CN113754746B - 水稻雄性育性调控基因、其应用以及利用CRISPR-Cas9调控水稻育性的方法 - Google Patents
水稻雄性育性调控基因、其应用以及利用CRISPR-Cas9调控水稻育性的方法 Download PDFInfo
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Abstract
本发明涉及生物技术领域,具体涉及水稻雄性育性调控基因、其应用以及利用CRISPR‑Cas9调控水稻育性的方法。本发明提供具有调控水稻雄性生殖细胞发育和花粉育性功能的水稻基因GMS2,其核苷酸序列如SEQ ID NO:1所示,CDS序列如SEQ ID NO:2所示,氨基酸序列如SEQ ID NO:3所示。本发明还提供GMS2基因的CRISPR‑Cas9敲除突变体。本发明提供的水稻基因GMS2及其突变体可用于水稻杂交种的不育化制种和生产,具有巨大的应用价值和经济价值。
Description
技术领域
本发明涉及生物技术领域,具体地涉及一个植物育性调控基因GMS2、GMS2编码蛋白、GMS2的基因敲除突变体,以及GMS2基因、蛋白和突变体在杂交育种中的应用。
背景技术
水稻是世界上最重要的粮食作物之一。随着人口的增长和生活品质的提升,据预计到2050年水稻的年产量要提高1-2倍才能满足人类发展的需求。杂交水稻是父母本杂交后获得的子一代,其产量往往比常规稻亲本提高15%以上,抗性和适应性也远胜于亲本。因此,应用和推广杂交水稻是提高水稻产量的一个重要途径。
雄性不育系是杂交水稻育制种技术的关键节点。雄性不育系是指雄配子发育异常而丧失生育能力,雌配子发育正常的植物株系。它只能作为母本接受父本的花粉,自交不能结实。目前杂交水稻生产上应用的雄性不育系有核质互作型和光温敏型两种。核质互作型雄性不育系的不育基因在细胞质中,细胞核中没有育性恢复基因。当细胞核中有育性恢复基因的恢复系与其配组杂交时可以生产可育的子一代杂交种,当细胞核中没有育性恢复基因而细胞质中也没有不育基因的保持系与其杂交时可以繁殖不育系种子。由于需要不育系、保持系和恢复系三系配套,这种杂交水稻育制种技术常被称为“三系法”。一些控制核质互作型不育及相应育性恢复的基因已经被克隆(Chen and Liu,2014,Male sterility andfertility restoration in crops,Annu Rev Plant Biol,65:579-606)。核质互作型不育系是杂交水稻育制种中第一种大规模应用的不育系,为杂交水稻产业的建立和发展奠定了材料基础。然而由于核质互作型不育系的组配受到恢复系基因型的限制,导致只有约5%的种质资源能被利用。而细胞质的不育基因有导致米质差、特定病虫害流行的潜在风险。
光温敏型雄性不育系是一种育性受光温环境调控的不育系。在一定的光温条件下这种不育系保持不育,可用于组配杂交。当条件改变时不育系恢复育性,可用于不育系繁殖。由于光温敏雄性不育系实现了不育系和保持系的合二为一,只需要父本与其配组生产子一代杂交种,因此相应的育制种技术常被称为“两系法”。调控光温敏雄性不育的基因在细胞核中,目前已经克隆的基因包括PMS3、TMS5、CSA和TMS10(Chen and Liu,2014,Malesterilityand fertility restoration in crops,Annu Rev Plant Biol,65:579-606;Zhou H,et al,2014,RNase ZS1processes UbL40mRNAs and controls thermosensitivegenic male sterility in rice,Nature Communications,5:4884-4892)。与核质互作型不育系相比,光温敏型不育系繁殖程序简单,配组因恢复基因广泛存在而更自由。光温敏不育系的大规模应用极大地巩固和推动了杂交水稻产业发展。然而,由于该型不育系的育性受光温环境影响,也导致制种风险高,制种地域受到限制。
为了克服目前杂交水稻育制种技术中存在的关键性缺陷,创造和利用新类型的不育系将是重要的突破口。细胞核雄性不育由细胞核基因突变产生,有显性突变和隐性突变,有孢子体基因突变和配子体基因突变。显性突变和配子体基因突变只能通过雌配子遗传,隐性突变既可通过雌配子也可通过雄配子进行遗传,而且遵循孟德尔定律。本发明提供了一种植物育性调控基因及基于该基因突变所产生的隐性核不育类型的雄性不育系。该不育系育性稳定,只受核编码的单基因调控,不受光温环境的影响。该不育系的育性恢复基因广泛存在于水稻种质资源中,也可以通过转野生型基因恢复育性。该基因和该基因突变产生的不育系为研发水稻新型杂交育制种技术提供了元件,为解决现有技术存在的问题奠定了基础。
发明内容
本发明的目的是提供一种植物育性相关蛋白、其编码基因以及通过操作该基因在调控植物雄性生育力中的应用。非限制性地举例而言,下文描述的任何方法都可与本发明所提供的植物育性相关蛋白的相应核苷酸序列一起使用,例如,使植株中所述植物育性相关蛋白的内源编码序列突变、向植株中引入该序列的反义序列、使用发卡形式、或将其与其它核苷酸序列连接起来调控植株的表型,或者是本领域技术人员己知的可用于影响植株的雄性生育力的多种方法中的任一方法。
