,本发明的发明人鉴别出来源于多毛番茄(L.hirsutum)的果实脱水性状的遗传模式为单个主要基因。本发明的发明人使用基于图谱的定位克隆策略,由野生番茄种多毛番茄(L.hirsutum)克隆出增加熟红番茄果实的角质层水透过性(CWP)并导致完整果实脱水的基因。
本发明的发明人表明,cwp的野生种等位基因允许该基因在发育中的番茄果实中表达,而标准的栽培普通番茄(L.esculentum)等位基因不表达,可被看作是无效等位基因。他们还表明,在表达水平上存在等位基因剂量作用,杂合HE基因型的特征在于其表达为具有两个野生种等位基因的纯合基因型的约一半。
生物信息学分析表明,cwp1编码生物功能未知的蛋白。该基因可能有助于新番茄产品以及其它作物的育种程序,因为其通过角质层控制失水量。而且,与该基因相关的微裂隙的结构表型表明了cwp在果实角质层发育中的作用。在栽培番茄中处于35S启动子控制下的cwp1基因表达在膨大果实中诱导形成微裂隙,这支持所提出的该基因在角质层水透过性调节中的作用。DNA印迹分析揭示了另外的番茄同源物cwp2。令人感兴趣的是,该同源物作图至番茄染色体2-1,在番茄染色体2-1中有番茄果实表皮网纹的报告QTL(Frary等,2004)。茄属栽培胡椒(辣椒(Capsicum annum))的发育中果实也在其表皮组织中表达与Lecwp1基因高度相似(87%)的cwp同源物,胡椒果实的特征在于收获后失水的园艺问题,以及在红辣椒栽培品种中果实脱水的需要性状。因此,CWP基因的同源物有可能还有助于角质层修饰和水透过性。
这些结果表明,cwp基因的表达产生结构修饰的角质层(基于重量和TEM),推测其在果实膨大过程中由于弹性下降而经历缝裂。但是,此现象仅在具有高度发育的果实角质层的果实中观察到,例如无气孔厚皮栽培番茄,在以较薄角质层为其特征的野生种果实中不明显。在栽培番茄果实发育过程中角质层组分的沉积在由生绿期向熟绿期的转变过程中出现突然上升(Baker,1982),这是合理的,因为其与微裂隙表型的观察结果一致。
非囿于理论,考虑到成熟和收获果实的稳定性,已表明相对不透水的果实角质层的遗传性状在栽培番茄的进化和驯化过程中是一种良性发育。脱水性状的遗传控制指明了在作物进化和驯化的某些阶段对无效Cwp的选择过程。
系统发生分析(图10a-b)表明,本发明的CWP基因属于一个较大的基因家族,其可用于控制广泛作物的角质层水透过性。
因此,本发明的一个方面提供一种包含核酸序列的分离的多核苷酸,该核酸序列编码的多肽具有与SEQ ID NO:22至少约30%、至少约40%、至少约50%、至少约55%、至少约60%、至少约65%、至少约70%、至少约75%,、至少约80%、至少约85%、至少约90%、至少约91%、至少约92%、至少约93%、至少约94%、至少约95%、至少约96%、至少约97%、至少约98%、至少约99%或100%同源的氨基酸序列,所述多肽能够增加表达其的植物的角质层水透过性。
本文使用的表述“角质层水透过性”指角质层抑制被角质层环绕的植物组织(植物的气生组织,例如果实)的水分蒸发的能力。要认识到的是,增加的角质层水透过性将导致被角质层环绕的组织由于增强的蒸发而产生脱水。
本文使用的表述“增加角质层水透过性”指与不表达其的相似亲本栽培品种或基因型的植物相比,角质层水透过性增加至少约5%、至少约10%、至少约15%、至少约20%、至少约30%、至少约40%、至少约50%。
测定角质层水透过性的方法在本领域众所周知,详尽地描述于以下的实施例章节(例如果实的裂隙严重性和失重百分率)。参见以下实施例章节的实施例5。另外,检测角质层水透过性的方法还包括但不限于检测穿过分离的果皮的水扩散、检测极性孔大小和水动力学透过性(Sch_nherr,1976)。这些功能测定连同本文提供的结构性指引一起使得可以鉴别本发明的多核苷酸和多肽的功能同源物。
同源性(例如同源性百分率)可使用任何同源性比较软件测定,包括例如美国国家生物技术信息中心(NCBI)的BlastP软件,例如使用默认参数。
同一性(例如同源性百分率)可使用任何同源性比较软件测定,包括例如美国国家生物技术信息中心(NCBI)的BlastN软件,例如使用默认参数。
本文使用的表述“分离的多核苷酸”指单链或双链核酸序列,其是分离的或以RNA序列、互补多核苷酸序列(cDNA)、基因组多核苷酸序列和/或复合多核苷酸序列(例如以上的组合)的形式提供。
本文使用的表述“互补多核苷酸序列”指使用逆转录酶或任何其它RNA依赖性DNA聚合酶由信使RNA的逆转录产生的序列。这种序列随后可使用DNA依赖性DNA聚合酶在体内或体外扩增。
本文使用的表述“基因组多核苷酸序列”指来源于(分离自)染色体的序列,因此其代表染色体的连续部分。
本文使用的表述“复合多核苷酸序列”指至少部分互补并至少部分为基因组的序列。复合多核苷酸序列可包括编码本发明多肽需要的某些外显子序列,以及在其间插入的某些内含子序列。内含子序列可为任何来源,包括其它基因的内含子序列,通常包括保守的剪接信号序列。这种内含子序列还可包括顺式作用表达调节元件。
按照本发明此方面的一个优选实施方案,本发明的上述分离的多核苷酸的核酸序列如SEQ ID NO:21、23、24、26、28、30、32、34、36、38、40、42、44、46、48、50、52、54或56所述。
按照本发明此方面的另一个优选实施方案,本发明的编码多肽的氨基酸序列如SEQ ID NO:22、25、27、29、31、33、35、37、39、41、43、45、47、49、51、53、55或57所述。
本发明此方面的分离的多核苷酸可使用杂交测定如下检定:在中等至严格杂交条件下,在寡核苷酸探针或引物存在时,温育上述分离的多核苷酸。
本文使用的术语“寡核苷酸”指核糖核酸(RNA)或脱氧核糖核酸(DNA)或其模拟物的单链或双链寡聚物或多聚物。该术语包括由天然碱基、糖和共价核苷酸间键合(例如骨架)组成的寡核苷酸,以及具有起对应天然部分类似作用的非天然部分的寡核苷酸。
按照本发明教导设计的寡核苷酸可按照本领域已知的任何寡核苷酸合成法产生,例如酶合成或固相合成。执行固相合成的设备和试剂可由例如Applied Biosystems市售。还可使用用于这种合成的任何其它工具;所述寡核苷酸的实际合成完全属于本领域技术人员的能力范围,可通过在例如:Sambrook,J.和Russell,D.W.(2001),“Molecular Cloning:A Laboratory Manual”;Ausubel,R.M.等编辑,(1994,1989),“Current Protocols in Molecular Biology,”I-III卷,JohnWiley & Sons,Baltimore,Maryland;Perbal,B.(1988),“A Practical Guideto Molecular Cloning,”John Wiley & Sons,New York;和Gait,M.J.编辑,(1984),“Oligonucleotide Synthesis”中详述的现有方法实现;使用固相化学法例如氰乙基氨基磷酸酯来实现,后接去保护、脱盐和利用例如自动化三苯甲基保护(trityl-on)法或HPLC纯化。
本发明的寡核苷酸为至少17个、至少18个、至少19个、至少20个、至少22个、至少25个、至少30个或至少40个碱基,可与本发明的多核苷酸序列特异性杂交。
中等至严格杂交条件的特征在于杂交溶液例如含10%硫酸葡聚糖、1M NaCl、1%SDS和5×106cpm32P标记探针,于65℃进行,最终漂洗溶液为0.2×SSC和0.1%SDS,最终漂洗于65℃进行,而中等杂交使用以下条件实现:杂交溶液含10%硫酸葡聚糖、1M NaCl、1%SDS和5×106cpm32P标记探针,于65℃进行,最终漂洗溶液为1×SSC和0.1%SDS,最终漂洗于50℃进行。
使用杂交方法,本发明的发明人能够分离cwp1的另一个番茄同源物cwp2,其作图至番茄果实表皮网纹的报告QTL(Frary等,2004),这支持其在角质层水透过性方面的作用。
因此,本发明包含上文所述核酸序列、其片段、与其杂交的序列、与其同源的序列、编码具有不同密码子选择的相似多肽的序列、特征在于突变的改变序列,所述突变例如为一个或多个核苷酸的缺失、插入或置换,或者是天然的,或者是人工诱导的,或者是随机的,或者为靶向方式。
因为本发明的多核苷酸序列编码先前未鉴别出的多肽,所以本发明还包括新多肽或其部分,其由上文所述的分离多核苷酸及其相应的核酸片段编码。
因此,本发明还包括由本发明的多核苷酸序列编码的多肽。这些新多肽的氨基酸序列如SEQ ID NO:22、25、27、29、31、33、35、37、39、41、43、45、47、49、51、53、55或57所述。
本发明还包括这些多肽的同源物,这种同源物可与SEQ ID NO:22至少约70%、至少约75%、至少约80%、至少约81%、至少约82%、至少约83%、至少约84%、至少约85%、至少约86%、至少约87%、至少约88%、至少约89%、至少约90%、至少约91%、至少约92%、至少约93%、至少约94%、至少约95%、至少约96%、至少约97%、至少约98%、至少约99%或更多(例如100%)同源。
本发明还包括上述多肽的片段和具有突变的多肽,所述突变例如为一个或多个氨基酸的缺失、插入或置换,或者是天然的,或者是人工诱导的,或者是随机的,或者为靶向方式。
本发明多肽的氨基酸序列信息可用于产生与本发明多肽特异性结合的抗体。例如,这种抗体可针对SEQ ID NO:22的氨基酸序列坐标55-160。序列坐标55-160包括大部分保守序列和多重对比分析的基序(图11)。由于在该氨基酸序列区域中的高序列同源性,预期这种抗体与本发明的各种多肽(例如SEQ ID NO:22、25、27、29、31、33、35、37、39、41、43、45、47、49、51、53、55或57)交叉反应。
本发明的多核苷酸和多肽序列可用于产生具有增加的角质层水透过性的植物。
例如,遗传修饰的植物可通过在植物中表达本发明的分离的多核苷酸来产生。
本文使用的术语“植物”指其中在商业上需要增加角质层水透过性的作物(全株或其部分,例如果实、种子),例如单子叶或双子叶作物,以及其它球果植物、藓类或藻类植物。优选地,本发明的植物产生脱水具有商业价值的果实。这种植物的实例包括但不限于番茄、葡萄、胡椒、苹果、桃、杏、枣、无花果、茄子、洋葱、草莓、葫芦、干草植物、饲料植物、香料植物、草本植物等。
为在植物细胞中表达外源多核苷酸,本发明的多核苷酸序列优选连接入适于植物细胞表达的核酸构建物中。这种核酸构建物包含顺式作用调节区,例如以组成型方式或诱导型方式引导细胞中的多核苷酸序列转录的启动子序列。所述启动子对于转化的植物/细胞可为同源的或异源的。
可按照本发明的该方面使用的优选启动子序列是果实特异性或种子特异性启动子。
例如,cwp1基因(或其功能片段)的新启动子序列优选可用于本发明的核酸构建物(SEQ ID NO:58)。
果实特异性启动子的其它实例描述于美国专利第4,943,674号。
可按照本发明的该方面使用的其它启动子是确保仅在植物的特定气生暴露器官(例如叶、块茎、种子、茎、花)或特定细胞类型(例如薄壁组织、表皮、毛状体或维管细胞)中表达的启动子。
在种子中增强表达的优选启动子包括phas启动子(Geest等,PlantMol.Biol.32:579-588(1996))、GluB-1启动子(Takaiwa等,Plant Mol.Biol.30:1207-1221(1996))、γ-玉米醇溶蛋白启动子(Torrent等,PlantMol.Biol.34:139-149(1997))以及油体蛋白启动子(Sarmiento等,ThePlant Journal 11:783-796(1997))。
介导组成型、诱导型、组织特异性或发育期特异性表达的启动子序列公开于WO 2004/081173。
核酸构建物可为例如质粒、杆粒、噬菌粒、粘粒、噬菌体、病毒或人工染色体。优选地,本发明的核酸构建物是质粒载体,更优选为双元载体。
表述“双元载体”指携带Ti质粒的修饰T区的表达载体,其在大肠杆菌和农杆菌细胞这二者中均能复制,通常在两个边界区之间包含植物转化的报告基因。适于本发明的双元载体包括pBI2113、pBI121、pGA482、pGAH、pBIG、pBI101(Clonetech)、pPI和pBIN PLUS(参见以下的实施例章节的实施例5)或其修饰形式。
本发明的核酸构建物可用于转化宿主细胞(例如细菌、植物)或植物。
本文使用的术语“转基因的”或“转化的”可交互使用,指克隆的遗传物质已转移入其中的细胞或植物。
在稳定转化中,本发明的核酸分子整合入植物基因组中,因而其代表稳定的和遗传的性状。在瞬时转化中,核酸分子由被转化但未整合入基因组中的细胞表达,因而代表暂时性状。
有多种将外源基因导入单子叶植物和双子叶植物这二者中的方法(Potrykus,I.(1991).Annu Rev Plant Physiol Plant Mol Biol 42,205-225;Shimamoto,K.等(1989).Fertile transgenic rice plants regeneratedfrom transformed protoplasts.Nature(1989)338,274-276)。
外源DNA稳定整合入植物基因组DNA中的主要方法包括两个主要途径:
(i)农杆菌介导的基因转移。参见:Klee,H.J.等,(1987).Annu RevPlant Physiol 38,467-486;Klee,H.J.和Rogers,S.G.(1989).Cell Cultureand Somatic Cell Genetics of Plants,第6卷,Molecular Biology of PlantNuclear Genes,2-25页,J.Schell和L.K.Vasil编辑,AcademicPublishers,San Diego,CaL;和Gatenby,A.A.(1989).Regulation andExpression of Plant Genes in Microorganisms,93-112页,PlantBiotechnology,S.Rung和C.J.Arntzen编辑,Butterworth Publishers,Boston,Mass。
(ii)直接DNA摄取。参见例如:Paszkowski,J.等,(1989).CellCulture and Somatic Cell Genetics of Plants,第6卷,Molecular Biologyof Plant Nuclear Genes,52-68页,J.Schell和L.K.Vasil编辑,AcademicPublishers,San Diego,CaL;和Toriyama,K.等,(1988).Bio/Technol 6,1072-1074(将DNA直接摄取入原生质体中的方法)。另参见:Zhang等,(1988).Plant Cell Rep 7,379-384;和Fromm,M.E.等,(1986).Stabletransformation of maize after gene transfer by electroporation.Nature 319,791-793(通过短暂电击植物细胞诱导的DNA摄取)。另参见:Klein等,(1988).Bio/Technology 6,559-563;McCabe,D.E.等,(1988).Stabletransformation of soybean(Glycine max)by particle acceleration.Bio/Technology 6,923-926;和Sanford,J.C.(1990).Biolistic planttransformation.Physiol Plant 79,206-209(通过粒子轰击将DNA注射入植物细胞或组织中)。另参见:Neuhaus,J.M.等,(1987).Theor ApplGenet 75,30-36;以及Neuhaus,J.M.和Spangenberg,G.C.(1990).Physiol Plant 79,213-217(使用微量移液管系统)。参见美国专利第5,464,765号(细胞培养物、胚或愈伤组织的玻璃纤维或碳化硅晶须转化)。另参见:De Wet,J.M.J.等,(1985).“Exogenous gene transfer inmaize(Zea mays)using DNA-treated pollen,”Experimental Manipulationof Ovule Tissue,G.P.Chapman等编辑,Longman,New York-London,197-209页;和Ohta,Y.(1986).High-Efficiency Genetic Transformationof Maize by a Mixture of Pollen and Exogenous DNA.Proc Natl Acad SciUSA 83,715-719(DNA与萌发花粉直接温育)。
农杆菌介导的系统包括使用含限定DNA节段的质粒载体,所述DNA节段整合入植物基因组DNA中。嫁接植物组织的方法随植物物种和农杆菌传递系统而变。广泛使用的方法是叶盘法,该方法可用提供启动全株分化的良好来源的任何组织外植体实施(Horsch,R.B.等,(1988).“Leaf disc transformation.”Plant Molecular Biology ManualA5,1-9,Kluwer Academic Publishers,Dordrecht)。补充方法使用农杆菌传递系统和真空浸润的组合。农杆菌系统对建立转基因双子叶植物尤其有用。
有多种将DNA直接转移入植物细胞中的方法。在电穿孔中,原生质体短暂暴露于强电场,打开小孔以允许DNA进入。在微注射中,使用微量移液器将DNA机械地注射入细胞中。在微粒轰击中,DNA吸附在微粒如硫酸镁晶体或钨颗粒上,微粒被物理加速进入细胞或植物组织中。
在稳定转化后,发生植物繁殖。最常见的植物繁殖法是播种。但是,播种繁殖再生的缺点是由于杂合性而使作物缺乏一致性,因为植物按照孟德尔法则支配的遗传变异产生种子。换句话说,每个种子在遗传上都是不同的,每个种子都将以其自身的特定性状生长。因此,优选要实现的再生使再生植物具有相同性状,并具有亲本转基因植物的特征。再生转化植物的优选方法是微繁殖,其提供快速一致的转化植物再生。
微繁殖是一种由选定亲本植物或栽培品种切离的单个组织样品培养第二代植物的方法。该方法允许大量再生具有优选组织并表达融合蛋白的植物。新生植物在遗传上与原植物相同,并具有其全部特征。微繁殖允许在短时间段内大量生产优质植物体,并提供选定栽培品种的快速繁殖,其保留了原转基因或转化植物的特征。该植物克隆法的优势包括植物繁殖速度以及所产生植物的质量和一致性。
微繁殖是一个多阶段方法,需要改变各阶段间的培养基或培养条件。微繁殖法包括4个基本阶段:第1阶段,初始组织培养;第2阶段,组织培养物繁殖;第3阶段,分化和植物形成;和第4阶段,温室培养和锻炼。在第1阶段中,建立组织培养物并证明其无污染物。在第2阶段中,繁殖初始组织培养物,直至产生足够数量的组织样品,以满足生产目标。在第3阶段中,新培养的组织样品分裂,生长成单株幼苗。在第4阶段,转化的幼苗转移至温室进行锻炼,在温室中逐渐增加植物对光的耐受,以使它们可在天然环境中继续生长。
瞬时转化可通过上述的任何直接DNA转移法或通过使用修饰植物病毒的病毒感染来实现。
已表明对植物宿主转化有用的病毒包括花椰菜花叶病毒(CaMV)、烟草花叶病毒(TMV)和杆状病毒(BV)。使用植物病毒转化植物描述于例如:美国专利第4,855,237号(菜豆金黄花叶病毒,BGMV);EPA 67,553(TMV);日本特许申请号63-14693(TMV);EPA194,809(BV);EPA 278,667(BV);和Gluzman,Y.等,(1988).Communications in Molecular Biology:Viral Vectors,Cold SpringHarbor Laboratory,New York,172-189页。假病毒颗粒在许多宿主(包括植物)中表达外源DNA的用途描述于WO 87/06261。