本发明在水稻中发现了一个具有雄性育性调控功能的花粉发育调控基因GMS2。GMS2位于水稻第4号染色体上,其在粳稻品种日本晴中的基因组核苷酸序列如SEQ ID NO:1所示,CDS序列如SEQ ID NO:2所示,氨基酸序列如SEQ ID NO:3所示。在籼稻品种9311中其基因组核苷酸序列如SEQ ID NO:4所示,其CDS序列如SEQ ID NO:69所示,其氨基酸序列与粳稻相同。在琴叶拟南芥(Arabidopsis lyrata)中该育性基因的氨基酸序列如SEQ ID NO:9所示;在香蕉(Musa acuminata)中该育性基因的氨基酸序列如SEQ ID NO:10所示;非洲栽培稻(Oryza glaberrima)中该育性基因的氨基酸序列如SEQ ID NO:11所示;在短药野生稻(Oryza brachyantha)中该育性基因的氨基酸序列如SEQ ID NO:12所示;大麦(Hordeumvulgare)中该育性基因的氨基酸序列如SEQ ID NO:13所示:高粱(Sorghum bicolor)中该育性基因的氨基酸序列如SEQ ID NO:14所示;玉米(Zea mays)中该育性基因的氨基酸序列如SEQ ID NO:15所示;小米(Setaria italica)中该育性基因的氨基酸序列如SEQ ID NO:16所示。
上述所述育性基因,可从各种植物中分离获得。本领域技术人员应该理解,本发明所述的育性基因包括与GMS2基因高度同源,并且具有同样的育性调控功能的高度同源的功能等价序列。所述高度同源的功能等价序列包括在严谨条件下能够与本发明所公开的GMS2基因的核苷酸序列杂交的DNA序列。本发明中所使用的“严谨条件”是公知的,包括诸如在含400mM NaCl、40mM PIPES(pH6.4)和l mM EDTA的杂交液中于60℃杂交12-16小时,然后在65℃下用含0.1%SDS、和0.1×SSC的洗涤液洗涤15-60分钟。
功能等价序列还包括与本发明所公开的GMS2基因的核苷酸序列具有至少85%、90%、95%、96%、97%、98%、或99%序列相似性,且具有育性调控功能的DNA序列,可以从任何植物中分离获得。其中,序列相似性的百分比可以通过公知的生物信息学算法来获得,包括Myers和Miller算法(Bioinformatics,4(1):1117,1988)、Needleman-Wunsch全局比对法(J Mol Biol,48(3):443-453,1970)、Smith-Waterman局部比对法(J Mol Biol,147:195-197,1981)、Pearson和Lipman相似性搜索法(PNAS,85(8):2444-2448,1988)、Karlin和Altschul的算法(Altschul等,J Mol Biol,215(3):403-410,1990;PNAS,90:5873-5877,1993)。这对于本领域技术人员来说是熟悉的。
基于本发明的上述发现,本发明的第一个方面是提供一种植物雄性育性相关蛋白,所述植物雄性育性相关蛋白为如下(1)或(2)所述的蛋白:
(1)具有SEQ ID NO:3、9、10、11、12、13、14、15或16所示氨基酸序列的蛋白;
(2)将SEQ ID NO:3、9、10、11、12、13、14、15或16经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的具有调控植物雄性育性活性的蛋白。
本发明提供编码所述植物雄性育性相关蛋白的核酸。
本发明所述的核酸可以从任何植物中分离获得,包括但不限于芸苔属、玉米、小麦、高梁、短药野生稻、非洲栽培稻、短柄草属、两节荠属、白芥、草麻子、芝麻、棉籽、亚麻子、大豆、拟南芥属、菜豆属、花生、茵宿、燕麦、油菜籽、大麦、燕麦、黑麦(Rye)、小米、蜀黍、小黑麦、单粒小麦、斯佩尔特小麦(Spelt)、双粒小麦、亚麻、格兰马草(Gramma grass)、摩擦禾、假蜀黍、羊茅、多年生麦草、甘廉、红莓苔子、番木瓜、香蕉、红花、油棕、香瓜、苹果、黄瓜、石角斗、剑兰、菊花、百合科、棉花、校、向日葵、芸苔、甜菜、咖啡、观赏植物和松类等。优选地,植物包括玉米、小米、拟南芥、二穗短柄草、大豆、红花、芥菜、小麦、大麦、黑麦、短药野生稻、非洲栽培稻、棉花和高粱。
以水稻为例,所述核酸的序列为以下任一:
(1)具有SEQ ID NO:1或2所示的核苷酸序列的核酸;
(2)具有SEQ ID NO:4或69所示的核苷酸序列的核酸;
(3)在严格条件下能够与(1)、(2)之任一所述序列的DNA杂交的DNA片段;
(4)与(1)、(2)之任一所述序列互补的DNA片段;
(5)在(1)、(2)之任一所述序列的基础之上,经过一至数个碱基的替换和/或一至数个碱基的插入和/或缺失,或经过大片段的核苷酸序列插入/缺失/易位/倒位所形成能够影响植物花粉生育能力的DNA片段;
(6)与(1)、(2)之任一所述序列的DNA片段具有85%、90%、95%、96%、97%、98%、或99%以上的同一性且编码水稻雄性育性相关蛋白的DNA片段。
本发明提供编码所述植物雄性育性相关蛋白的核酸的抑制因子,所述抑制因子导入植物中能够使编码所述植物雄性育性相关蛋白的核酸的表达量降低、不表达或发生突变而失活。所述抑制因子可为蛋白或核酸(包括但不限于反义基因、siRNA及其DNA、dsRNA及其DNA、sgRNA及其DNA等)。
本发明提供一种生物材料,其含有编码所述植物雄性育性相关蛋白的核酸,或含有编码所述植物雄性育性相关蛋白的核酸的抑制因子,所述生物材料为表达盒、载体、宿主细胞、转基因细胞系或转基因植物。
本发明提供一种植物、植物组织或植物细胞,其表现为雄性不育性状,由编码所述植物雄性育性相关蛋白的核酸的突变所造成,所述突变为一个或多个核苷酸的缺失、插入或取代突变,或为通过反义基因的转入、共抑制或发夹结构的引入所产生的突变;所述突变导致所述的植物雄性育性相关蛋白的表达量降低、不表达或失活。
所述植物、植物组织或植物细胞可通过自然突变或人工诱变获得,可以是转基因植物、植物组织或植物细胞也可以是非转基因植物、植物组织或植物细胞。
所述人工诱变包括理化诱变、插入突变、基因打靶敲除、反义基因的转入、共抑制或发夹结构的引入等。
所述植物包括但不限于芸苔属、玉米、小麦、高梁、短药野生稻、非洲栽培稻、短柄草属、两节荠属、白芥、芝麻、大豆、拟南芥属、菜豆属、花生、茵宿、燕麦、油菜籽、大麦、燕麦、黑麦(Rye)、小米、蜀黍、小黑麦、单粒小麦、斯佩尔特小麦(Spelt)、双粒小麦、亚麻、格兰马草(Gramma grass)、摩擦禾、假蜀黍、羊茅、多年生麦草、甘廉、红莓苔子、番木瓜、香蕉、红花、油棕、香瓜、苹果、黄瓜、石角斗、剑兰、菊花、百合科、棉花、向日葵、芸苔、甜菜、咖啡、观赏植物和松类等。优选地,包括玉米、小米、拟南芥、二穗短柄草、芥菜、小麦、大麦、黑麦、短药野生稻、非洲栽培稻、棉花和高粱。
可选地,所述植物、植物组织或植物细胞为采用CRISPR-Cas9方法获得,所述CRISPR-Cas9方法使用的靶序列位于编码所述植物雄性育性相关蛋白的核酸的序列中,靶序列的反向互补序列具有5’-(N)X-NGG-3’结构,其中,N表示A,T,C和G中的任意一个,X为19或20nt的任意核苷酸序列。
具体地,所述植物、植物组织或植物细胞衍生系为采用CRISPR-Cas9方法以GCGGTCGGTGGCGGCCATGG(SEQ ID NO:17)和CGCCTCCCTCGCCGTCGCG G(SEQ ID NO:18)为靶位点得到的在靶位点或靶位点相邻区域发生突变的植物。
本发明的第二个方面提供所述植物雄性育性相关蛋白或编码所述植物雄性育性相关蛋白的核酸或所述核酸的抑制因子或所述生物材料或所述植物、植物组织或植物细胞的如下任一种应用:
(1)在调控植物雄性育性中的应用;
(2)在制备雄性不育植物中的应用;
(3)在恢复由编码所述植物雄性育性相关蛋白的核酸突变导致的隐性核不育的雄性育性中的应用;
(4)在植物杂交育种中的应用;
(5)在植物种质资源改良中的应用。
上述(1)中,调控植物雄性育性可为使植物的雄性育性降低或丧失。具体可通过调控植物雄性生殖细胞、花粉的发育实现。其中,使植物的雄性育性降低或丧失可通过将植物中所述植物雄性育性相关蛋白的编码基因突变,使其表达量降低或不表达,或者通过将编码所述植物雄性育性相关蛋白的核酸的抑制因子引入植物中实现。
上述(2)中,所述雄性不育植物为隐性核不育系,带有编码所述植物雄性育性相关蛋白的核酸的纯合突变。
上述(3)中,通过将编码所述植物雄性育性相关蛋白的核酸引入植株来恢复由所述植物雄性育性相关蛋白突变或失活导致的植株隐性核不育的雄性育性,以便导入外源基因从而获得优质的转基因作物。
上述(4)中,使用带有编码所述植物雄性育性相关蛋白的核酸的纯合突变的隐性核不育系进行杂交育、制种。
上述(5)中,所述改良包括产量提高、品质提高、抗病虫害、抗逆、抗倒伏等。
以上所述植物是自花授粉或异花授粉作物,包括但不限于水稻、玉米、小麦、高梁。
本发明的第三个方面是提供通过影响所述植物雄性育性相关蛋白或编码该蛋白的核酸的序列,或者通过影响该核酸的转录、翻译从而影响植物育性的方法。所述影响植物育性是指使所述植物的育性发生改变,如导致植物雄性不育。具体地,取决于实际应用需求,可以通过多种方法来影响所述植物雄性育性相关蛋白或编码该蛋白的核酸的序列或其在植物体内的表达和翻译,从而达到调控植物雄性育性的效果。更具体地,影响所述植物雄性育性相关蛋白或编码该蛋白的核酸的序列或其在植物体内的表达和翻译可以使用许多本领域普通技术人员可获得的工具进行,例如,通过理化诱变、插入突变、基因打靶敲除、反义基因的转入、共抑制或发夹结构的引入等,都可以用于破坏所述植物雄性育性相关蛋白的正常表达,从而获得雄性不育的植株。
本发明的第四个方面是提供所述植物雄性育性相关蛋白的突变体,其为在所述植物雄性育性相关蛋白的编码基因中插入、和/或缺失、和/或取代若干个核苷酸得到,该突变体能够导致水稻雄性不育。
本发明提供一种适用于CRISPR-Cas9方法对编码所述植物育性相关蛋白的核酸进行定向敲除的靶位点,其为靶位点1:GCGGTCGGTGGCGGCCATGG(SEQ ID NO:17)和/或靶位点2:CGCCTCCCTCGCCGTCGCGG(SEQ ID NO:18)。
本发明还提供特异性靶向上述靶位点1和靶位点2的sgRNA。
含有上述sgRNA的DNA序列的CRISPR-Cas9打靶载体也属于本发明的保护范围。
本发明的第五个方面是提供所述靶位点或靶向该靶位点的sgRNA或含有该sgRNA的DNA的CRISPR-Cas9打靶载体的如下任一种应用:
(1)在调控植物雄性育性中的应用;
(2)在制备雄性不育植物中的应用;
(3)在植物杂交育种中的应用;
(4)在植物种质资源改良中的应用。
上述(1)中,调控植物雄性育性可为使植物的雄性育性降低或丧失。具体可通过调控植物雄性生殖细胞、花粉的发育实现。其中,使植物的雄性育性降低或丧失可通过将植物中所述植物雄性育性相关蛋白的编码基因突变,使其表达量降低或不表达,或者通过将编码所述植物雄性育性相关蛋白的核酸的抑制因子引入植物中实现。
上述(2)中,所述雄性不育植物为隐性核不育系,带有编码所述植物雄性育性相关蛋白的核酸的纯合突变。