用于在植物中导入和表达非病毒外源核酸序列的植物RNA病毒的构建阐述于以上参考文献以及:Dawson,W.O.等(1989)A tobaccomosaic virus-hybrid expresses and loses an added gene.Virology 172,285-292;French,R.等(1986)Science 231,1294-1297;和Takamatsu,N.等(1990).Production of enkephalin in tobacco protoplasts using tobaccomosaic virus RNA vector.FEBS Lett 269,73-76。
如果转化病毒是DNA病毒,本领域技术人员可针对病毒自身做出合适修饰。或者,所述病毒首先可克隆入细菌质粒中,以方便构建具有外源DNA的目标病毒载体。然后可由质粒中切出病毒。如果所述病毒是DNA病毒,则可将细菌复制起点连接至病毒DNA,然后通过细菌复制病毒DNA。DNA的转录和翻译将产生外壳蛋白,其将壳体化病毒DNA。如果所述病毒是RNA病毒,则一般将病毒克隆为cDNA,并插入质粒中。然后使用所述质粒制备所有植物遗传构建物。然后由质粒的病毒序列转录RNA病毒,接着翻译病毒基因,以生产壳体化病毒RNA的外壳蛋白。
在以上参考文献以及美国专利第5,316,931号中论述了用于在植物中导入和表达非病毒外源核酸序列(例如包含在本发明构建物中的序列)的植物RNA病毒的构建。
在一个实施方案中,提供用于插入的植物病毒核酸,其包含病毒核酸天然外壳蛋白编码序列的缺失、非天然(外源)植物病毒外壳蛋白编码序列以及非天然启动子,优选非天然外壳蛋白编码序列的亚基因组启动子,并能够在植物宿主中表达,包装重组植物病毒核酸,并确保重组植物病毒核酸系统感染宿主。或者,天然外壳蛋白编码序列可通过在其中插入非天然核酸序列而变成不可转录的,以产生非天然蛋白。重组植物病毒核酸构建物可包含一个或多个另外的非天然亚基因组启动子。每个非天然亚基因组启动子均能够在植物宿主中转录或表达邻近基因或核酸序列,且不能彼此重组和与天然亚基因组启动子重组。另外,重组植物病毒核酸构建物可包含一个或多个顺式作用调节元件,例如增强子,其结合反式作用调节子,并调节位于其下游的编码序列的转录。非天然核酸序列可邻近天然植物病毒亚基因组启动子插入,或者如果包含一个以上的核酸序列,则可邻近多个天然和非天然植物病毒亚基因组启动子插入。非天然核酸序列在宿主植物中于亚基因组控制下转录或表达,以生产目标产物。
在第二个实施方案中,如在第一个实施方案中一样提供重组植物病毒核酸构建物,例外之处是将天然外壳蛋白编码序列置于其中一个非天然外壳蛋白亚基因组启动子附近,而不是在非天然外壳蛋白编码序列附近。
在第三个实施方案中,提供重组植物病毒核酸构建物,其包含处于其亚基因组启动子附近的天然外壳蛋白基因,一个或多个非天然亚基因组启动子插入到病毒核酸构建物中。插入的非天然亚基因组启动子能够在植物宿主中转录或表达邻近基因,且不能彼此重组和与天然亚基因组启动子重组。非天然核酸序列可插入到非天然亚基因组植物病毒启动子附近,使得所述序列在宿主植物中于亚基因组控制下转录或表达,以生产目标产物。
在第四个实施方案中,如在第三个实施方案中一样提供重组植物病毒核酸构建物,例外之处是天然外壳蛋白编码序列被非天然外壳蛋白编码序列取代。
病毒载体被表达的外壳蛋白壳体化,以产生重组植物病毒,所述外壳蛋白由如上文所述的重组植物病毒核酸构建物编码。重组植物病毒核酸构建物或重组植物病毒用于感染合适的宿主植物。重组植物病毒核酸构建物能够在宿主中复制、在宿主中系统扩散以及在宿主中转录或表达一个或多个外源基因(分离的核酸),以生产目标蛋白。
除了上述以外,本发明的核酸分子还可导入到叶绿体基因组中,由此能进行叶绿体表达。
将外源核酸序列导入叶绿体基因组中的技术是已知的。该技术包括以下步骤。首先,化学处理植物细胞,以便将每个细胞的叶绿体数降至约1。然后,优选通过粒子轰击将外源核酸导入细胞中,目标是将至少一个外源核酸分子导入叶绿体中。由本领域一般技术人员选择能通过同源重组整合入叶绿体基因组中的外源核酸,同源重组可容易地通过叶绿体固有的酶实现。为此,所述外源核酸除了包含目标基因以外还包含至少一个来源于叶绿体基因组的核酸序列。另外,外源核酸包含选择标记,其通过顺序选择方法用于使操作者确定在此选择之后的全部或基本全部的叶绿体基因组拷贝都包含外源核酸。关于此技术的进一步细节可见于美国专利第4,945,050号和5,693,507号,其通过引用结合到本文中。因此,多肽可通过叶绿体蛋白表达系统生产,并整合入叶绿体内膜中。
本领域已知许多使作物和杂草间的基因流动最小的方法。以下是此类方法的非限制性描述[另参见美国专利申请第20040098760、20040172678号以及Daniell(2002)Nat.Biotech.20:581]。其它方法包括雄性和/或种子不育性(其阻止远交、自生种子传播)、闭花受精(其中授粉在开花前发生,由此防止远交)和无配生殖(种子来自植物源且不来自有性杂交,其控制远交和自生种子传播。参见美国专利第6,825,397号)。
母体遗传
胞质器的母体遗传为植物(叶绿体)和动物(线粒体)系统所共有。已提出了几种解释来说明此现象。它通过在个体中占过多比例的自私胞质因子促使群侵入1。另外,胞质因子的母体遗传是防止与雄性相关的疾病或病原体有性传播的进化机制;只有细胞核(不是胞质)在受精过程中被允许进入胚珠中[Gressel J.Molecular Biology in WeedControl(Taylor and Francis,London,2002)]。它还可能是在受精后发生在植物中的雄性核基因一般抑制的延伸[Avni Mol.Gen.Genet.225,273-277(1991)]。
在涉及遗传修饰作物中间或遗传修饰作物和杂草之间的远交可能性的情况下,高度需要使用叶绿体遗传工程来促进转基因的母体遗传。存在于大部分被子植物中的质体遗传的普遍模式是单亲母体的,叶绿体基因组在大部分作物中是母体遗传的。
叶绿体基因组的母体遗传在植物中于形成精细胞的生殖细胞发育过程中获得,然后精细胞与雌配子在受精过程中融合。生殖细胞是花粉形成过程中不等分裂的结果,不接受任何叶绿体[HagemannProtoplasma 152,57-64(1989)]。
在叶绿体转基因植物中转基因的母体遗传和通过花粉防止基因流动已在几个植物物种中成功地被证明,包括烟草和番茄[Daniell Nat.Biotechnol 16,345-348;Ruf Nat.Biotechnol.19,870-875(2001)]。尽管已转化了几种其它植物物种(包括马铃薯)的叶绿体基因组,但在这些研究中还没有证实母体遗传。但是,30多种转基因已稳定地整合入叶绿体基因组中,赋予目标植物性状,或者将转基因叶绿体用作生物工厂,以产生功能性生物药物或可食用疫苗或生物聚合物[DaniellTrends Plant Sci 7,84-91(2001);Daniell Curr.Opin.Biotechnol.13,136-141]。
同许多其它防范策略不同,母体遗传方法已在许多领域得到检验。Scott和Wilkinson[Nat.Biotechnol.17,390-392(1999)]研究了由两个野生群收集的天然杂合体中的质体遗传,这两个野生群沿着34公里的英国泰晤士河紧邻油菜生长,评价3年当中18个野生油菜群的持久性。他们分析了几种会影响叶绿体基因由作物向野生亲缘植物移动的可变因素,包括质体遗传模式和同域性(物种在相同区域一起出现)发生率,以量化形成混合群的机会和作物在基因渗入的农业限制之外的持久性。尽管作物和杂草物种间的同域性约为0.6-0.7%,但混交成分在3年中显示出植物数量快速下降、种子返回并最终消失的强烈趋势。因此,Scott和Wilkinson得出结论:如果经由叶绿体基因组对植物进行遗传工程,则基因流动应是罕见的。
因此,叶绿体基因组的母体遗传对于基因防范是一个有前景的选项。尽管质体转化在几种主要作物物种中仍待实现,但现已表明叶绿体遗传工程赋予对除草剂[Daniell Nat.Biotechnol.16,345-348(1998)],昆虫、疾病[DeGray Plant Physiol.127,852-862(2001)]和干旱的抗性,以及产生抗体[Daniell Trends Plant Sci.7,84-91(2001)]、生物药品[Daniell Trends Plant Sci.7,84-91(2001)]和可食用疫苗。近期欧洲环境署(Copenhagen,Denmark)的报告推荐叶绿体遗传工程作为基因防范方法[Eastham Genetically Modified Organisms(GMOs):TheSignificance of Gene Flow Through Pollen Transfer.Environmental IssueReport 28(European Environmental Agency,Copenhagen,Denmark,2002)]。
基因组不相容性-许多栽培作物具有多个基因组。这些作物基因组中仅有一个适于与杂草种间杂交。例如,小麦的D基因组与美国一种难控制杂草柱穗山羊草(Aegilops cylindrica)的D基因组相容;相反,只要具有AABB四倍体B基因组[Gressel.Molecular Biology inWeed Control(Taylor and Francis,London,2002)]的硬质小麦的多倍体水平没问题,实现杂草与硬质小麦之间的种间杂交就会难得多。同样,有可能由芥菜型油菜(Brassica juncea)(印度芥菜或棕芥菜)的B基因组向许多野生种的油菜(Brassica)杂草进行基因转移;但是,迄今为止大部分遗传工程对甘蓝型油菜(Brassica napus)实施,甘蓝型油菜具有AACC四倍体基因组,因此不可能相容。转基因性状扩散入杂草中的风险可通过仅释放具有不相容基因组的转基因系而急剧下降。
由于基因组信息的可用性,使得有可能通过不相容性机制工程改造出与杂草远交的可能性降低的作物。
时间和组织特异性控制-化学诱导启动子可用于基因防范策略。例如,可在除草剂喷洒大田前用化学物质诱导赋予除草剂抗性的基因瞬时表达。显然,通过将外源基因表达限制在作物未开花的时间有可能进行遗传隔离。这种启动子目前是可获得的(参见参考文献WO97/06269)。
仅在作物未处于开花期时启动外源基因的备选方法应是在开花前物理去除基因。Keenan和Stemmer[Nat.Biotechnol.20,215-216(2002)]提出,这可通过化学诱导型或果实特异性启动子来实现,以活化会在开花前切除外源基因的位点特异性重组酶如Cre的表达。这种系统可诱导Cre表达,导致位于种子(使用种子特异性启动子)或完整植株(使用化学诱导型启动子)中两个lox位点侧翼的基因的去除。
转基因缓和-另一种包含基因扩散的方法应是使杂草适合度失效,所述杂草通过基因渗入由作物基因获得了积极的生存性状[GresselTrends Biotechnol.17,361-366(1999)]。该方法称为转基因缓和(TM),基于以下前提:(1)串联构建物起紧密连锁基因的作用,其彼此的分离是极为罕见的;(2)TM性状对作物是中性的或积极的,但对杂草有害;和(3)即便温和有害的TM性状也由杂草群中去除,因为这种植物在其自身之间强烈竞争,具有大的种子产量。可由TM控制的过程的实例包括种子休眠、种子成熟和脱粒以及生长。
杂草种子通常表现出二次休眠,一次收获的种子在整个下一季和以后年份萌发,由此在降低同胞竞争的同时最大化适合度(和防止所有杂草遭受单一处理的控制)。二次休眠的废除对作物是中性的,但对杂草有害。Steber等已鉴别出一种拟南芥(Arabidopsis)突变体,其对脱落酸不敏感,完全没有二次休眠。这种与休眠相关的基因(工程过的或突变的)可用于TM[Genetics 149,509-521(1998)]。
杂草植物的另一个特征是其在一段时间内散播其种子,其成熟种子的大部分落粒至地面,确保连续性。结果,均一的成熟和抗落粒基因对杂草有害,但对作物是中性的,作物的种子均一成熟,不容易落粒;实际上,抗落粒基因对油菜甚至有利,油菜也具有落粒和自生杂草问题。只有无杂草的“有资格”种子才撒种,由此消除转基因杂草种子。一般认为,改变种子离层带的激素平衡影响落粒倾向。细胞因子过量生产可延迟落粒。近来已由拟南芥(Arabidopsis)分离出SHATTERPROOF基因,该基因通过延迟长角果上的瓣开裂防止种子落粒。这可能是用于密切相关的油菜的理想策略。
矮化由于产生水稻和小麦的“绿色革命”变化和为印度和中国带来粮食自足而特别有价值。但是,矮化性状对杂草是不利的,因为它们不能再与作物竞争光。通过防止赤霉素33的生物合成有可能得到遗传工程过的高度降低。另外,已分离出缺陷型赤霉酸报告基因,其通过与天然受体竞争赋予赤霉素不稳定性,由此诱导矮化。
启动子序列信息(例如SEQ ID NO:58)使得可通过修饰栽培植物的启动子序列产生本发明多肽表达增加的植物。因此,例如,“敲入”技术或诱变(例如化学或辐射)可用于直接或间接产生本发明多肽的表达被上调的植物。
要认识到的是,通过将本发明的cwp1基因定位到野生番茄的番茄染色体4,并精确作图至小于由端粒标记TG464延伸至着丝粒标记CT173的染色体片断的基因渗入,有可能使用经典育种技术产生角质层水透过性增加的栽培番茄植物。
例如,普通番茄植物可与野生番茄植物杂交。然后,令普通番茄植物的果实成熟,并收集杂合(F1)种子。然后,种植收集的F1种子,F1植株生长并令其自花受粉。自交可持续至少另一个代,或者F1植株可与普通番茄亲本植株杂交。使自交或回交代的果实在过了正式成熟点后仍保留在藤上,这由果实颜色的变化确定,并根据以下情况筛选:(i)存在天然脱水;和(ii)上述基因渗入。例如,通过使用以下标记CT199、TG163、CT61并在覆盖CT61和TG464的区域内,可将含野生种等位基因的最小基因渗入限于10cM以下、5cM以下、2cM以下和1cM以下。例如,可用于产生仍能增加角质层水透过性的最小基因渗入的标记包括源于图3a所示的BAC末端的任何序列。
因此,本发明还提供一种栽培番茄植物,其具有的基因组含来源于野生番茄的基因渗入,所述基因渗入含所述番茄的染色体4的一部分,小于由端粒标记TG464延伸至着丝粒标记CT173的染色体片断,所述基因渗入能够增加栽培番茄植物的角质层水透过性。
一旦产生了本发明的栽培的和遗传修饰的植物(如上所述),就可如下产生脱水果实。
让果实在过了正常成熟点之后仍保留在藤上。以果皮具皱为证的脱水外观表明果实中的水含量减少。一旦获得脱水果实就可收集它们。或者,由藤收集果实,随后令其脱水(例如在背景章节详述的晒干)。
因此,本发明提供在表达其的植物中控制角质层水透过性的多核苷酸和多肽,以及使用这些多核苷酸和多肽生产有商业价值作物的脱水果实的方法。
本文使用的术语“约”指±10%。
在检验以下的实施例时,本发明的其它目标、优势和新特征对本领域一般技术人员变得显而易见,这些实施例无限制意图。另外,如上文所描绘的和下文权利要求书章节要求保护的本发明的各种实施方案和方面各自都在以下实施例中得到实验支持。
实施例
现在论述以下实施例,其与上文描述一起以非限制性方式阐述本发明。
一般来说,本文使用的术语和本发明中使用的实验室方法包括分子、生物化学、微生物学和重组DNA技术。这种技术在文献中被详尽阐明。参见例如“Molecular Cloning:A laboratory Manual”Sambrook等,(1989);“Current Protocols in Molecular Biology”I-III卷Ausubel,R.M.编辑,(1994);Ausubel等,“Current Protocols inMolecular Biology”,John Wiley and Sons,Baltimore,Maryland(1989);Perbal,“A Practical Guide to Molecular Cloning”,John Wiley & Sons,New Yor(1988);Watson等,“Recombinant DNA”,Scientific AmericanBooks,New Yor;Birren等(编辑)“Genome Ahalysis:A LaboratoryManual Series”,1-4卷,Cold Spring Harbor Laboratory Press,New York(1998);如美国专利第4,666,828、4,683,202、4,801,531、5,192,659和5,272,057号所述的方法;“Cell Biology:A Laboratory Handbook”I-III卷Cellis,J.E.编辑,(1994);“Current Protocols in Immunology”I-III卷Coligan J.E.编辑,(1994);Stites等(编辑),“Basic and ClinicalImmunology”(第8版),Appleton & Lange,Norwalk,CT(1994);Mishell和Shiigi(编辑),“Selected Methods in Cellular Immunology”,W.H.Freeman和Co.,New York(1980);可用的免疫测定详尽描述于专利和科学文献,参见例如美国专利第3,791,932、3,839,153、3,850,752、3,850,578、3,853,987、3,867,517、3,879,262、3,901,654、3,935,074、3,984,533、3,996,345、4,034,074、4,098,876、4,879,219、5,011,771和5,281,521号;“Oligonucleotide Synthesis”Gait,M.J.编辑,(1984);“Nucleic Acid Hybridization”Hames,B.D.,和Higgins S.J.编辑,(1985);“Transcription and Translation”Hames,B.D.,和Higgins S.J.编辑,(1984);“Animal Cell Culture”Freshney,R.I.编辑,(1986);“Immobilized Cells and Enzymes”IRL Press,(1986);“A Practical Guideto Molecular Cloning”Perbal,B.,(1984)和“Methods in Enzymology”1-317卷,Academic Press;“PCR Protocols:A Guide To Methods AndApplications”,Academic Press,San Diego,CA(1990);Marshak等,“Strategies for Protein Purification and Characterization-A LaboratoryCourse Manual”CSHL Press(1996);所有这些文献都通过引用结合到本文中,如同其在本文完整陈述一样。其它一般参考文献贯穿本文件提供。其中的方法一般被认为是本领域周知的,提供其是为了方便读者。其中包含的所有信息都通过引用结合到本文中。
材料和方法
植物材料和检测-如本文概述的先前所述(WO 0113708)开发一组近等基因基因渗入系,其来源于回交育种程序,基于普通番茄(E)和野生种多毛番茄(H)的种间特异性杂交,通过果实脱水性状区分。E育种系1630的植物用野生种H(LA1777)授粉。杂合体F1植物自花授粉,产生F2种子。基于其成熟时的高糖含量选择3株F2植株。播种F3种子,培养这3株F2选择株的每个F3植物的10株植株,在过了正规成熟和收获期后仍让果实保留在藤上。在F3植物中,一个植株(F3-203-10)显示出果实脱水迹象的特征,以果皮起皱为证。开发谱系选择程序,其由自交该F3个体,直至在针对果实脱水率深度选择后获得F4代组成。此后,植物与E育种系回交,该杂交的产物对4个另一代自交,产生BC1F4群。该群的脱水个体与E进行另一次回交,产生具备该性状的杂合植物。由这些F1个体构建两个F2群(2394和2395)。
最初,选择步骤基于果实脱水和果实角质层上微裂隙的表型。在开发出与所述性状连锁的分子标记之后,按照基因型进行选择。酶切扩增多态性(CAPS)标记用作分子标记。使用由内切核酸酶切割的特异性PCR产物开发CAPS(参见下文进一步描述的“DNA分析”)。