上述(3)中,使用所述核苷酸的抑制子是所述育性调控蛋白失活,从而创制出隐性核雄性不育的植株,以便用于杂交育、制种。
上述(4)中,所述改良包括产量提高、品质提高、抗病虫害、抗逆、抗倒伏等。
本发明还提供一种制备雄性不育植物的方法,其为使植物中的所述植物雄性育性相关蛋白的表达量降低、不表达或失活。
作为本发明的一种优选方案,本发明提供利用CRISPR-Cas9技术制备雄性不育水稻的方法,其为利用CRISPR-Cas9技术敲除或突变水稻中编码所述植物育性相关蛋白的核酸。
具体地,利用CRISPR-Cas9技术以靶位点GCGGTCGGTGGCGGCCATGG(SEQ ID NO:17)和/或靶位点CGCCTCCCTCGCCGTCGCGG(SEQ ID NO:18)为靶位点,使得靶位点或靶位点及相邻核苷酸序列突变。
本发明还提供了一种获得GMS2基因在植物中的直系同源基因片段的方法,以及利用该方法获得拟南芥、香蕉、非洲栽培稻、短药野生稻、大麦、高粱、玉米、小米同源GMS2的氨基酸序列及其应用。
本发明提供的一种获得GMS2基因在植物中的直系同源基因片段的方法包括:使用前述GMS2基因的DNA片段在核苷酸数据库中进行blastx搜索;获得的所有Identities大于或等于35%、Positives大于或等于50%即为GMS2基因的直系同源的基因片段。
与现有技术相比,本发明具有如下的有益效果:本发明提供的水稻花粉发育调控基因GMS2直接参与花粉发育调控,该基因被敲除或表达受到抑制后,花粉完全不育,导致植物雄性不育。本发明利用CRISPR-Cas9基因编辑技术对GMS2基因进行了基因编辑,获得了GMS2基因突变的水稻雄性不育突变体。由GMS2突变导致的水稻不育突变体与现有三系和两系不育系相比,其不育性状稳定,不受环境条件影响。利用本发明的GMS2基因及其突变体可通过转基因等方法培育新的核不育系并提供恢复所述不育系育性的方法,为水稻核不育系培育和繁殖奠定了基础,在农作物的杂种优势利用和不育化杂交种制种生产中都将发挥重要作用。
附图说明
图1为本发明实施例2中灌浆期野生型(左)和gms2突变体(右)的植株形态。
图2为本发明实施例2中野生型(左)和gms2突变体(右)的小穗形态。
图3为本发明实施例2中野生型(左)和gms2突变体(右)穗开花的形态。
图4为本发明实施例2中解剖后野生型(左)和gms2突变体(右)的小花形态。
图5为本发明实施例2中野生型(左)和gms2突变体(右)的花药形态。
图6为本发明实施例2中野生型(左)和gms2突变体(右)花粉碘染。
图7为本发明实施例4中利用InD48490标记鉴定定位群体中不育单株的基因型。上带大小149bp,下带大小140bp。左边前2个泳道的DNA模板分别为gms2突变体和明恢63,后面泳道为定位群体中的不育单株。
图8A为本发明实施例4中GMS2基因图位克隆图。
图8B为本发明实施例4中gms2突变体的突变位点示意图。
图9为本发明实施例4中GMS2基因在9311(48490-9311),明恢63(48490-MH63),日本晴(48490-Nip)和gms2突变体(48490-3148)材料中的核苷酸序列差异。有差异的地方用灰色背景突出。每行最后一个核苷酸在整个基因序列中的位置标示在行末。起始密码子ATG和终止密码子TGA分别用方框标出。
图10为本发明实施例4中GMS2编码蛋白在9311(48490-9311)和gms2突变体(48490-3148)中的氨基酸序列差异。有差异的地方用浅灰色背景突出。每行最后一个氨基酸残基在整个蛋白序列中的位置标示在行末。
图11为本发明实施例4中GMS2杂合株后代基因型鉴定。上带大小149bp,下带大小140bp。箭头所指为来自雄性不育的样品。
图12为本发明实施例5中GMS2在水稻不同组织和不同发育时期幼穗中的表达量。花1-花9代表幼穗发育的颖花原基分化期至花粉成熟时期。
图13为本发明实施例6中pC9M-GMS2载体示意图。T1代表靶位点1,T2代表靶位点2。
图14A为本发明实施例6中利用CRISPR-Cas9系统对GMS2基因敲除后部分转基因阳性植株的靶位点序列分析。
图14B为本发明实施例6中转基因植株PC9M-GMS2-Line17在靶位点1和靶位点2处的测序峰图。其中,在靶位点1处的测序峰图中,箭头指向缺失位点;在靶位点2处的测序峰图中,箭头指向插入位点。
图15为本发明实施例6中GMS2野生型(左)、敲除植株PC9M-GMS2-Line17(右)的整株形态。
图16为本发明实施例6中GMS2野生型(左)、敲除植株PC9M-GMS2-Line17(右)的颖壳形态。
图17为本发明实施例6中GMS2野生型(左)、敲除植株PC9M-GMS2-Line17(右)的花药形态。
图18为本发明实施例6中GMS2野生型(左)、敲除植株PC9M-GMS2-Line17(右)的花粉碘染结果。
图19为本发明实施例7中pUbi1301-48490-CDS载体示意图。
图20为本发明实施例7中GMS2在超表达植株中的RT-PCR表达量分析。柱状图为对RT-PCR胶图中的条带亮度进行量化后,用48490的亮度值分别除以对应GAPDH的亮度值得到的结果。
图21为本发明实施例8中pC1300-48490-genome载体示意图。
图22为本发明实施例8中野生型植株(左)和gms2突变体互补植株(右)的植株形态。
图23为本发明实施例8中野生型植株(左)和gms2突变体互补植株(右)的颖壳形态。
图24为本发明实施例8中野生型植株(左)和gms2突变体互补植株(右)的花药形态。
图25为本发明实施例8中野生型植株(左)和gms2突变体互补植株(右)的花粉碘染结果。
图26为本发明实施例9中水稻GMS2基因编码蛋白与其他物种基因组中的同源蛋白的序列比对图。包括琴叶拟南芥(Arabidopsis lyrata)蛋白AT3G60900.1、香蕉(Musaacuminata)蛋白GSMUA_Achr11P03090_001、非洲栽培稻(Oryza glaberrima)蛋白ORGLA04G0194100.1、短药野生稻(Oryza brachyantha)蛋白OB04G29380.1、大麦(Hordeumvulgare)蛋白MLOC_7985.1、高粱(Sorghum bicolor)蛋白Sb06g026030.1、玉米(Zea mays)蛋白GRMZM2G003752_P01、小米(Setaria italica)蛋白Si010135m。NxYL保守序列用方框标出。
图27为本发明实施例9中水稻GMS2基因编码蛋白系统进化树分析。
具体实施方式
以下的实施例便于更好地理解本发明,但并不限定本发明的应用范围。下述实施例中的所有技术和科学术语,如无特殊说明,均为本发明所属领域普通技术人员通常所理解的相同含义。除非有相反指明,本发明所使用或提及的技术均为本领域普通技术人员公认的标准技术。所述试验材料,如无特别注明,均为本发明领域通用的试验材料。下述实施例中所用的试验试剂,如无特殊说明,均为自常规生化试剂商店购买得到的。
本发明所述的雄性不育,特指由植物细胞核基因发生功能变化导致植物雄性生殖器官发育出现异常(无法产生正常雄蕊、花药或者正常的雄性配子体)并出现育性的丧失,即通常所说的雄性核不育(Genic male sterility)而非细胞质核不育(Cytoplasmic malesterility)。雄性生殖器官育性的异常和恢复均由细胞核内的基因加以控制。
因此,本发明也包括利用序列表所述序列调控植株的雄配子生育能力,即利用本发明提供的基因序列在基因组、和/或转录组、和/或蛋白质组水平影响其它植物中相同或同源基因的功能从而达到控制雄性生殖器官育性的目的。例如,包括但不限于下述方法:通过天然序列的变异导致基因表达抑制或蛋白质功能的丧失、通过向植物中转入所述基因的反义序列或引入发卡结构、或将所述基因与其它序列(DNA或RNA)相结合产生新的具有功能活性的DNA或RNA链,来影响或改变植物基因的功能。或其它本领域技术人员己知的可用于影响植物雄性育性的技术方法中的任何一种技术方法。
本发明包括水稻GMS2基因,其显性等位基因对植物雄性育性具有关键作用,功能缺失性的隐性等位基因会导致雄性不育。该基因位于水稻4号染色体,其基因具体位置如图8A、图8B所示。
该基因序列及其同源序列可从各种植物中获得,包括但不限于卷柏(Selaginellamoellendorffii)、毛果杨(populus trichocarpa)、芜菁(Brassica rapa)、琴叶拟南芥(Arabidopsis lyrata)、拟南芥(Arabidopsis thaliana)、大豆(Glycine max)、马铃薯(Solanum tuberosum)、葡萄(Vitis vinifera)、小果野芭蕉(Musa acuminata)、小米(Setaria italica)、高粱(Sorghum bicolor)、玉米(Zea mays)、二穗短柄草(Brachypodium distachyon)、大麦(Hordeum vulgare)、短药野生稻(Oryzabrachyantha)、非洲栽培稻(Oryza glaberrima)、籼稻(Oryza sativa Indica Group)、粳稻(Oryza sativa Japonica Group)、小立碗藓(Physcomitrella patens)等。获得方法包括但不限于:使用水稻GMS2基因序列通过blastx、blastn或使用水稻GMS2氨基酸序列通过blastp从其它植物的基因组序列数据库、和/或cDNA序列数据库、和/或蛋白质序列数据库中调取;以水稻GMS2基因的DNA或cDNA或RNA序列为参考序列设计引物,从其它植物的基因组DNA或cDNA或RNA中利用PCR的方法直接获得;以水稻GMS2的基因序列设计探针,利用核酸杂交的方法从基因组文库中分离含有同源基因序列的DNA或cDNA或RNA片段。
GMS2基因同源序列指在与SEQ ID NO:3的氨基酸序列进行blastp比较分析后,Identities大于或等于35%、Positives大于或等于50%的植物基因序列。进行blastp时,所有参数均遵照http://blast.ncbi.nlm.nih.gov/所示的默认设置进行。
下文通过说明和阐述提供了更为详细的描述,但这并非意欲对本发明的范围加以限制。
实施例1水稻雄性不育突变体gms2的筛选
2013年6月用钴60辐射93-11种子10公斤得到M0代。辐射后的种子种植于海南省临高县试验田,成熟后分单株收种,共获得M1代材料约6500份。2014年春,选种子量较多的3617个M1代材料种植成株系,每个株系种50个单株。分别在分蘖期、孕穗期、抽穗期、开花期、灌浆期筛选株型、穗型、育性、产量等各种类型突变体,并收种保存。其中在编号为3148的株系中发现一个不育突变体,命名为gms2。
实施例2水稻雄性不育突变体gms2的表型分析