如前所述(Miron和Schaffer,1991),植物在温室中按照标准方法在15-1盆中生长。如下测定果实平均重量和脱水率:由每株植株采摘和称重5个熟红果实,将它们置于室温(约25℃)的净重平台(net-table)上,每2-3天对其称重。通过放大镜(2X)或双目显微镜(10X)验证果实角质层上微裂隙(MF)的存在情况。
DNA分析-按照Fulton等(1995)提取基因组DNA。如下由选自高密度番茄图谱(Tanksley等,1992)的RFLP标记开发CAPS(酶切扩增多态性)标记。含代表选定RPLP标记的番茄DNA插入片段的BlueScript质粒载体(Stratagene)由Weizmann Institute of Science,Rehovot,Israel的Tomato Genome Center惠赠。在Hebrew University,Jerusalem,Israel的DNA Analysis Unit使用T7和SP6引物(分别为SEQID NO:1和2)对基因组DNA插入节段部分测序。按照这些序列分析结果,使用1.0版的Primer Express Program(Perkin Elmer Biosystems)设计序列特异性PCR引物。设计出总共约20个标记,检验这些标记,以确定普通番茄和多毛番茄亲本基因型之间以及多毛番茄来源性状不同的番茄系之间存在的多态性。
以下是两个标记TG163和TG587的PCR引物,代表染色体4上的位置。
TG163 F:5′-TGCAATCCCGAACATGAAGAC-3′(SEQ ID NO:3)
TG163 R:5′-CCTTCTGGTCGCATCTGTGTCT-3′(SEQ ID NO:4)
TG587 F:5′-TCAGGGTGAGGGGTAATAATTGAG-3′(SEQ ID NO:5)
TG587 F:5′-GCTTAAAACTCAAGTCTCCTCGCA-3′(SEQ ID NO:6)
在自动化热循环仪(Mastercycle Gradient,Eppendorf Germany)中使用热稳定性Taq DNA聚合酶(SuperNova Taq Polymerase,JMRProducts,Kent,UK)进行扩增反应。在含10×反应缓冲液、0.125mM的各脱氧核苷酸、0.5μ的各引物、2.5单位Taq聚合酶和50-100ng番茄基因组DNA的25μl终体积中进行反应。所述条件对各组引物的退火温度和产物片段大小优化。为鉴别应在普通番茄和多毛番茄等位基因之间产生多态性的限制性内切核酸酶,在含3.5μl PCR产物、1μl 10×浓缩限制性酶缓冲液和1-3单位的适宜限制性内切核酸酶的10μl终体积中进行反应。消化产物在1%凝胶上分析。发现DraI和HinFl分别对TG163和TG587适合,将其用于分离群。对其它CAPS标记的设计应用类似方法。
在该工作中使用的所有BACS(细菌人工染色体)都由ClemsonUniversity Genomic Institute(Clemson,North Carolina,USA)使用Tomato Heinz 1706 BAC Library Filters(LE_HBa)是供。如下所述通过放射性探针筛选Tomato BAC文库过滤器的特异性BAC克隆,所述放射性探针使用NEBlotTM试剂盒(New England BioLabs inc.#N1500S)并按照供应商的说明标记。使用磷成像装置(FLA-5000;FujiFilm)检测滤器上的标记BAC菌落。使用QIAGEN_Maxi Plasmid PurificationKit(#12263)由匹配的大肠杆菌菌株纯化BAC质粒。对于“染色体步查”法,使用SP6和T7引物测序BAC末端,并按照BAC末端序列开发PCR产物。新纯化的PCR产物进行放射性标记,并用于另一轮的番茄滤器菌落检测。
将LE_HBa 37B8 BAC克隆(Clemson University Genomic Institute,Clemson,North Carolina USA)亚克隆入BlueScript II ks+载体(Stratagene)中并测序。如上所述通过开发引物并通过PCR克隆以及测序相关部分来完整测序15kb部分。使用NCBI核酸和翻译蛋白数据库通过使用BLAST软件(Altschul等,1990)分析DNA序列。
RNA和定量RT-PCR分析-为制备cDNA,如前所述(Miron等,2002)提取总RNA。总RNA用作首链cDNA合成的模板,首链cDNA的合成采用Super-script II扩增前系统逆转录酶试剂盒(Gibco BRL,LifeTechnologies,UK)于42℃按照供应商的说明进行。
PCR引物-用1.0版的Primer Express Program(Perkin ElmerBiosystems),基于得自3个ORF的BAC测序的序列,设计具有用于在线定量PCR的短扩增子的特异性引物,所述3个ORF为:1)ZINC基因,正向,5′-AATAATGCGAATCGAATCACTA-3′(SEQ ID NO:7),和反向,5′-AAGGCTAAATCTCCTCCTTTCT-3′[SEQ ID NO:8,扩增子140bp(SEQ ID NO:9)],2)DBP基因,正向,5′-TGGATAAGCGGACGACTCTATTG-3′(SEQ ID NO:10),和反向,5′-CTGTTGTTTGGGAAGTGGCTTCT-3′[SEQ ID NO:11,扩增子116bp (SEQ ID NO:12)],3)PUT基因,正向,5′-CTCTCCTTGGCCCAAGGCTCAA-3′(SEQ ID NO:13),和反向,5′-CAGCTTTAGTGGTATCTCTCATCA-3′[SEQ ID NO:14,扩增子205bp(SEQ ID NO:15)]。肌动蛋白用作参比基因,使用基于Gene bank登录号BF096262的以下引物:正向,5′-CACCATTGGGTCTGAGCGAT-3′(SEQ ID NO:16),和反向,5′-GGGCGACAACCTTGATCTTC-3′[SEQ ID NO:17,扩增子251bp(SEQ ID NO:18)]。
cDNA用作定量PCR扩增的模板,PCR扩增在GeneAmp 5700序列检测系统(PE Biosystems)上使用含AmpliTaq Gold的SYBR GreenMaster Mix按照生产商(PE Biosystems)的说明进行。热循环仪的程序设定为所有反应40个循环,第一步为95℃、30秒变性,退火温度62℃、15秒,以及延伸温度72℃、30秒。数据采集于77℃、30秒进行。PCR产物的初步离散分析表明,在77℃以上保留的产物是特异性PCR产物。除了用于标准化的肌动蛋白引物以外,还用每组引物获得含对数增加的已知cDNA水平的标准曲线。所有实时PCR产物都在2%琼脂糖凝胶上检验,并送去测序以证明身份。
全长put基因的克隆-使用以下引物:Put正向,5′-GTAGTACTATATAAACCATGTGAG-3′(SEQ ID NO:19),和反向5′-CATATGTTGACATATCTAATG-3′(SEQ ID NO:20),由从HH系果实(开花后10天)中提取的cDNA扩增推定蛋白基因(put)的全长序列。使用T-A克隆法将全长基因[(SEQ ID NO:20),930bp)克隆至pGEM-T Easy Vector(promega),然后使用EcorI(NEB #R0101)内切核酸酶亚克隆入BlueScript II ks+载体(Stratagene)。再使用XhoI(NEB#R0146)和XbaI(NEB #R0145)内切核酸酶将put基因(SEQ ID NO:21)亚克隆在pBIN PLUS双元载体(Ghosh等,2002)的花椰菜35S启动子和n-终止子位点之间。
转基因植物-将含处于35S启动子控制下的put基因的构建载体转化入大肠杆菌(菌株DH5α,Stratagene)中,然后使用Walkerpeach和Velten(1994)描述的方法再转化入EHAl05农杆菌电感受态细胞中。使用Mini-Prep试剂盒(Qiagen#12143)制备质粒,并再转化入pBINPLUS中用于测序,以确保没有缺失和其它克隆错误。
使用如Meissner等(1997)描述的MicroTom栽培品种和如Barg等(1997)描述的MP1栽培品种进行番茄转化实验。将转基因插枝种植在补加1mgL-1玉米素(Duchefa #Z0917)、100mgL-1卡那霉素和100mgL-1 Chlaforan的Murashige和Skoog基础培养基(Duchefa,Haarlem,The Netherlands)上。执行培养转化植株的标准操作规程。
实施例1
脱水性状的遗传分析
基于标准小果栽培品种(1815系)和表现出脱水性状的远代基因渗入系(1881系)的杂交,在两个独立分离F5群(2394系和2395系)中测定果皮上微裂隙(MF)外观性状的遗传。分离群中微裂隙外观的分布模式符合3∶1的微裂隙:标准角质层比率,2394和2395群的卡方概率值分别为0.546和0.864(以下表1)。
表1:分离群2394和2395中微裂隙和脱水表型的分离模式
2395 | 2394 | ||||
表型 | 数量 | 概率 | 表型 | 数量 | 概率 |
N | 16 | 0.272 | N | 15 | 0.234 |
Y | 39 | 0.709 | Y | 49 | 0.765 |
总数 | 55 | 1.000 | 总数 | 64 | 1.000 |
X2值:0.029X2的概率:0.864 | X2值:0.424X2的概率:0.546 |
N-非脱水;Y-脱水;数量-群体中个体的数量
该分布模式特征为具有显性/隐性等位基因关联的单个基因遗传。
在2394群中果实脱水性状(CWP)按照3∶1比率分离,而在2395群中按照1∶2∶1比率分离,所述群约一半脱水,但以中间速度脱水。由此得出结论:等位基因关联是完全显性或者是半显性的,这取决于该群的遗传背景(图1a-b)。由上述可得出结论:果实CWP性状作为单一基因性状被遗传,其在本文被称为Cwp.-。
实施例2
cwp基因的精细作图
基于高密度番茄RFLP图谱(Tanksley等,1992)设计一组CAPS(酶切扩增多态性)标记。研究代表各种基因组位置的基因座,包括与网状表皮的QTL连锁的标记(Fulton等,2000,分别位于染色体4、6、8、12的标记TG464、TG477、CT68和TG68),用于分析与微裂隙性状的连锁。将基于各个多态性PCR的分子标记应用于两个亲本和一组48个按所述性状分离的F2个体。
基于初始标记组将Cwp基因作图到染色体4的端粒部分,以约3cM的估测距离与CT199标记连锁(在96个配子中有2个重组事件,图2a)。为更精细作图染色体4端粒部分,设计另一组CAPS标记作为定位于整个该染色体节段中的标记簇。多毛番茄亲本的染色体基因渗入节段位于CT163和TG464标记之间(图2b)。该基因渗入在近等基因系中代表多毛番茄节段,该近等基因系在该分析中用作脱水供体亲本。
为进一步缩窄基因渗入大小,用位于CT199和TG464标记之间的基于PCP的引物研究较大的F2群(超过200个个体)。将紧密连锁的分子标记簇(<1.5cM)定义为侧接Cwp基因(图2c),基于该研究,Cwp基因位于TG464和CT61之间(0.5cM)。
实施例3
cwp基因的定位克隆
该小基因渗入的定位使得可定位克隆Cwp。为此,采用标记TG464和CT61,对来源于近等基因系的杂合子个体的另3500个分离子代(7000个配子)进行CAPS标记分析,显示出12个重组体(0.34cM,相比之下在“第一轮”精细作图中相同标记之间为0.5cM)。使用染色体步查技术鉴别和组装桥接连锁标记TG464和CT61的一组5个连续BAC。简单地讲,这可通过测序BAC末端并使用BAC末端作为探针鉴别连续BAC来完成。
为了相对于基因渗入定位新BAC,并产生较高分辨率图谱,开发两个物种的多态性CAP,检验重组体的这些新标记。
5个连续BAC产生CT61和TG464 CAPS标记之间的桥(图3a)。对于12株重组植株中的每一个,生长10株自交子代,以合适的分离标记为基因型,分析其脱水和微裂隙外观。在12个初始鉴别的重组事件中,3个事件进一步定位于BAC 37B8的两个末端之间(图3a-由两条虚线限定的区域),表明Cwp位于37B8 BAC中。为进一步解析所述重组事件,亚克隆BAC 37B8,按顺序组装较小的片段,鉴别Cwp基因位于其中的约15,000bp(15kb)的节段。(图3b,未提供较低分辨率的图谱和亚克隆序列重叠群数据)。
实施例4
候选基因的生物信息学分析
将在实施例3中描述的BAC 37B8中的15kb节段亚克隆入Bluescript载体(Stratagene),测序并使用SEQUENCHER软件包(GeneCodes Corporation)组装。
在通过BLAST程序(BLASTP,NCBI,http://www.ncbi.nlm.nih.gov)分析后,15kb序列的生物信息学分析揭示了3个候选可读框(ORP,图4)。第一个ORF显示出与拟南芥(Arabidopsis thaliana)未知功能蛋白(GenBank登录号NP_189369.1)的相似性(蛋白同一性-44%,同源性-61%)。该蛋白具有两个结构域。第一个结构域是环指结构域(rpsBLAST-NCBI Conserved Domain Search),其是一种结合两个锌原子的40-60个残基的特化型锌指,可能涉及介导蛋白-蛋白相互作用(Borden,1998)。已在蛋白中鉴别出广泛的功能,例如病毒复制、信号转导和发育。其具有两个变体C3HC4型和C3H2C3型(RING-H2型锌指),它们具有不同的半胱氨酸/组氨酸模式。另一个结构域是DUF23,其是功能未知的结构域。其是由存在于线虫(C.elegans)和果蝇蛋白中的约300个残基长区域组成的家族的一部分。该区域包含几个保守半胱氨酸残基和几个可起催化残基作用的带电氨基酸。该ORF称为“Zinc”。令人感兴趣的是,番茄Zinc与拟南芥同系物的同源性不在“环指”部位,而是仅在DUF23一个部位,Zinc番茄基因没有“环指”结构域区。
第二个0RF显示出与结合DNA的含溴区结构域的蛋白(拟南芥GenBank登录号NP_74153.1)的相似性(蛋白同一性-37%,同源性56%)。该基因是与蛋白、DNA和染色质的乙酰化调节相关的DNA结合蛋白家族的一部分,该DNA结合蛋白家族是组蛋白乙酰转移酶调节的一部分(Dhalluin等,2000)。我们称该基因为“DBP”(DNA结合蛋白)。
第三个ORF与仅被描述为“表达蛋白”的蛋白(拟南芥At4g38260,GenBank登录号NP_568038.1)具有相似性(蛋白同一性-48%,同源性-67%)。其包含未知功能的结构域(DUF833)。其是存在于真核生物、原核生物和病毒中的一个家族的一部分,功能未知。已发现一个成员在小鼠的早期胚胎形成过程中被表达(Halford等,1993)。该基因称为“PUT”(推定的)。这3个候选基因中没有一个显示出与参与角质层生物合成机制已知步骤的基因的任何相似性或同源性。
实施例5
候选基因的表达分析
为确定3个候选基因中哪一个与番茄果实角质层发育相关,检测其Cwp等位基因不同的近等基因系中3个基因每一个的表达水平[多毛番茄脱水等位基因(HH),和普通番茄不脱水等位基因(EE),图5a-b]。由以下阶段的子房和果实提取mRNA:开花期,开花后5-15天,以及在生绿期、熟绿期和花端着色发育期(图5a-b)。果实标本取自用于定位克隆法的同一分离群。每个基因的表达都通过RT-PCR检验。DBP仅在子房期表达,在两个基因型(HH和EE)中相等,由此表明该基因的表达与表型性状无关(图5b)。Zinc基因的表达在任一个基因型的任何果实期均未观察到,同样表明其表达与脱水性状无关(未显示)。
只有PUT在发育中果实的幼年期表达,而且仅在具有Cwp的多毛番茄等位基因的脱水基因型(HH)果实中差异性表达(图5a)。在该研究中观察到的最高表达是在开花后15天的小果期。
为了证实PUT基因的差异表达模式,分析该基因在来源于M82番茄工业栽培品种的其余群中的表达。一个群是来源于杂合子个体的F2群,所述杂合子个体起源于脱水系(2168系)与M82确定栽培品种的杂交。我们检查了全部3种分离基因型(HH、HE、EE)在开花后5-15天时期(在第一个表达分析中具有最高表达水平的时期)的表达。如图6所示,发现了PUT基因的经典Mendelian表达模式,HH基因型显示出最高表达水平,杂合子HE个体显示出约一半的表达水平,EE基因型无表达(图6中的前3个条)。
另外,检查PUT基因在另一个NIL(近等基因系)群基因渗入系4.4中的表达,基因渗入系4.4来源于普通番茄(M82)和另一个野生种潘那利番茄的种间杂交,包含与本文所述来源于多毛番茄的基因型类似的基因渗入(Eshed和Zamir,1994)。该群代表PUT基因的另一个野生等位基因,IIA.4的果实还显示出微裂隙和脱水。与多毛番茄来源的群相似,含Cwp的潘那利番茄等位基因的潘那利番茄来源的基因渗入(IL 4.4)表明,与M82相比,PUT基因在在幼年小果中表达(图6,后两个条柱)。
表达PUT基因的转基因番茄植物
为了表明Put基因的表达与特有的角质层发育性状有关,开发具有处于35S启动子控制之下的PUT基因的转基因番茄植物(使用如所述的pBIN PLUS双元载体)。在转基因植物中观察到表型性状,表明Put的表达与所述性状相关。
为了确定来源于初始转基因植物自交的个体分离T1植株的基因剂量,将各个T1植株的50-70颗种子播种在含100mg/ml卡那霉素的1/2MS培养基中。在发芽后,测定具有正常根的幼苗的百分率。当100%幼苗表现出正常根生长时,T1植株被认为是转基因纯合子。约75%的T2幼苗具有正常根表明T1植株是转基因杂合子。尽管在本文中没有观察到其它比率,但其可能表明存在两个或多个未连锁的转基因拷贝。分析两个独立T1分离群的16个T1个体,以确定其等位基因组成。然后使用这些分类确定PUT基因的等位基因剂量和表型性状之间的关系。
如图7a-b所示,果实角质层上微裂隙(MF-)的表型性状已在T0代观察到。来自20个独立T0转基因个体的4株植株(MF1-1、MF1-4、MF1-8、MF1-12)表明,果皮上的MF水平可变。另外,这些转基因植物显示出高于野生型果实的脱水率(图7b)。
培养两个分离T1群,并测试其MF的存在情况和脱水率。图8a-b显示了PUT转基因拷贝数对微裂隙严重性(1-5之间记分,图8a)和果实失重百分率(于室温14天后,图8b)的作用。如在材料和方法章节中一样测定PUT基因拷贝数。
图9a-b显示了类似于野生型的无拷贝表达的转基因番茄个体(T1代)和表达两个拷贝的野生番茄种Solatium habrochaites S.的PUT基因的转基因番茄个体(T1代)之间的对比。图9a-扫描电子显微照片,呈现了无PUT基因拷贝(0拷贝)的个体果实和具有两个拷贝转基因的个体微裂隙果实的完整表面。图9b-无PUT基因拷贝(0拷贝)个体和具有两个拷贝的个体之间的脱水率比较。
这些结果清楚表明,PUT基因的表达是微裂隙和果实脱水表型的起因。
基于基因序列的系统发生分析表明,cwp是由拟南芥中3个成员代表的基因家族的一部分(图10a-b)。有另一个番茄同源物(CWP2)显示与Lecwp1基因30%同源,其实际上在栽培番茄中表达(EST No.AW621927)。
令人感兴趣的是,该同源物作图至番茄染色体2-1,在番茄染色体2-1中有番茄果实表皮网纹的报告QTL(Frary等,2004)。茄属栽培胡椒(辣椒(Capsicum annum))的发育中果实也在其表皮组织中表达与Lecwp1基因高度相似(87%)的cwp同源物,胡椒果实的特征在于收获后失水的园艺问题以及红辣椒栽培品种中果实脱水的需要性状。因此,CWP基因的同源物还有可能有助于角质层修饰和水透过性。
这些结果表明,cwp基因的表达产生结构修饰的角质层(基于重量和TEM),推测其在果实膨大过程中由于弹性下降而经历缝裂。但是,该现象仅在具有高度发育的果实角质层的果实中观察到,例如无气孔厚皮栽培番茄,在以较薄角质层为特征的野生种果实中不明显。在栽培番茄果实发育过程中角质层组分的沉积在由生绿期向绿熟期的转变过程中出现突然上升,这是合理的,因为这与微裂隙表型的观察结果一致。
要认识到的是,为清楚起见而在单独的实施方案背景中描述的本发明某些特征也可与单个实施方案组合提供。相反,为简短起见而在单个实施方案背景中描述的本发明各种特征也可单独地或以任何合适的次组合提供。