与野生型相比,gms2突变体植株(图1)和小穗(图2)形态正常,花期稍迟。内、外稃大小、小花开张尺寸、开张时间与野生型没有明显差异(图3)。体式显微镜下观察突变体小花形态,发现子房,花柱,柱头均比野生型略大(图4),但花药比野生型瘦小,颜色较浅(图5)。用碘-碘化钾溶液(0.6%KI,0.3%I2,w/w)溶液对花粉进行染色,如图6所示,野生型花粉粒大而圆并且被染成蓝黑色,而突变体花粉粒皱缩并且不能被染色。同一家系野生型植株套袋自交后正常结实,而gms2突变体不结实。而以水稻品种93-11为父本给gms2突变体授粉则可以结实。这表明该突变体为雄性不育突变体。
实施例3水稻雄性不育突变体gms2的遗传分析
在M5代种植gms2的分离群体80株,其中64株育性正常,16株不育,可育与不育株分离比符合3:1(χ2=0.57,P>0.05)。用gms2与93-11回交,F1代植株全部可育。在F2代种植gms2的分离群体70株,其中57株育性正常,13株不育,可育与不育株分离比符合3:1(χ2=0.85,P>0.05)。上述结果表明gms2的不育性状是由隐性单基因控制。
实施例4水稻雄性不育基因GMS2的克隆
使用图位克隆的方法对GMS2基因进行定位。以明恢63为父本与gms2突变体杂交构建了一个包含66棵突变植株的F2群体。利用该群体将GMS2定位于4号染色体SSR标记RM17332和RM280之间6861.252Kb范围内,与SSR标记RM303及Indel标记4826紧密连锁。GMS2基因与上述四个标记之间的交换单株分别为8个,1个,1个,32个。利用连锁标记挑选F2群体中的gms2杂合单株发展了一个F3群体,包含1937个突变体单株。在F3群体中RM303、4826、S10与GMS2基因之间的交换单株分别为10个,7个,8个。在RM303和S10之间通过分析和比较93-11和明恢63基因组的序列,开发并实验证实了5个新的单核苷酸多态性标记S4b,S3b,S2,S1,S8。在F3群体中,上述标记的交换单株分别为6个,6个,1个,4个,8个,(图7)。以S2上下游77kb为候选区段,发现在该区段内共有11个注释基因,其中LOC_Os04g48490预测编码一个成束蛋白样阿拉伯半乳聚糖蛋白,推测是GMS2基因。在日本晴中,LOC_Os04g48490基因组核苷酸序列长1582bp(记为48490-Nip,序列如SEQ ID NO:1),CDS核苷酸序列长1296bp(序列如SEQ ID NO:2),包含1个外显子(图8A和图8B),编码一个长432个氨基酸残基的蛋白(序列如SEQ ID NO:3)。用于定位GMS2基因的标记引物对序列如表1所示(SEQ ID NO.39-68)。
表1用于定位GMS2基因的标记引物对序列
根据48490-Nip序列设计引物对LOC_Os04g48490基因在93-11、明恢63和gms2突变体中的等位基因进行扩增和测序,引物序列如表2所示。所有PCR扩增均使用KOD FX DNAPolymerase(TOYOBO CO.,LTD.Life Science Department,Osaka,Japan),并按照产品说明的反应体系和条件,在Thermo scientific Arktik thermal cycler上进行PCR扩增。PCR产物送往南京金斯瑞生物科技有限公司进行测序。测序结果用DNAman 6.0进行拼接。93-11、明恢63和gms2突变体中的LOC_Os04g48490基因分别记为48490-9311(序列如SEQ ID NO:4),48490-MH63(序列如SEQ ID NO:5),48490-3148(序列如SEQ ID NO:6)。
表2用于扩增LOC_Os04g48490的引物对序列
对48490-9311、48490-3148、48490-MH63和48490-Nip进行多序列比对,结果如图9所示。48490-9311和48490-3148相比,48490-3148只在ATG后第118个碱基开始存在一个AACAGCTAC的缺失(图8B和图9)。氨基酸序列分析显示,该突变将导致LOC_Os04g48490基因编码蛋白中第40到第42位的天冬酰胺、丝氨酸和酪氨酸残基缺失(图10)。48490-MH63和48490-Nip与48490-3148也在ATG后118个碱基处存在上述相同差异(图9)。这表明ATG后118位碱基开始的AACAGCTAC的缺失突变是造成gms2突变体雄性不育的原因。此外48490-9311和48490-MH63的序列完全一致,而与48490-Nip相比,在第8位有一个A变C的SNP,在第109位有一个C变G的SNP,在1288位有一个T变C的SNP,在第1515位有一个G碱基的插入(图9)。两处核苷酸差异分别落在5’UTR和3’UTR中,另外两处核苷酸差异虽然落在外显子中,但并不影响编码蛋白。这表明LOC_Os04g48490基因在水稻中高度保守,其核苷酸序列即使在籼、粳亚种间也只存在4个碱基的差异,而蛋白质序列则没有差异。在93-11中LOC_Os04g48490的CDS核苷酸序列如SEQ ID NO:69所示,编码蛋白序列如SEQ ID NO:3。在gms2突变体中LOC_Os04g48490的CDS核苷酸序列和氨基酸序列分别如SEQ ID NO:7和SEQ ID NO:8所示。
基于LOC_Os04g48490基因突变位点的测序结果,在突变位点两侧设计特异引物InD48490_F:GCTCCGGCTGTTGATCT(SEQ ID NO:19)和InD48490_R:GCCTGCTCTTCCTCCTG(SEQID NO:20)。当InD48490_F和InD48490_R配对扩增野生型LOC_Os04g48490基因时将产生149bp条带,扩增突变型LOC_Os04g48490基因时将产生140bp条带。使用InD48490_F和InD48490_R引物对对41株gms2的M6分离群体进行基因型检测。如图11所示,野生型要么扩增出149bp和140bp两条带,要么扩增出149bp一条带,而不育突变体均只能扩增出140bp一条带。这说明突变基因型与不育表型共分离,LOC_Os04g48490就是GMS2基因。
实施例5 GMS2基因的表达分析
取93-11各时期组织提取总RNA,反转录成cDNA。使用引物InD48490_F:GCTCCGGCTGTTGATCT(SEQ ID NO:19)和InD48490_R:GCCTGCTCTTCCTCCTG(SEQ ID NO:20)检测GMS2基因的表达量,使用引物GAPDH-RTF:GAATGGCTTTCCGTGTT(SEQ ID NO:25)和GAPDH-RTR:CAAGGTCCTCCTCAACG(SEQ ID NO:26)检测内参基因GAPDH的表达量。采取实时定量PCR方法进行表达量分析。如图12所示,GMS2基因的表达量在根和茎中明显低于其它组织,在种子中明显高于其它组织。在茎节、叶、叶鞘和穗中,GMS2基因的表达量处于中等,但并不相同。在花1(花长1mm)、花2(花长2mm)、花3(花长3mm)、花4(花长4mm)、花5(花长5mm)、花6(花长5.5mm)、花7(花长6mm)、花8(花长7mm)、花9(花长8mm)到即颖花原基分化期至花粉成熟时期的穗中,GMS2的表达量呈现先降低后升高最后再降低的波动。
实施例6 GMS2基因敲除株系的获得及表型分析
利用CRISPR-Cas9系统对GMS2基因进行定向敲除。为了提高敲除效率,选择两个靶位点同时进行敲除。靶位点1位于外显子的负链上,序列为GCGGTCGGTGGCGGCCATGG(SEQ IDNO:17,位于序列SEQ ID NO:1的第45位第64位),靶位点2位于外显子的上,序列为CGCCTCCCTCGCCGTCGCGG(SEQ ID NO:18,位于SEQ ID NO:1序列的第85位至第104位)。根据Ma等(Ma X,et al.A Robust CRISPR-Cas9System for Convenient,High-EfficiencyMultiplex Genome Editing in Monocot and Dicot Plants.Mol Plant,2015,8:1274-84)的方法将靶位点1和靶位点2连入载体pC9M中,获得载体pC9M-GMS2(图13)。有pC9M-GMS2的大肠杆菌被命名为E.coli-pC9M-GMS2。将pC9M-GMS2通过电击转入农杆菌菌株EHA105中,得到的菌株命名为Ab-pC9M-GMS2。
利用重组农杆菌Ab-pC9M-GMS2侵染粳稻中花11(ZH11)愈伤组织,经潮霉素抗性筛选、分化、生根获得再生转基因株系25株。炼苗移栽后得到存活植株22株,提取所述植株叶片的总DNA,利用引物SP1:CCCGACATAGATGCAATAACTTC(SEQ ID NO:29)和SP2:GCGCGGTGTCATCTATGTTACT(SEQ ID NO:30)做阳性检测,全部为株阳性株。用靶位点1两侧的引物靶1-F:AAACCCACGCCCAGAAA(SEQ ID NO:31)和靶1-R:GCCAGGAGGAAGAGCAG(SEQ ID NO:32)以及靶位点2两侧的引物靶2-F:GCCTGCTCTTCCTCCTG(SEQ ID NO:33)和靶2-R:GTGCTCCGGCTGTTGAT(SEQ ID NO:34)扩增基因组DNA,扩增产物测序后与基因组进行比对。结果显示14株T0植株发生了基因编辑,其中一株发生纯合突变,8株T0代苗未被编辑。
植株PC9M-GMS2-Line17的基因组DNA在靶位点1和靶位点2处均发生了纯合突变,其中在靶位点1处发生了TG碱基缺失(SEQ ID NO:27),在靶位点2处发生了T碱插入(SEQ IDNO:28)(图14B)。PC9M-GMS2-Line1的基因组DNA在靶位点1处发生了双等位突变,其中等位基因1发生了A碱基插入,等位基因2发生了T碱基缺失;PC9M-GMS2-Line1的基因组DNA在靶位点2处也发生了双等位突变,在等位基因1发生了G/T碱基SNP,等位基因2发生了G/C碱基SNP。PC9M-GMS2-Line3的基因组DNA在靶位点1处发生了双等位突变,其中等位基因1发生了TG碱基缺失,等位基因2发生了A碱基插入;PC9M-GMS2-Line3的基因组DNA在靶位点2处也发生了双等位突变,在等位基因1发生了G/T碱基SNP,等位基因2发生了C碱基缺失。而转基因阴性单株PC9M-GMS2-Line2,PC9M-GMS2-Line5,PC9M-GMS2-Line7基因型并未改变(图14A)。
开花后对上述阳性株进行表型分析。与野生型ZH11相比,GMS2敲除植株PC9M-GMS2-Line17在株叶和小穗形态上并无明显差别(图15和图16)。但GMS2敲除植株的花药明显更加瘦小(图17)。花粉碘染结果表明,野生型ZH11的花粉大而圆,可以被染色,而GMS2敲除植株的花粉小而皱缩,不能被染色(图18)。其它GMS2双等位突变植株也表现出雄性不育的性状。
实施例7 GMS2基因超表达株系的获得及表型分析