尽管已描述了本发明连同其具体实施方案,但显然许多备选、修改和变更对本领域技术人员显而易见。因此,有意包含所有这些属于所附权利要求书的精神和广泛范围的备选、修改和变更。在本说明书中提及的所有出版物、专利和专利申请以及GenBank登录号都通过引用整体结合到本说明书中,其程度如同各个单独的出版物、专利或专利申请或GenBank登录号被明确和个别指出通过引用结合到本文中。另外,在本申请中的任何参考文献的援引或标识都不应被解释为认可该参考文献可作为本发明的先有技术使用。
参考文献
(本申请中提及的其它参考文献)
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gaatgacgag aaaatctgcg tcaaagatat gatagaaaaa ttgatgagag ataccactaa 2520
agctgataaa agtaaattgc cttgtatttg ttctacagac tgggagttgg aacttagctc 2580
tattttcgtg gaagttgaca ctgcactggg taattcatac cgcgttataa ctaatatgtt 2640
tgtttgattt taacgtactc aaacgatgat aaaggttaaa gtagatatac aaacatttta 2700
aaaataattg aaatagttca ataatagaag tgtacatatc attaacatag tttgatgggt 2760
ttttttggtg gtgtgaatat gtaggggtgt tatggtacta gaagtacaac agcattgaca 2820
attgaagtgg gaggagaagt aagcttttat gagttgtacc ttgagaacaa catgtggaaa 2880
gagcaaattg tcaactatcg gattgaaaaa ctccaaatgc aataaatgtt tttaatatgt 2940
tgatatatct aatgtttttc atgttcatat gttgacatat ctaatgtttt catttttttt 3000
ttttaattca aataagattt tttcttcaaa aaattaaact ttttgtcttt gaatggaaat 3060
tgttattcat tgtatttgta aaatgtacta cactacttgg aagacataat gtatgtttcg 3120
ggctcctttg ttttagcaac aattttagac tttca 3155
<210>24
<211>750
<212>DNA
<213>多毛番茄(Lycopersicon hrisutum)
<400>24
atgtctatac cggtgttcat atggaaagcg catccgttgc atcccttcct tctcttcctc 60
aacagagatg aataccacaa tcgtccaacg aaaccattgt catggtggga agatactgat 120
atacttggtg gaagggatga agttgctggt gggacttggt tggcttgtac tcgcactgga 180
agacttgctt ttcttactaa tgttcgagaa atcaattcaa attcacatac cagaagtagg 240
ggagaccttc ctcttcgatt cttaaagagt gtgaagagcc ctcgtgattt ttcagagcaa 300
ctattgatag aagcaggtga atataatggg tttaatttga tagtaactga tctttgttca 360
atgactatgc tatatataac taaccgaccg aaacacacg gtatgtccgt cactgaggtt 420
tcacccggta ttcatgtttt atcaaatgca tcactaaact ctccatggcc taagtctcaa 480
cggctggagt gcagtttcaa gcaattattg gatgaatatg gcgaatcgga aattccaata 540
gggcatgcag ctgaaagaat atgagagacg tggctcaaga agatagtaac ccgccaggca 600
tcatattctc ccgagtgtga gtaccaattg agctccctat ttgttgacac tgaaatgtgc 660
atggggcgtt tttgcccaag aagcacttct tcactggccg tgaagaagtc ttgtgacgcc 720
accttttatg agcggttcct gagaaggttt 750
<210>25
<211>250
<212>PRT
<213>多毛番茄(Lycopersicon hrisutum)
<400>25
Met Ser Ile Pro Val Phe Ile Trp Lys Ala His Pro Leu His Pro Phe
1 5 10 15
Leu Leu Phe Leu Asn Arg Asp Glu Tyr His Asn Arg Pro Thr Lye Pro
20 25 30
Leu Ser Trp Trp Glu Asp Thr Asp Ile Leu Gly Gly Arg Asp Glu Val
35 40 45
Ala Gly Gly Thr Trp Leu Ala cys Thr Arg Thr Gly Arg Leu Ala Phe
50 55 60
Leu Thr Asn Val Arg Glu Ile Asn Ser Asn Ser His Thr Arg Ser Arg
65 70 75 80
Gly Asp Leu Pro Leu Arg Phe Leu Lys Ser Val Lys Ser Pro Arg Asp
85 90 95
Phe Ser Glu Gln Leu Leu Ile Glu Ala Gly Glu Tyr Asn Gly Phe Asn
100 105 110
Leu Ile Val Thr Asp Leu Cys Ser Met Thr Met Leu Tyr Ile Thr Asn
115 120 125
Arg Pro Lys His Thr Gly Met Ser Val Thr Glu Val Ser Pro Gly Ile
130 135 140
His Val Leu Ser Asn Ala Ser Leu Asn Ser Pro Trp Pro Lys Ser Gln
145 150 155 160
Arg Leu Glu Cys Ser Phe Lys Gln Leu Leu Asp Glu Tyr Gly Glu Ser
165 170 175
Glu Ile Pro Ile Gly His Ala Ala Glu Arg Ile Met Arg Asp Val Ala
180 185 190
Gln Glu Asp Ser Asn Pro Pro Gly Ile Ile Ser Pro Glu Cys Glu Tyr
195 200 205
Gln Leu Ser Ser Leu Phe Val Asp Thr Glu Met Cys Met Gly Arg Phe
210 215 220
Cys Pro Arg Ser Thr Ser Ser Leu Ala Val Lys Lys Ser Cys Asp Ala
225 230 235 240
Thr Phe Tyr Glu Arg Phe Leu Arg Arg Phe
245 250
<210>26
<211>762
<212>DNA
<213>拟南芥(Arabidopsis thaliana)
atgaagatca caacagggcg acagaggcgc tgcgtttggt gggaagacgg agagacggtg 60
ggaggaagag accttgttgg cggcgggacg tggctgggct gcacgaggca tggccgtctg 120
ggtttcctca ccaatttcaa ggaagcctcc tccttccctg ctgctaaatc ccgtggagat 180
ctgcctcttc gttacttgca gagcgaaaag agtccggccg agtttgccga ggagatccaa 240
gacgaaattt cactctacaa tggctttaac ctggttgtcg ctaatgtctt gtccaaatcc 300
atgatttaca ttaccaaccg accaccccac ggtgacaagc tcgtgacgca agtctctccc 360
gggatccatg tcctttccaa cgccaacctc gactcccctt ggcccaagtg tctgaggctg 420
agggagggtt tccaacagct tctggatgag aacgggagcg gtgaattccc ggtgaagacc 480
atggtggagg aggtgatgac caatactgtc aaggacgaag aaaccgagct acctcacgtt 540
ttcacaccag agacggaata ccatctcagc tccatcttcg tcgacatgca gagaccaact 600
gggcgttatg ggaccagaag catctctgcg atcatcgtca agtcccatgg agatggtggt 660
ggtgatggtg agatttgctt ctacgagagg catcttgaag aaggcgatto atggaaggaa 720
cacactcaac agtttgtaat aatacaaaac caaagcattt ga 762
<210>27
<211>253
<212>PRT
<213>拟南芥(Arabidopsis thaliana)
<400>27
Met Lys Ile Thr Thr Gly Arg Gln Arg Arg Cys Val Trp Trp Glu Asp
1 5 10 15
Gly Glu Thr Val Gly Gly Arg Asp Leu Val Gly Gly Gly Thr Trp Leu
20 25 30
Gly Cys Thr Arg His Gly Arg Leu Ala Phe Leu Thr Asn Phe Lys Glu
35 40 45
Ala Ser Ser Phe Pro Ala Ala Lys Ser Arg Gly Asp Leu Pro Leu Arg
50 55 60
Tyr Leu Gln Ser Glu Lys Ser Pro Ala Glu Phe Ala Glu Glu Ile Gln
65 70 75 80
Asp Glu Ile Ser Leu Tyr Asn Gly Phe Agn Leu Val Val Ala His Val
85 90 95
Leu Ser Lys Ser Net Ile Tyr Ile Thr Asn Arg Pro Pro His Gly Asp
100 105 110
Lys Leu Val Thr Gln Val Ser Pro Gly Ile His Val Leu Ser Asn Ala
115 120 125
Asn Leu Asp Ser Pro Trp Pro Lys Cys Leu Arg Leu Arg Glu Gly Phe
130 135 140
Gln Gln Leu Leu Ala Glu Asn Gly Ser Gly Glu Phe Pro Val Lys Thr
145 150 155 160
Met Val Glu Glu Val Met Thr Asa Thr Val Lys Asp Glu Glu Thr Glu
165 170 175
Leu Pro His Val Phe Thr Pro Glu Thr Glu Tyr His Leu Ser Ser Ile
180 185 190
Phe Val Asp Met Gln Arg Pro Thr Gly Arg Tyr Gly Thr Arg Ser Ile
195 200 205
Ser Ala Ile Ile Val Lys Ser His Gly Asp Gly Gly Gly Asp Gly Glu
210 215 220
Ile Cys Phe Tyr Glu Arg His Leu Glu Glu Gly Asp Ser Trp Lys Glu
225 230 235 240
His Thr Gln Gln Phe Val Ile Ile Gln Asn Gln Ser Ile
245 250
<210>28
<211>801
<212>DNA
<213>拟南芥(Arabidopsis thaliana)
<400>28
atggggagag ggagaaaaca cactgacgct gctgcagaac agagagaact ggcaattaag 60
gcaaacattg ttgatgaacc tttttatgta toggcgattg ataggtaaat aagaaaggcg 120
gaatgggtta aaactgaaac tgaccagata ttaagtggtc gttgccaaga gaccgatggg 180
acgtggttag gtatttctac tcgaggccga gtcgcttttc ttgtggaggc agggactatt 240
aacagagaca gattcaacgg cgccgagagt cgtactcttg agttcttaga gagcaacgag 300
agtccagagg actttgcaaa gtcatcggct gcagattaca tacgtaacaa gaacacagcc 360
gcctttcatc taattgtggc cgacatagct tcaaactcaa tgctttatat ctccaaaccg 420
cgtttctctg actatggcat tgtctataca gagcctgttg gtcctggtgt tcacacacta 480
tcttcagctg gactcgattc cgacgttgga tacagggact tacgtatgag acactctttt 540
tgtgagatga ttaacagaga acgactacca ccaataaggg acattgctga gattatgtat 600
gatccagtca aagcttacga aagcgtgcta ctgagctcta tttttttcgt cgacatgaag 660
attggatacg aacactatgg aacaagaatt acgacagcat tggttgtgaa acgcaccaag 720
gaagtgttgt tctttgagag gtacagggag atatttaatg atgattggga cgaccacgac 780
ttcgcgttca ccatcatcta g 801
<210>29
<211>266
<212>PRT
<213>拟南芥(Arabidopsis thaliana)
<400>29
Met Gly Arg Gly Arg Lye Hie Thr Asp Ala Ala Ala Glu Gln Arg Gla
1 5 10 15
Leu Ala Ile Lys Ala Asn Ile Val Asp Glu Pro Phe Ser Val Ser Ala
20 25 30
Ile Asp Arg Ser Ile Arg Lys Ala Glu Trp Val Lye Thr Glu Thr Asp
35 40 45
Gln Ile Leu Ser Gly Arg Cys Pro Glu Thr Asp Gly Thr Trp Leu Gly
50 55 60
Ile Ser Thr Arg Gly Arg Val Ala Phe Leu Val Glu Ala Gly Thr Ile
65 70 75 80
Asn Arg Asp Arg Phe Asn Gly Ala Glu Ser Arg Thr Leu Glu Phe Leu
85 90 95
Glu Ser Asn Glu Ser Pro Glu Asp Phe Ala Lye Ser Ser Ala Ala Asp
100 105 110
Tyr Ile Arg Asn Lye Asn Thr Ala Ala Phe His Leu Ile Val Ala Asp
115 120 125
Ile Ala Ser Asn Ser Met Leu Tyr Ile Ser Lys Pro Arg Phe Ser Asp
130 135 140
Tyr Gly Ile Val Tyr Thr Glu Pro Val Gly Pro Gly Val Hia Thr Leu
145 150 155 160
Ser Ser Ala Gly Leu Asp Ser Asp Val Gly Tyr Arg Asp Leu Arg Met
165 170 175
Arg His Ser Phe Cye Glu Met Ile Asn Arg Glu Arg Leu Pro Pro Ile
180 185 190
Arg Asp Ile Ala Glu Ile Met Tyr Asp Pro Val Lya Ala Tyr Glu Ser
195 200 205
Val Leu Leu Ser Ser Ile Phe Phe Val Asp Met Lys Ile Gly Tyr Glu
210 215 220
His Tyr Gly Thr Arg Ile Thr Thr Ala Leu Val Val Lys Arg Thr Lys
225 230 235 240
Glu Val Leu Phe Phe Glu Arg Tyr Arg Glu Ile Phe Asn Asp Asp Trp
245 250 255
Asp Asp His Asp Phe Ala Phe Thr Ile Ile
260 265
<210>30
<211>1122
<212>DNA
<213>拟南芥(Arabidopsis thaliana)
<400>30
atgcctggag aatcgaatat catcgagtgg ccagcaagta gagtcagggt cataagtggg 60
gcatcttgga gtcgaaacgg tcagattttg agtggtcggt gcaaagctaa caacggaacc 120
tggtttggta ttactaaagg tggccgagtc gcttttctcg tgaatacatc gttgttgttg 180
gaccgtgtta agtcatacag cggctcggag ttgtatcccg ttcgtttctt ggagggcaac 240
atgagtccag agcagtttgc caacgaagtg aaagtgcatg aaaaggagac taatgaaagg 300
catgcctata gtcttgtcgt tgcagacatg acttcgagtt caatggttca tatcctgaaa 360
ccctcggata ctaagtctga tgtcgtgata gagactgttc cgtttggtgt gcatacactt 420
tcttcttacg aaggtctcga ttctacagat tctgccaggg atttactcct gagacgcttg 480
tttacccaga tggttggtaa tttgggaaac gttcaacaac gacagatgga ggagattgct 540
gggaggttta tgtatgatgc tcaagcagga agagacgcgg tgttttacca tagtagagat 600
gagcatccta atggaaaact tggaacgcaa cgctttggaa caacaagtac gacagcattg 660
gttgtgaaac gcactagaga agtgatgctc tttgagaagt acatggagca gaatggtgca 720
tggaacacga acaacttcgc tttcaacatc caaaaacagc aaaagctata tccaaatttg 780
gataaagaag ctcttaagcg cgttggggta tttgcgttgg aagaagttaa caaccatgag 840
catgatattc accctgacct gatgcccagt ttcttcgagg atgatatgct gaaagtaaaa 900
tttaatgaga tgattgctag acatgcaaaa ctgccgccaa ttaagaacat tgttgaggat 960
cttatgatga agtctccatt ttttatcgac agtgtcgatg gtgctggcaa gaaggtgagg 1020
tatcgaacag tacgtacatt gggaatggac ataaaagcca acagaccaca agggcggttc 1080
tatgagaggc atttgaatga taatggtgaa tgggtaggct ag 1122
<210>31
<211>373
<212>PRT
<213>拟南芥(Arabidopsis thaliana)
<400>31
Met Pro Gly Glu Ser Asn Ile Glu Trp Pro Ala Ser Arg Val Arg
1 5 10 15
Val Ile Ser Gly Ala Ser Trp Ser Arg Asn Gly Gln Ile Leu Ser Gly
20 25 30
Arg Cys Lys Ala Asn Aan Gly Thr Trp Phe Gly Ile Thr Lys Gly Gly
35 40 45
Arg Val Ala Phe Leu Val Asn Thr Ser Leu Leu Leu Asp Arg Val Lys
50 55 60
Ser Tyr Ser Gly Ser Glu Leu Tyr Pro Val Arg Phe Leu Glu Gly Asn
65 70 75 80
Met Ser Pro Glu Gln Phe Ala Asn Glu Val Lys Val His Glu Lys Glu
85 90 95
Thr Asn Glu Arg His Ala Tyr Ser Leu Val Val Ala Asp Met Thr Ser
100 105 110
Ser Ser Met Val His Ile Leu Lys Pro Ser Asp Thr Lys Ser Asp Val
115 120 125
Val Ile Glu Thr Val Pro Phe Gly Val His Thr Leu Ser Ser Tyr Glu
130 135 140
Gly Leu Asp Ser Thr Asp Ser Ala Arg Asp Leu Leu Leu Arg Arg Leu
145 150 155 160
Phe Thr Gln Met Val Gly Asn Leu Gly Asp Val Gln Gln Arg Gln Met
165 170 175
Glu Glu Ile Ala Gly Arg Phe Met Tyr Asp Ala Gln Ala Gly Arg Asp
180 185 190
Ala Val Phe Tyr His Ser Arg Asp Glu His Pro Asn Gly Lys Leu Gly
195 200 205
Thr Gln Arg Pha Gly Thr Thr Ser Thr Thr Ala Leu Val Val Lya Arg
210 215 220
Thr Arg Glu Val Met Leu Phe Glu Lys Tyr Met Glu Glu Asn Gly Ala
225 230 235 240
Trp Asn Thr Asn Asn Phe Ala Phe Asn Ile Gln Lys Gln Gln Lys Leu
245 250 255
Tyr Pro Asn Leu Asp Lys Glu Ala Leu Lys Arg Val Gly Val Phe Ala
260 265 270
Leu Glu Glu Val Asn Asn His Glu His Asp Ile His Pro Asp Leu Met
275 280 285
Pro Ser Phe Phe Glu Asp Asp Met Leu Lys Val Lys Phe Asn Glu Met
290 295 300
Ile Ala Arg His Ala Lys Leu Pro Pro Ile Lys Asn Ile Val Glu Asp
305 310 315 320
Leu Met Met Lys Ser Pro Phe Phe Ile Asp Ser Val Asp Gly Ala Gly
325 330 335
Lys Lys Val Arg Tyr Arg Thr Val Arg Thr Leu Gly Met Asp Ile Lys
340 345 350
Ala Asn Arg Pro Gln Gly Arg Phe Tyr Glu Arg His Leu Asn Asp Aan
355 360 365
Gly Glu Trp Val Gly
370
<210>32
<211>713
<212>DNA
<213>葡萄(Vitis vinifera)
<400>32
ggtcgcactg tatatattca acggagagga gcagtgacgg cgtttggagg ccgagaaagt 60
aagagatttc agtttctgag gcgggaaagt acggaagcat gtgtatagca gtattcttat 120
ggcaagctca cccgatttat cctttccttc tgttgctcaa cagagacgaa tatcataatc 180
ggcctactga ggctctggca tggtggcaag gtggggagat actgggcggg cgagatgggc 240
tcgccggtgg gacatggttg gcttgtagca gagatgggag gttggctttt cttacaaatg 300
tgcgagaagt tcacccaatc cccgaagcca agagcagagg agacctaatt gttcggttct 360
tggagagcaa gaagaatccc atggaatttg cagaggaagt tgtgaaggag gcagataagt 420
ataatgggtt taacttgata atggctgatc tttgttccaa aactatgatc tatataacca 480
acagaccaag agaagctaat gtttctgttg tcgaggtttc acctggtatt catgtgctgt 540
caaatgcaag tttggactca ccttggccta aggtacgaag actaggtcat aatttcaaag 600
agctcttgga taaatatggt gaaggtgaga tccccacaga ggagatggtt gagaaattaa 660
tgaagaaaca caatcaaaga cgatgaaatc gtgctgcctc gcatctatcc tcc 713
<210>33
<211>207
<212>PRT
<213>葡萄(Vitis vinifera)
<400>33
Glu Ile Ser Val Ser Glu Ala Gly Lys Tyr Gly Ser Met Cys Ile Ala
1 5 10 15
Val Phe Leu Trp Gln Ala His Pro Ile Tyr Pro Phe Leu Leu Leu Leu
20 25 30
Asn Arg Asp Glu Tyr His Asn Arg Pro Thr Glu Ala Leu Ala Trp Trp
35 40 45
Gln Gly Gly Glu Ile Leu Gly Gly Arg Asp Gly Leu Ala Gly Gly Thr
50 55 60
Trp Leu Ala Cys Ser Arg Asp Gly Arg Leu Ala Phe Leu Thr Asn Val
65 70 75 80
Arg Glu Val His Pro Ile Pro Glu Ala Lys Ser Arg Gly Asp Leu Ile
85 90 95
Val Arg Phe Leu Glu Ser Lys Lys Asn Pro Mer Glu Phe Ala Glu Glu
100 105 110
Val Val Lys Glu Ala Asp Lys Tyr Aen Gly Phe Asn Leu Ile Met Ala
115 120 125
Asp Leu Cys Ser Lys Thr Met Ile Tyr Ile Thr Asn Arg Pro Arg Glu
130 135 140
Ala Asn Val Ser Val Val Glu Val Ser Pro Gly Ile His Val Leu Ser
145 150 155 160
Asn Ala Ser Leu Asp Ser Pro Trp Pro Lys Val Arg Arg Leu Gly His
165 170 175
Asn Phe Lys Glu Leu Leu Asp Lye Tyr Gly Glu Gly Glu Ile Pro Thr
180 185 190
Glu Glu Met Val Glu Lys Leu Met Lys Lys His Asn Gln Arg Arg
195 200 205
<210>34
<211>1068
<212>DNA
<213>玉米(Zea mays)
<400>34
gatcagctaa gatagctgca aaacaagcga gttacttaca accaaacaga agggtagaaa 60
ccacctgaag ccatgtgcat tgctgcatgg atttggcagg ctcaccctgt gcaccaactc 120
ctcctgcttc tcaacagaga tgagttccac agcaggccta caaaagcagt aggatggtgg 180
ggtgaaggct caaagaagat ccttggtggc agggatgtgc ttggtggagg aacatggatg 240
gggtgcacca aggatggaag gcttgccttc ctgaccaatg tgcttgaacc agatgccatg 300
cccggtgcac ggactagggg agatctgcct ctcaaattcc tgcagagcaa caagagccca 360
ctcgaagttg caactgaagt ggcagaagaa gctgatgaat acaatggctt caacctcata 420
ctagctgatc taacaacaaa tatcatggtt tatgtgtcaa accggcctaa gggtcagcct 480
gcaacaattc aactcgtgtc accaggactc catgtgctgt ccaatgcaag gctagatagc 540
ccttggcaga aggcaattct cctcggtaaa aacttcaggg agcttcttag ggagcatggt 600
gctgatgagg ttgaagtgaa ggatatagtt gagaggctaa tgactgacac cacaaaggct 660
gacaaagata gactgccaaa cactggttgt gatcccaact gggagcatgg tctgagctcc 720
atcttcattg aggtgcaaac tgaccaaggg ccctatggga cacggagcac agccgtttta 780
tcagtgaact atgatggcga agctagcttg tacgagaagt atcttgagag tggtatatgg 840
aaggatcaca cagtgagtta ccagatagag tagtaggcat tgcacaggaa aagttggcga 900
cctcaaataa atagaaatat gaagcagaca caattgtgaa tttcattatt tccctgatct 960
ctagtcatct tcgtgattat ctaagatcct accataatgc caattacatt attcactgta 1020
agcagatttt tcacttgacg ataaaatgtc aaccaaaact ttggtttt 1068
<210>35
<211>271
<212>PRT
<213>玉米(Zea mays)
<400>35
Lys Pro Pro Glu Ala Met Cys Ile Ala Ala Trp Ile Trp Gln Ala His
1 5 10 15
Pro Val His Gln Leu Leu Leu Leu Leu Asn Arg Asp Glu Phe His Ser
20 25 30
Arg Pro Thr Lys Ala Val Gly Trp Trp Gly Glu Gly Ser Lys Lys Ile
35 40 45
Leu Gly Gly Arg Asp Val Leu Gly Gly Gly Thr Trp Met Gly Cys Thr
50 55 60
Lys Asp Gly Arg Leu Ala Pha Leu Thr Asn Val Leu Glu Pro Asp Ala
65 70 75 80
Met Pro Gly Ala Arg Thr Arg Gly Asp Leu Pro Leu Lys Phe Leu Gln
85 90 95
Ser Asn Lys Ser Pro Leu Gul Val Ala Thr Glu Val Ala Glu Glu Ala
100 105 110
Asp Glu Tyr Asn Gly Phe Asn Leu Ile Leu Ala Asp Leu Thr Thr Asn
115 120 125
Ile Met Val Tye Val Ser Asn Arg Pro Lys Gly Gln Pro Ala Thr Ile
130 135 140
Gln Leu Val Ser Pro Gly Leu His Val Leu Ser Asn Ala Arg Leu Asp
145 150 155 160
Ser Pro Trp Gly Lys Ala Ile Leu Leu Gly Lys Asn Phe Arg Glu Leu
165 170 175
Leu Arg Glu His Gly Ala Asp Glu Val Glu Val Lys Asp Ile Val Glu
180 185 190
Arg Leu Met Thr Asp Thr Thr Lys Ala Asp Lys Asp Arg Leu Pro Asn
195 200 205
Thr Gly Cys Asp Pro Asn Trp Glu His Gly Leu Ser Ser Ile Phe Ile
210 215 220
Glu Val Gln Thr Asp Gln Gly Pro Tyr Gly Thr Arg Ser Thr Ala Val
225 230 235 240
Leu Ser Val Asn Tyr Asp Gly Glu Ala Ser Leu Tyr Glu Lys Tyr Len
245 250 255
Glu Ser Gly Ile Trp Lys Asp His Thr Val Ser Tyr Gln Ile Glu
260 265 270
<210>36
<211>1152
<212>DNA
<213>大麦(Hordeum vulgare)
<400>36
cacacacaca caaggcgcac ggttgcaaaa caagggagtt atttttagaa gcagggagta 60
aggaaccacc tgaagccatg tgtatcgctg catggcattt ggcaggctca cccacagcat 120
cagctcctgc tgctgctcaa cagagatgag ttccatagca ggcctacaaa ggcagtagga 180
tggtggggcg agggctcaat gaagattctt ggtggcaggg atgtactcgg tggaggaaca 240
tggatgggga gcaccaaaga tggcaggctt gccttcctga ccaatgtgct cgagcctgat 300
gcaatgcccg gcgcacgcac taggggagac ctgcccctca ggttcctgca gggaaacaag 360
agcccactgg aggttgcgac tgaagtggca aaagaagctg atgagtacaa tggcttcaac 420
cttatactag ctgatctaac caggaatgtc atggtctacg tgtcaaaccg gccaaagggg 480
cagcctgcga caattcagct cgtctcacca ggactccatg tgttgtccaa tgcaaggctt 540
gatagccctt ggcagaaggc aattcgcctt ggtaaaaact tcagggagtt tataaggaag 600
catggtgatg atgaagttga agcgaaggat atagctgaca gactaatgac tgacacgacg 660
agggctgata aagataggct gccaaacacc ggttgtgatc ccacctggga gcacggtctg 720
agctccatct tcatcgaggt gcaaactgac gaagggctct atgggacaag gagcacagca 780
gttctttcag tgaactatga tggagaagct agcttatatg aaaagtacct cgagagtggt 840
atatggaaga accacacagt gcattaccag atagagtagc caatgcggac ctaaaggcgg 900
gagcccaaaa taggaagaaa gaatgaatag ctacaattgt gcatgctgtt atttccacag 960
ttgcgcttta agatcatata atgatctcta gttatggcga ttaaattatt tactgtatgc 1020
agatttatca attcagagag agatcattca aattgttgaa tatatataca taataataat 1080
aatatgatat gatatgtata tttacagact tcatgttgcc acctttgtct atgaacatac 1140
atgctttact ac 1150
<210>37
<211>291
<212>PRT
<213>大麦(Hordeun vulgare)
<400>37
Lys Gln Gly Val Arg Asn His Leu Lys Pro Cys Val Ser Leu His Gly
1 5 10 15
Ile Trp Gln Ala His Pro Gln His Gln Leu Leu Leu Leu Leu Asn Arg
20 25 30
Aap Glu Phe His Ser Arg Pro Thr Lys Ala Val Gly Trp Trp Gly Glu
35 40 45
Gly Ser Met Lys Ile Leu Gly Gly Arg Asp Val Leu Gly Gly Gly Thr
50 55 60
Trp Met Gly Ser Thr Lys Asp Gly Arg Leu Ala Phe Leu Thr Asn Val
65 70 75 80
Leu Glu Pro Asp Ala Met Pro Gly Ala Arg Thr Arg Gly Asp Leu Pro
85 90 95
Leu Arg Phe Leu Gln Gly Asn Lys Ser Pro Leu Glu Val Ala Thr Glu
100 105 110
Val Ala Lys Glu Ala Asp Glu Tyr Asn Gly Phe Asn Leu Ile Leu Ala
115 120 125
Asp Leu Thr Arg Asn Val Met Val Tyr Val Ser Asn Arg Pro Lys Gly
130 135 140
Gln Pro Ala Thr Ile Gln Leu Val Ser Pro Gly Leu His Val Leu Ser
145 150 155 160
Asn Ala Arg Leu Asp Ser Pro Trp Gln Lys Ala Ile Arg Leu Gly Lys
165 170 175
Asn Phe Arg Glu Phe Ile Arg Lys His Gly Asp Asp Glu Val Glu Ala
180 185 190
Lys Asp Ile Ala Asp Arg Leu Met Thr Asp Thr Thr Arg Ala Asp Lys
195 200 205
Asp Arg Leu Pro Asn Thr Gly Cys Asp Pro Thr Trp Glu His Gly Leu
210 215 220
Ser Ser Ile Phe Ile Glu Val Gln Thr Asp Glu Gly Leu Tyr Gly Thr
225 230 235 240
Arg Ser Thr Ala Val Leu Ser Val Asn Tyr Asp Gly Glu Ala Ser Leu
245 250 255
Tyr Glu Lys Tyr Leu Glu Ser Gly Ile Trp Lys Asn His Thr Val His
260 265 270
Tyr Gln Ile Glu Pro Met Arg Thr Arg Arg Glu Pro Lys Ile Gly Arg
275 280 285
Lys Asn Glu
290
<210>38
<211>801
<212>DNA
<213>水稻(Oryza sativa)
<400>38
atgtgtatag ctgcatgggt ttggcaagct cacccacagc accagctcct cctgctgctc 60
aaccgggatg agttccatag caggccaacc aaggcagtag gatggtgggg ggagggctcg 120
aagaagattc ttggtggtag agatgttctt ggtggaggga catggatggg ttgcacaaag 180
gatggcaggc tcgccttcct caccaatgtg ctcgagccgg acgccatgcc gggggcgcgc 240
acaaggggag atctccccct caggttcctg cagagcaaca agagcccact tgaagttgca 300
actgaggtgg caaaagaagc tgacgagtac aacggcttca accttgtact ggctgatctg 360
accacaaacg tcatggttta tgtgtcaaat cggccaaagg ggcagcctgc aacgatccaa 420
cttgtctcac cagggctcca tgtgttgtcc aatgcaaggc cagacagccc ttggcagaag 480
gcgattcgcc tcggtaagaa cttcagggag catcttagga agcatggtga tgatgaggtt 540
gaagccaagg acatagttga gaggctaatg actgacacca caaaggctga caaagatagg 600
ctgccaaaca ctggctgtga tccaaactgg gagcacggcc tgagctccat tttcattgag 660
gtgcagactg accagggact ctacgggaca cggagcacgg ccgttctatc agtgaactac 720
gacggtgaag ctagcttgta cgagaaatac ctggagagtg gtatatggaa ggatcacacg 780
gtgcattacc agatagagta g 801
<210>39
<211>266
<212>PRT
<213>水稻(Oryza sativa)
<400>39
Wet Cys Ile Ala Ala Trp Val Trp Gln Ala His Pro Gln His Gln Leu
1 5 10 15
Leu Leu Leu Leu Asn Arg Asp Glu Phe His Ser Arg Pro Thr Lys Ala
20 25 30
Val Gly Trp Trp Gly Glu Gly Ser Lys Lys Ile Leu Gly Gly Arg Asp
35 40 45
Val Leu Gly Gly Gly Thr Trp Met Gly Cys Thr Lys Asp Gly Arg Leu
50 55 60
Ala Phe Leu Thr Asn Val Leu Glu Pro Asp Ala Met Pro Gly Ala Arg
65 70 75 80
Thr Arg Gly Asp Leu Pro Leu Arg Phe Leu Gln Ser Asn Lys Ser Pro
85 90 95
Leu Glu Val Ala Thr Glu Val Ala Lys Glu Als Asp Glu Tyr Asn Gly
100 105 110
Phe Aen Leu Val Leu Ale Asp Leu Thr Thr Asn Val Met Val Tyr Val
115 120 125
Ser Asn Arg Pro Lys Gly Gln Pro Ala Thr Ile Gln Leu Val Ser Pro
130 135 140
Gly Leu His Val Leu Ser Asn Ala Arg Leu Asp Ser Pro Trp Gln Lys
145 150 155 160
Ala Ile Arg Leu Gly Lye Asn Phe Arp Glu His Leu Arg Lys His Gly
165 170 175
Asp Asp Glu Val Glu Ala Lys Asp Ile Val Glu Arg Leu Met Thr Asp
180 185 190
Thr Thr Lys Ala Asp Lys Asp Arg Leu Pro Asl Thr Gly Cys Asp Pro
195 200 205
Asn Trp Glu His Gly Leu Ser Ser Ile Phe Ile Glu Val Gln Thr Asp
210 215 220
Gln Gly Leu Tyr Gly Thr Arg Ser Thr Ala Val Leu Ser Val Asn Tyr
225 230 235 240
Asp Gly Glu Ala Ser Leu Tyr Glu Lys Tyr Leu Glu Ser Gly Ile Trp
245 250 255
Lys Asp His Thr Val His Tyr Gln Ile Glu
260 265
<210>40
<211>707
<212>DNA
<213>蒺藜苜蓿(Medicago truncatula)
<220>
<221>misc_feature
<222>(689)..(689)
<223>n是a、c、g或t
<220>
<221>mis_feature
<222>(692)..(692)
<223>n是a、c、g或t
<400>40
ttgtacttag ttatagtatt attgaagcta gccaactcaa aatttgtgaa caatgtgtat 60
agctttgttt ctttggcaat ctcatccacc tttatccttt tcttcttttg aataatagag 120
atgaatatca caataggcct acaaagaaag tgtcatggtg ggaagaatgt gatatagtgg 180
gaggaaggga tgaaatagga ggagggacat ggttggcttg ttcttcacaa ggaaaagtgg 240
cttttcttac caatgttttg gagcttcata cttgccctga ggccaaaact cgtggagacc 300
tacccctcat gtttctcaag agcagcaaga atcccaaaga atttgcagaa agcttaaaaa 360
gagaagctca atattacaat ggattcaatt tagtcattgc tgatattaat tccaaatcca 420
tggtatacat atcaaataga cccaagggac agccaattac tgtccaagag gttcctcctg 480
gtctacatgt actttcaaat gctaagttaa attcaccatg gcataaggct cagcgccttc 540
aatttagatt caaagagcat cttgctaaaa atggggaagg tgagatacat gtaaaggaag 600
taattaaaaa gctaatgaag gacaaaatta aagcagacaa aagcatgcta cctaatatat 660
gctcacttga ttggggaatt caatcttanc tncatttttg ttgaaga 707
<210>41
<211>213
<212>PRT
<213>蒺藜苜蓿(Medicago truncatula)
<220>
<221>misc_feature
<222>(209)..(210)
<223>Xaa可为任何天然氨基酸
<400>41
Leu Cys Phe Phe Gly Asn Leu Ile His Leu Tyr Pro Phe Leu Leu Leu
1 5 10 15
Asn Asn Arg Asp Glu Tyr His Asn Arg Pro Tnr Lys Lys Val Ser Trp
20 25 30
Trp Glu Glu Cys Asp Ile Val Gly Gly Arg Asp Glu Ile Gly Gly Gly
35 40 45
Thr Trp Leu Ala Cys Ser Ser Gln Gly Lys Val Ala Phe Leu Thr Asn
50 55 60
Val Leu Glu Leu His Thr Cys Pro Glu Ala Lys Thr Arg Gly Asp Leu
65 70 75 80
Pro Leu Met Phe Leu Lys Ser Ser Lys Asn Pro Lys Glu Phe Ala Glu
85 90 95
Set Leu Lys Arg Glu Ala Gln Tyr Tyr Asn Gly Phe Asn Leu Val Ile
100 105 110
Ala Asp Ile Asn Ser Lys Ser Met Val Tyr Ile Ser Asn Arg Pro Lys
115 120 125
Gly Gln Pro Ile Thr Val Gln Glu Val Pro Pro Gly Leu Hls Val Leu
130 135 140
Ser Asn Ale Lys Leu Asn Ser Pro Trp His Lys Ala Gln Arg Leu Gln
145 150 155 160
Phe Arg Phe Lys Glu His Leu Ala Lys Asn Gly Glu Gly Glu Ile His
165 170 175
Val Lys Glu Val Ile Lys Lys Leu Met Lys Asp Lys Ile Lys Ala Asp
180 185 190
Lys Ssr Met Leu Pro Asn Ile Cys Ser Leu Asp Trp Gly Ile Gln Ser
195 200 205
Xaa Xaa His Phe Cys
210
<210>42
<211>627
<212>DNA
<213>铃薯(Solanum tuberosum)
<400>42
gatatggaaa gcgcatccgc tgtatccctt cctcctattc ctcaacagag atgaatacca 60
caatcgtgat atacttggtg gaagggatga agttgctggt gggacttggt tggcttgtac 120
tcgcaccgga agacttgctt tccttactaa tgttcgagaa atcaattcaa attcacatac 180
caaaagtagg ggagaccttc ctcttcgatt cttgaagagt gtaaagagcc ctcatgattt 240
ttcagagcaa cttttgaaag aagcaggcga atataatggg tttaacttga tagtagctga 300
tctttgttca atgactatgc ttgatataac caaccgacca aaacacaccg gtatgtccgg 360
cactgaagtt tcacccggta ttcacgtttt atcaaatgca acactagact ctccatggcc 420
taagtctcaa cggctggagt acagtttcaa gcaattattg gatgaatatg gcgaatctga 480
aattccaata gggcagacag ctgaaagaat aatgagagac ttggctaaag aagatagcaa 540
cctgccaggc atctattccc ctgagtgtga gtaccagttg agctccatat tcgttgacac 600
tgaaatgtcc atggggcgtt ttggcac 627
<210>43
<211>208
<212>PRT
<213>马铃薯(Solanum tuberosum)
<400>43
Ile Trp Lys Ala His Pro Leu Lyr Pro Phe Leu Leu Phe Leu Asn Arg
1 5 10 15
Asp Glu Tyr His Asn Arg Asp Ile Leu Gly Gly Arg Asp Glu Val Ala
20 25 30
Gly Gly Thr Trp Leu Ala Cys Thr Arg Thr Gly Arg Leu Ala Phe Leu
35 40 45
Thr Asn Val Arg Glu Ile Asn Ser Asn Ser His Thr Lys Ser Arg Gly
50 55 60
Asp Leu Pro Leu Arg Phe Leu Lys Ser Val Lys Ser Pro His Asp Phe
65 70 75 80
Ser Glu Gln Leu Leu Lys Glu Ala Gly Glu Tyr Asn Gly Phe Asn Leu
85 90 95
Ile Val Ala Asp Leu Cys Ser Met Thr Met Leu Asp Ile Thr Asn Arg
100 105 110
Pro Lys His Thr Gly Met Ser Gly Thr Glu Val Ser Pro Gly Ile His
115 120 125
Val Leu Ser Asn Ala Thr Leu Asp Sar Pro Trp Pro Lys Ser Gln Arg
130 135 140
Leu Glu Tyr Ser Phe Lys Gln Leu Leu Asp Glu Tyr Gly Glu Ser Glu
145 150 155 160
Ile Pro Ile Gly Gln Thr Ala Glu Arg Ile Met Arg Asp Leu Ala Lys
165 170 175
Glu Asp Ser Asn Leu Pro Gly Ile Tyr Ser Pro Glu Cys Glu Tyr Gln
180 185 190
Leu Ser Ser Ile Phe Val Asp Thr Glu Met Ser Met Gly Arg Phe Gly
195 200 205
<210>44
<211>423
<212>DNA
<213>大豆(Glycine max)
<400>44
gtgctcccca gattgttcta ttttggcttt tataaagaat tgtttagatc ctttgaattg 60
aagaatgtgt atagctttgt ttctttggca agcccaccca ctctaccctt tccttctttt 120
gaacaacaga gatgaatatc acaacaggcc tacgaagcca gtgtcatggt gggaagatat 180
tgatatagtt ggaggaagag atgagattgc tggaggaaca tggttggctt gttcaagaga 240
aggaagagtt gcttttctga ccaatgtttt ggagcttcgt tcccttcctg aggctaaaag 300
cagaggagac ctacctgtct catttcttaa gagtggaaag catccgaaag aatttgcaga 360
aagtctaaaa atggaagctc attattacaa tgggttcaac ttgattgtgg ccgatattcc 420
gtc 423
<210>45
<211>119
<212>PRT
<213>大豆(Glycine max)
<400>45
Met Cys Ile Ala Leu Phe Leu Trp Gln Ala His Pro Leu Tyr Pro Phe
1 5 10 15
Leu Leu Leu Asn Asn Arg Asp Glu Tyr His Asn Arg Pro Thr Lys Pro
20 25 30
Val Ser Trp Trp Glu Asp Ile Asp Ile Val Gly Gly Arg Asp Glu Ile
35 40 45
Ala Gly Gly Thr Trp Leu Ala Cys Ser Arg Glu Gly Arg Val Ala Phe
50 55 60
Leu Thr Asn Val Leu Glu Leu Arg Ser Leu Pro Glu Ala Lys Ser Arg
65 70 75 80
Gly Asp Leu Pro Val Ser Phe Leu Lys Ser Gly Lys His Pro Lys Glu
85 90 95
Phe Ala Glu Ser Leu Lys Met Glu Ala His Tyr Tyr Asn Gly Phe Asn
100 105 110
Leu Ile Val Ala Asp Ile Pro
115
<210>46
<211>542
<212>DNA
<213>甜菜(Beta vulgaris)
<400>46
cccacgcgtc cgcccacgcg tccgcccacg cgtccgcgga cgcgtgggtc gacccacgcg 60
tccgtttgaa ccacttttca attttcgagc tgaaacatga aagtgcatta attcacaccc 120
aaacctgcaa cacatctttc tgaatagctc aaaattcgaa attccactca tgcaagagca 180
agaatttagc atgaacatga aatgacaaat ttgaaatttc caccactaat catgaaaaac 240
ccatgaaaaa gaaacgtgat gtgcatcgca atatttcaat ggcaatccca cccactttac 300
ccatttcttc tactcctcaa ccgcgacgaa tatcataccc ggccaacaaa tccagcaggg 360
tggtgggaag gtgaagaaat tgttggtggg aaagatgaag ttggtggtgg gacatggttg 420
gcttgttcca aaggtggaag aattgctttt cttaccaatt ttagagagag agaatcaatt 480
cctcatgcta aaagtagagg agatttgcct gttcgttttc ttaagtgtaa gaaagatccg 540
gc 542
<210>47
<211>94
<212>PRT
<213>甜菜(Beta vulgaris)
<400>47
Mgt Cys Ile Ala Ile Phe Gln Trp Gln Ser His Pro Leu Tyr Pro Phe
1 5 10 15
Leu Leu Leu Leu Asn Arg Asp Glu Tyr His Thr Arg Pro Thr Asn Pro
20 25 30
Ala Gly Trp Trp Glu Gly Glu Glu Ile Val Gly Gly Lys Asp Glu Val
35 40 45
Gly Gly Gly Thr Trp Leu Ala Cys Ser Lys Gly Gly Arg Ile Ala Phe
50 55 60
Leu Thr Asn Phe Arg Glu Arg Glu Ser Ile Pro His Ala Lys Ser Arg
65 70 75 80
Gly Asp Leu Pro Val Arg Phe Leu Lys Cys Lys Lys Asp Pro
85 90
<210>48
<211>190
<212>DNA
<213>辣椒(Capsicum annuum)
<400>48
agaatgtgca tagcagtgtt tatttggcaa gcagacagta gatattcatt agtgttgttg 60
ttgaacagag atgaatatca caataggcca acaaaggcag ttcattggtg ggaaggtgga 120
gatcaaatag ttggtggtaa agatgacgtt ggtggtggta cttggttacc ttcttcaaca 180
aatggtaaat 190
<210>49
<211>62
<212>PRT
<213>辣椒(Capsicum annuum)
<400>49
Met Cya Ila Ala Val Pha Ila Trp Gln Ala Asp Ser Arg Tyr Ser Leu
1 5 10 15
Val Leu Leu Leu Asn Arg Asp Glu Tyr His Asn Arg Pro Thr Lys Ala
20 25 30
Val His Trp Trp Glu Gly Gly Asp Gln Ile Val Gly Gly Lys Asp Asp
35 40 45
Val Gly Gly Gly Thr Trp Leu Pro Ser Ser Thr Asn Gly Lys
50 55 60
<210>50
<211>719
<212>DNA
<213>莴苣(Lactuca sativa)
<220>
<221>miac_feature
<222>(687)..(687)
<223>n是a、c、g或t
<400>50
gggaggaaga gtgtcattcc ttactaacgt cttggagctt cacactctcc cggaagccaa 60
aactagagga gaccttccac ttcgtttctt ggagagcaat aagagtccag aggaatttgc 120
aaaggaattg gtgaaggagg ttcatgagta caatgggttc aacctcataa cccttgacat 180
ttcttcaaaa acgatgtttt atatatcaaa tagaccaaaa agtgaacctc caactgttca 240
acaggttcaa ccaggcatcc atgtcctctc caatgccaag ctcgactccc cttggccaaa 300
ggctcaacgt ttgaagttta attttaaaaa gttgcttagc gcatatgata aagacgaaga 360
tatacccatg aaggatatga tggacaaact aatgagagac accatgaaag cagaaaagag 420
tcaacttcct aatatttgtt ccattgattg ggagcataat ctaagctcga tatttgttga 480
agtagacacc ccgttgggtc gttatgggac gagaagcatg attgcactaa gtatcaaaga 540
taccgaagaa gcaagttttc atgagaccta cattgaaaga ggattttggt gggagaaaac 600
cgtcgattat tatgttactc cacaagttaa aataaaagat atcgtcttct aagactaaat 660
atacgttaca aatatttaaa atacagnctt tctctctata tatatcttat atataaaaa 719
<210>51
<211>216
<212>PRT
<213>莴苣(Lactuca sativa)
<400>51
Gly Gly Arg Val Ser Phe Leu Thr Asn Val Leu Glu Leu His Thr Leu
1 5 10 15
Pro Glu Ala Lys Thr Arg Gly Asp Leu Pro Leu Arg Phe Leu Glu Ser
20 25 30
Asn Lys Ser Pro Glu Glu Phe Ala Lys Glu Leu Val Lys Glu Val His
35 40 45
Glu Tyr Asn Gly Phe Asn Leu Ile Thr Leu Asp Ile Ser Ser Lys Thr
50 55 60
Met Phe Tyr Ile Ser Asn Arg Pro Lys Ser Glu Pro Pro Thr Val Gln
65 70 75 80
Gln Val Gln Pro Gly Ile His Val Leu Ser Asn Ala Lys Leu Asp Ser
85 90 95
Pro Trp Pro Lys Ala Gln Arg Leu Lys Phe Asn Phe Lys Lys Leu Leu
100 105 110
Ser Ala Tys Asp Lys Asp Glu Asp Ile Pro Met Lys Asp Met Met Asp
115 120 125
Lys Leu Met Arg Asp Thr Met Lye Ala Glu Lys Ser Gln Leu Pro Asn
130 135 140
Ile Cye Ser Ile Asp Trp Glu His Asn Leu Ser Ser Ile Phe Val Glu
145 150 155 160
Val Asp Thr Pro Leu Gly Arg Tyr Gly Thr Arg Ser Met Ile Ala Leu
165 170 175
Ser Ile Lys Asp Thr Glu Glu Ala Ser Phe His Glu Thr Tyr Ile Glu
180 185 190
Arg Gly Phe Trp Trp Glu Lys Thr Val Asp Tyr Tyr Val Thr Pro Gln
195 200 205
Val Lys Ile Lys Asp Ile Val Phe
210 215
<210>52
<211>522
<212>DNA
<213>高梁(Sorghum bicolor)
<400>52
agctaagata gttgcaaaca agcgagttac ttacaaccaa ccaaaggagt agaaaccacc 60
tgaagccatg tgcattgctg catggatttg gcaggctcac cctgtgcacc aactcctcct 120
gcttctcaac agagatgagt tccacagcag gcctacaaaa gcagtaggat ggtggggaga 180
aggctcaaag aagattcttg gtggcaggga tgtgcttggt ggaggaacat ggatggggtg 240
caccaaggat ggaaggcttg ccttcctgac caatgtgctt gaaccagatg ccatgcccgg 300
tgcacggact aggggagatc tgcctctcag gttcctgcag agcaacaaga gcccactcga 360
agttgcaact gaagtggcag aagaagctca taaatacaat ggcttcaacc tcatactagc 420
tgatctaaca acaaatatca tggtctatgt gtcaaaccgg cctaaggggc agcctgcaac 480
aattcaactc gtctcaccag gactccatgt gctgtccaat gc 522
<210>53
<211>151
<212>PRT
<213>高粱(Sorghum bicolor)
<400>53
Met Cys Ile Ala Ala Trp Ile Trp Gln Ala His Pro Val His Gln Leu
1 5 10 15
Leu Leu Leu Leu Asn Arg Asp Glu Phe His Ser Arg Pro Thr Lys Ala
20 25 30
Val Gly Trp Trp Gly Glu Gly Ser Lys Lys Ile Leu Gly Gly Arg Asp
35 40 45
Val Leu Gly Gly Gly Thy Trp Met Gly Cys Thr Lys Asp Gly Arg Leu
50 55 60
Ala Phe Leu Thr Asn Val Leu Glu Pro Asp Ala Met Pro Gly Ala Arg
65 70 75 80
Thr Arg Gly Asp Leu Pro Leu Arg Phe Leu Gln Ser Asn Lys Ser Pro
85 90 95
Leu Glu Val Ala Thr Glu Val Ala Glu Glu Ala His Lys Tyr Asn Gly
100 105 110
Phe Asn Leu Ile Leu Ala Aep Leu Thr Thr Asn Ila Met Val Tyr Val
115 120 125
Ser Aan Arg Pro Lys Gly Gln Pro Ala Thr Ile Gln Leu Val ser Pro
130 135 140
Gly Leu His Val Leu Ser Asn
145 150
<210>54
<211>1132
<212>DNA
<213>小麦(Triticum aestivum)
<220>
<221>misc_feature
<222>(17 )..(17)
<223>n是a.c.g或t
<400>54
aatcctcagg gttacgncga cccacgcgtc cgcaaacaca caaggcgcac ggttgcgaaa 60
caagggaatt atttagaagc aggaaggaac acctgaagcc atgtgtatcg ctgcatggat 120
ttggcaggct cacccacagc atcagctcct gcttctgctc aacagagatg agttccatag 180
caggcctaca aaggcagtag gatggtgggg ggagggctca atgaagattc ttggcggcag 240
ggatgtactt ggtggaggaa catggatggg gagcaccaaa gatggcagac ttgccttcct 300
gaccaatgtg ctcgagcctg atgcgatgcc tggcgcacgc actaggggag acctgcccct 360
caggttcctg cagggcaaca agagcccact ggaggttgca actgaagtcg caaaagaagc 420
tgatgagtac aatggcttca accttatact agctgatcta accaggaatg tcatggttta 480
tgtgtcaaac cggccaaagg ggcagcctgc gacgattcag ctcgtctcac caggactcca 540
tgtgttgtcc aatgcaaggc tagacagccc ttggcagaag gcaattcgcc ttggtaaaaa 600
cttcagggag tttataagga agcatggtga tgatgaagtt gaagcgaagg atatagctga 660
tagactaatg actgacacca cgagggctga taaagatagg ctgccaaaca ccggttgtga 720
tcccaactgg gagcacggtc tgagctccat cttcatcgag gtgcaaactg acgaagggct 780
ctatgggaca aggagcacag cagttctttc agtgaactat gatggagaag ctagcttata 840
tgagaagtac ctcgagagtg gtatatggaa gaaccacaca gtgcattacc agatagaatt 900
gccaatgcgc acctaaaggc aggagcctca aataggaaga aagaatgaat agctaccatt 960
gtgcatgctg ttatttccac agttgcgctt taagatcaca taatgatctc taattatggc 1020
aattaaatta tttactgtat gcggatctat aaattcagag acagatcaag tcaaattgtt 1080
gaatatatat acataataat aatatgatat agtatgtgta tttacagact tc 1132
<210>55
<211>271
<212>PRT
<213>小麦(Triticum aestivum)
<400>55
Met Cys Ile Ala Ala Trp Ile Trp Gln Ala His Pro Gln His Gln Leu
1 5 10 15
Leu Leu Leu Leu Asn Arg Asp Glu Phe His Ser Arg Pro Thr Lys Ala
20 25 30
Val Gly Trp Trp Gly Glu Gly Ser Met Lye Ile Leu Gly Gly Arg Asp
35 40 45
Val Leu Gly Gly Gly Thr Trp Met Gly Ssr Thr Lys Asp Gly Arg Leu
50 55 60
Ala Phe Leu Thr Asn Val Leu Glu Pro Asp Ala Met Pro Gly Ala Arg
65 70 75 80
Thr Arg Gly Asp Leu Pro Leu Arg Phe Leu Gln Gly Asn Lys Ser Pro
85 90 95
Leu Glu Val Ala Thr Glu Val Ala Lys Glu Ala Asp Glu Tyr Asn Gly
100 105 110
Phe Asn Leu Ile Leu Ala Asp Leu Thr Arg Asn Val Met Val Tyr Val
115 120 125
Ser Asn Arg Pro Lys Gly Gln Pro Ala Thr Ila Gln Leu Val Ser Pro
130 135 140
Gly Leu His Val Leu Ser Asn Ala Arg Leu Asp Ser Pro Trp Gln Lys
145 150 155 160
Ala Ile Arg Leu Gly Lys Asn Phe Arg Glu Phe Ile Arg Lys His Gly
165 170 175
Asp Asp Glu Val Glu Ala Lys Asp Ile Ala Asp Arg Leu Met Thr Asp
180 185 190
Thr Thr Arg Ala Asp Lys Asp Arg Leu Pro Asn Thr Gly Cys Asp Pro
195 200 205
Asn Trp Glu His Gly Leu Ser Ser Ile Phe Ile Glu Val Glu Thr Asp
210 215 220
Glu Gly Leu Tyr Gly Thr Arg Ser Thr Ala Val Leu Ser Val Asn Tyr
225 230 235 240
Asp Gly Glu Ala Ser Leu Tyr Glu Lys Tyr Leu Glu Ser Gly Ile Trp
245 250 255
Lys Asn His Thr Val His Tyr Gln Ile Glu Leu Pro Met Arg Thr
260 265 270
<210>56
<211>1090
<212>DNA
<213>甘蔗(Saccharum officinarum)
agcaaaacaa gcgagttact tacaaccaac caaggagtag aaaccacctg aagccatgtg 60
cattgctgca tggatttggc aggctcaccc tgtgcaccaa ctcctcctga ttctcaacag 120
agatgagttc cactgcaggc ctacaaaagc agtaggatgg tggggagaag gctcaaagaa 180
gattcttggc ggcagggatg tgcttggtgg aggaacatgg atgggttgca ccaaggatgg 240
caggcttgcc ttcctgacca atgtgcttga accagatgcc atgcccggtg cacggactag 300
gggagatctg cctctcaggt tcctgcagag caacaagagc ccactcgug ttgcaactga 360
agtggcagaa gaagctcatg aatacaatgg gttcaacctc atactagctg atctaacaac 420
aaatatcatg gtctatgtgt caaatcggcc taaggggcag cctgcaacaa ttcaactcgt 460
ctcaccagga ctccatgtgc tgtccaatgc aaggctagat agcccttggc agaaggcaat 540
tcgccttggt aaaaacttca aggagcttct tagggagcat ggtgacgatg agattgaagt 600
gaaggatata gttgagaggc taatgactga caccacaaag gctgacaaag atagactgcc 660
aaacactggt tgtgatccca actgggagca tggtctgagc tccatcttca tcgaggtgca 720
aactgaccaa gggctctacg ggacacggag cacagccgtt ttatcagtga actatgatgg 780
tgaagctagc ttgtacgaga agtaccttga gagtggtata tggaaggacc acacagtgaa 840
ttaccagata gagtagtagg cattgcacag gaaaagctgg caacctcaaa taaatagaga 900
tatgaagcag acacaattgt ggatttcatt ctttccctaa tccctagtca ccttcacgac 960
tatctaagat cccatcatga tgccaattac attatttact gtaagcagat ttgtcacttg 1020
acgataaaat gtcaagcaga agtttaagtt taaatatata caccaaatat ataaatttac 1080
agacttcgtg 1090
<210>57
<211>266
<212>PRT
<213>甘蔗(Saccharum officinarum)
<400>57
Met Cys Ile Ala Ala Trp Ile Trp Gln Ala His Pro Val His Gln Leu
1 5 10 15
Leu Leu Ile Leu Asn Arg Asp Glu Phe His Cys Arg Pro Thr Lya Ala
20 25 30
Val Gly Trp Trp Gly Glu Gly Ser Lye Lys Ile Leu Gly Gly Arg Asp
35 40 45
Val Leu Gly Gly Gly Thr Trp Met Gly Cys Thr Lys Asp Gly Arg Leu
50 55 60
Ala Phe Leu Thr Asn Val Leu Glu Pro Asp Ala Met Pro Gly Ala Arg
65 70 75 80
Thr Arg Gly Asp Leu Pro Leu Arg Phe Leu Gln Ser Asn Lys Ser Pro
85 90 95
Leu Glu Val Ala Thr Glu Val Ala Glu Glu Ala His Glu Tyr Asn Gly
100 105 110
Phe Asn Leu Ile Leu Ala Asp Leu Thr Thr Asn IIe Met Val Tyr Val
115 120 125
Ser Asn Arg Pro Lys Gly Gln Pro Ala Thr Ile Gln Leu Val Ser Pro
130 135 140
Gly Leu His Val Leu Ser Asn Ala Arg Leu Asp Ser Pro Trp Gln Lys
145 150 155 160
Ala Ile Arg Leu Gly Lys Asn Phe Lys Glu Leu Leu Arg Glu His Gly
165 170 175
Asp Asp Glu Ile Glu Val Lys Asp Ile Val Glu Arg Leu Met Thr Aap
180 185 190
Thr Thr Lys Ala Asp Lys Asp Arg Leu Pro Asn Thr Gly Cys Asp Pro
195 200 205
Asn Trp Glu His Gly Leu Ser Ser Ile Phe Ile Glu Val Gln Thr Asp
210 215 220
Gln Gly Leu Tyr Gly Thr Arg Ser Thr Ala Val leu Ser Val Asn Tyr
225 230 235 240
Asp Gly Glu Ala Ser Leu Tyr Glu Lys Tyr Leu Glu Ser Gly Ile Trp
245 250 255
Lya Asp His Thr Val Asn Tyr Gln Ile Glu
260 265
<210>58
<211>743
<212>DNA
<213>人工序列
<220>
<223>cwp1基因启动子的功能片段
<400>58
tgccgtccta ttcttagaat actcaagtaa tttaacgtag tggtgaaaat ttgataaatt 60
aattatatac taatttttca gtcttatttt atgtggtata tttaattgga tatgtagttt 120
aagaaataat aaaaacttta aaatatttat aaatttactt ttctaaaaaa gtgaattcaa 180
ttttttctct cctcataaat gtattagagt attatcatta aaattaagtg ggactaataa 240
aggtaaaaaa taaattattc ctttaaatta tttaaccata taagaaaatg tgacattctt 300
ttttagactt gactaaaata gaaaataatg tcatatatat aaaatgagac gaaaaaagta 360
aatattaatt taaaatttaa aactttaggg taatagctac tttgaattac ctagatttca 420
ataaaattca acatataata aaacatacta atttacaatt tttaaaataa tatgactaaa 480
agtcatatta ttcaaaaaac aatctatacc gccgtcacct agttacttta atttgtgtag 540
cttctagtac atacattttt aaactttatc tgaatttaat attttaatta tattaaacat 600
ttattaaaat ttataaaatt taaattgacg taatataatg aagagagtag tactatataa 660
accatgtgag tactaacatg atcttcatct tattcttgtt tttatttata gaaacaataa 720
aatagttata aaattaatca atc 743
Claims (19)
1.一种分离的多核苷酸,所述多核苷酸包含编码多肽的核酸序列,所述多肽具有与SEQ ID NO:22至少88%同源的氨基酸序列,所述多肽能够增加表达该多肽的植物的角质层水透过性。
2.权利要求1的分离的多核苷酸,其中所述核酸序列如SEQ IDNO:21或23所述。
3.权利要求1的分离的多核苷酸,其中所述氨基酸序列如SEQ IDNO:22所述。
4.一种核酸构建物,所述核酸构建物包含权利要求1的分离的多核苷酸。
5.权利要求4的核酸构建物,所述核酸构建物还包含有效连接至所述核酸序列的启动子。
6.一种宿主细胞,所述宿主细胞包含权利要求4的核酸构建物。
7.一种遗传修饰的植物,所述植物包含权利要求1的分离的多核苷酸。
8.一种寡核苷酸,所述寡核苷酸能够与权利要求1的分离的多核苷酸特异性杂交。
9.一种分离的多肽,所述多肽包含与SEQ ID NO:22至少88%同源的氨基酸序列,所述多肽能够增加表达该多肽的植物的角质层水透过性。
10.一种抗体,所述抗体能够特异性识别权利要求9的多肽。
11.一种栽培的番茄植株,所述番茄植株具有含基因渗入的基因组,所述基因渗入来源于野生番茄(Lycopersicon spp.),所述基因渗入包含所述野生番茄的染色体4的一部分,小于由端粒标记TG464延伸至着丝点标记CT173的染色体片断,所述基因渗入能够增加栽培番茄植物的角质层水透过性。
12.一种生产栽培植物的脱水果实的方法,所述方法包括遗传修饰植物,以表达具有与SEQ ID NO:22至少30%同源的氨基酸序列的多肽,所述多肽能够增加表达该多肽的植物的角质层水透过性。
13.权利要求12的方法,所述方法还包括:
使果实在植株上脱水;并随后
收集脱水果实。
14.权利要求12的方法,所述方法还包括:
在作物脱水前将果实由作物上取下;并随后
使果实脱水。
15.一种遗传修饰的种子,所述种子包含分离的多核苷酸,所述多核苷酸包含编码多肽的核酸序列,所述多肽具有与SEQ ID NO:22至少30%同源的氨基酸序列,所述多肽能够增加表达该多肽的植物的角质层水透过性。
16.一种遗传修饰的果实,所述果实包含分离的多核苷酸,所述多核苷酸包含编码多肽的核酸序列,所述多肽具有与SEQ ID NO:22至少30%同源的氨基酸序列,所述多肽能够增加表达该多肽的植物的角质层水透过性。
17.权利要求12、15或16的方法、遗传修饰的种子或遗传修饰的果实,其中所述核酸序列如SEQ ID NO:21、23、24、26、28、30、32、34、36、38、40、42、44、46、48、50、52、54或56所述。
18.权利要求12、15或16的方法、遗传修饰的种子或遗传修饰的果实,其中所述氨基酸序列如SEQ ID NO:22、25、27、29、31、33、35、37、39、41、43、45、47、49、51、53、55或57所述。
19.一种遗传修饰的植物,所述植物表达多肽,所述多肽具有与SEQ ID NO:22至少30%同源的氨基酸序列,所述多肽能够增加表达该多肽的植物的角质层水透过性。
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IL164125 | 2004-09-19 | ||
IL164125A IL164125A (en) | 2004-09-19 | 2004-09-19 | The visitor's garden has a scattering development in tomato fruit cuticle |
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CN101123870A true CN101123870A (zh) | 2008-02-13 |
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CNA2005800393423A Pending CN101123870A (zh) | 2004-09-19 | 2005-09-19 | 用于产生具有增加的角质层水透过性的植物的分离多肽以及编码该多肽的多核苷酸 |
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US (4) | US8481809B2 (zh) |
JP (1) | JP4808718B2 (zh) |
CN (1) | CN101123870A (zh) |
BR (1) | BRPI0515680A (zh) |
CA (1) | CA2580713C (zh) |
IL (1) | IL164125A (zh) |
WO (1) | WO2006030445A2 (zh) |
Cited By (3)
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CN110373425A (zh) * | 2019-07-25 | 2019-10-25 | 中国农业大学 | NOR-like1基因及其编码的蛋白质在调控番茄果实采后失水中的应用 |
CN111819285A (zh) * | 2018-01-09 | 2020-10-23 | 希博斯美国有限公司 | 防碎基因和突变 |
CN114644702A (zh) * | 2020-12-21 | 2022-06-21 | 中国农业大学 | Tango蛋白、其相关生物材料以及植物育种方法 |
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IL131509A (en) | 1999-08-19 | 2007-03-08 | Israel State | A method of growing tomatoes with a low water content |
IL164125A (en) | 2004-09-19 | 2014-06-30 | Arthur Schaffer | The visitor's garden has a scattering development in tomato fruit cuticle |
US20100162433A1 (en) | 2006-10-27 | 2010-06-24 | Mclaren James | Plants with improved nitrogen utilization and stress tolerance |
WO2008073578A2 (en) | 2006-12-08 | 2008-06-19 | Iowa State University Research Foundation, Inc. | Plant genes involved in nitrate uptake and metabolism |
IL181193A0 (en) * | 2007-02-06 | 2007-07-04 | Volcani Ct The State Of Israel | Means and methods of producing fruits with high levels of anthocyanins and flavonols |
MX2009010160A (es) | 2007-03-30 | 2009-10-12 | Unilever Nv | Proceso de produccion de pasta de tomate. |
US8692070B2 (en) | 2009-11-04 | 2014-04-08 | Iowa Corn Promotion Board | Plants with improved nitrogen utilization and stress tolerance |
JP2016527885A (ja) * | 2013-07-25 | 2016-09-15 | ザ ステイト オブ イスラエル ミニストリー オブ アグリカルチャー アンド ルーラル ディベロップメント アグリカルチュラル リサーチ オーガニゼイション (エー.アール.オー.) ザ ボルカニ センター | 加工トマト、および該トマトを製造し、そして用いる方法 |
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IL131509A (en) | 1999-08-19 | 2007-03-08 | Israel State | A method of growing tomatoes with a low water content |
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IL164125A (en) | 2004-09-19 | 2014-06-30 | Arthur Schaffer | The visitor's garden has a scattering development in tomato fruit cuticle |
MX2009010160A (es) | 2007-03-30 | 2009-10-12 | Unilever Nv | Proceso de produccion de pasta de tomate. |
US20090064367A1 (en) | 2007-08-28 | 2009-03-05 | Douglas Heath | Tomato line chd 15-2062 |
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JP2016527885A (ja) | 2013-07-25 | 2016-09-15 | ザ ステイト オブ イスラエル ミニストリー オブ アグリカルチャー アンド ルーラル ディベロップメント アグリカルチュラル リサーチ オーガニゼイション (エー.アール.オー.) ザ ボルカニ センター | 加工トマト、および該トマトを製造し、そして用いる方法 |
-
2004
- 2004-09-19 IL IL164125A patent/IL164125A/en active IP Right Grant
-
2005
- 2005-09-19 BR BRPI0515680-7A patent/BRPI0515680A/pt not_active IP Right Cessation
- 2005-09-19 WO PCT/IL2005/001000 patent/WO2006030445A2/en active Application Filing
- 2005-09-19 CN CNA2005800393423A patent/CN101123870A/zh active Pending
- 2005-09-19 US US11/663,151 patent/US8481809B2/en not_active Expired - Fee Related
- 2005-09-19 CA CA2580713A patent/CA2580713C/en not_active Expired - Fee Related
- 2005-09-19 JP JP2007531955A patent/JP4808718B2/ja not_active Expired - Fee Related
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- 2013-05-27 US US13/902,885 patent/US9497909B2/en active Active
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- 2016-11-20 US US15/356,597 patent/US20170066803A1/en not_active Abandoned
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Cited By (4)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
CN111819285A (zh) * | 2018-01-09 | 2020-10-23 | 希博斯美国有限公司 | 防碎基因和突变 |
CN110373425A (zh) * | 2019-07-25 | 2019-10-25 | 中国农业大学 | NOR-like1基因及其编码的蛋白质在调控番茄果实采后失水中的应用 |
CN114644702A (zh) * | 2020-12-21 | 2022-06-21 | 中国农业大学 | Tango蛋白、其相关生物材料以及植物育种方法 |
CN114644702B (zh) * | 2020-12-21 | 2023-10-03 | 中国农业大学 | Tango蛋白、其相关生物材料以及植物育种方法 |
Also Published As
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WO2006030445A3 (en) | 2007-01-11 |
US20180244733A1 (en) | 2018-08-30 |
JP2008513004A (ja) | 2008-05-01 |
JP4808718B2 (ja) | 2011-11-02 |
US20170066803A1 (en) | 2017-03-09 |
BRPI0515680A (pt) | 2008-07-29 |
US20090311398A1 (en) | 2009-12-17 |
CA2580713C (en) | 2014-04-22 |
US9497909B2 (en) | 2016-11-22 |
US8481809B2 (en) | 2013-07-09 |
IL164125A0 (en) | 2005-12-18 |
CA2580713A1 (en) | 2006-03-23 |
WO2006030445A2 (en) | 2006-03-23 |
US10611808B2 (en) | 2020-04-07 |
IL164125A (en) | 2014-06-30 |
US20130316057A1 (en) | 2013-11-28 |
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