以9311的RNA反转录产物为模板,用引物3148OX-F CggggtaccATGGCCGCCACCGAC:(SEQ ID NO:35)和3148OX-R:CGCggatccTCACAAGAACGACGC(SEQ ID NO:36)扩增获得带有GMS2完整编码核苷酸序列(SEQ ID NO:2)的DNA片段。将该片段用Kpn I和BamH I双酶切后连入pBLU5获得质粒pUbi1301-48490-CDS(图19)。有pUbi1301-48490-CDS的大肠杆菌被命名为E.coli-pUbi1301-48490-CDS。将pUbi1301-48490-CDS通过电击转入农杆菌菌株EHA105中,得到的菌株命名为Ab-pUbi1301-48490-CDS。
利用重组农杆菌Ab-pUbi1301-48490-CDS侵染粳稻中花11愈伤组织,经潮霉素抗性筛选、分化、生根获得转基因阳性植株6株。使用实时定量PCR方法,利用实施例5中的引物InD48490-F:GCTCCGGCTGTTGATCT(SEQ ID NO:19)和InD48490-R:GCCTGCTCTTCCTCCTG(SEQID NO:20),GAPDH-RTF:GAATGGCTTTCCGTGTT(SEQ ID NO:25)和GAPDH-RTR:CAAGGTCCTCCTCAACG(SEQ ID NO:26)对转基因阳性植株中GMS2的表达量进行分析。如图20所示,和转基因阴性单株2和8相比,超表达植株20和28中GMS2的表达量分别上升了9倍和45倍以上,但超表达植株并未出现与表达量共分离的明显表型,说明GMS2基因超表达对水稻表型并没有明显影响。
实施例8 gms2突变体转基因互补株系的获得及表型分析
以9311的基因组DNA为模板,用引物3148HB-F:CgcgtttcgaaatttTGATTTCTTCATCGCACT(SEQ ID NO:37)和3148HB-R:GtcgcgatcgcatgcACAACATGGTGCAACAGTG(SEQ ID NO:38)扩增获得带有GMS2起始密码子ATG上游2kb和终止密码子TGA下游515bp的基因全长片段。将该片段用Kpn I和BamH I双酶切后连入pC1300获得质粒pC1300-48490-genome(图21)。有pC1300-48490-genome的大肠杆菌被命名为E.coli-pC1300-48490-genome。将pC1300-48490-genome通过电击转入农杆菌菌株EHA105中,得到的菌株命名为Ab-pC1300-48490-genome。利用重组农杆菌Ab-pC1300-48490-genome侵染gms2突变体愈伤组织,经抗性筛选、分化、生根共获得转基因阳性植株4株,4株育性均恢复正常(图22、图23、图24和图25)。这进一步证明GMS2基因调控花粉发育,该基因突变会导致花粉败育。
实施例9 GMS2编码蛋白与其同源蛋白的序列比对及进化树分析
利用blastp工具在NCBI的Genbank数据库中对水稻GMS2基因编码蛋白的氨基酸序列进行同源性搜索,得到了琴叶拟南芥(Arabidopsis lyrata)、香蕉(Musa acuminata)、非洲栽培稻(Oryza glaberrima)、短药野生稻(Oryza brachyantha)、大麦(Hordeumvulgare)、高粱(Sorghum bicolor)、玉米(Zea mays)、小米(Setaria italica)基因组中预测的同源蛋白,将这些蛋白序列进行比对分析,结果显示来自不同植物的同源蛋白都具有非常相似的保守序列,彼此之间同源性很高(图26和图27),表明该蛋白在植物花的雄性器官发育过程中生物学功能保守,起着非常重要的作用。
在琴叶拟南芥(Arabidopsis lyrata)中该育性基因的氨基酸序列如SEQ ID NO:9所示;香蕉(Musa acuminata)中该育性基因的氨基酸序列如SEQ ID NO:10所示;非洲栽培稻(Oryza glaberrima)中该育性基因的氨基酸序列如SEQ ID NO:11所示;短药野生稻(Oryza brachyantha)中该育性基因的氨基酸序列如SEQ ID NO:12所示;大麦(Hordeumvulgare)中该育性基因的氨基酸序列如SEQ ID NO:13所示:高粱(Sorghum bicolor)中该育性基因的氨基酸序列如SEQ ID NO:14所示;玉米(Zea mays)中该育性基因的氨基酸序列如SEQ ID NO:15所示;小米(Setaria italica)中该育性基因的氨基酸序列如SEQ ID NO:16所示。
实施例10转育带有GMS2基因的隐性核不育系
用gms2突变体与育性正常的受体,如H28B,进行杂交、回交和自交,并在此过程中用分子标记进行gms2基因和遗传背景选择,最终获得H28B背景下带有纯合GMS2突变基因的隐性核不育系。具体实施步骤如下:
1、以受体亲本,如H28B,为父本与gms2杂交获得F1。
2、以F1为母本与受体亲本,如H28B,回交获得BC1F1。
3、种植BC1F1,使用引物InD48490_F:GCTCCGGCTGTTGATCT(SEQ ID NO:19)和InD48490_R:GCCTGCTCTTCCTCCTG(SEQ ID NO:20)检测gms2基因型。选择gms2杂合基因型,即同时能扩增出149bp和140bp条带的植株。
4、使用一组(例如100个,或200个等)基因型在gms2突变体和轮回亲本基因组之间存在多态性,且分布均匀的分子标记(可以是但不限于SSR、SNP、INDEL、EST、RFLP、AFLP、RAPD、SCAR等类型标记),对步骤3中选出的单株进行遗传背景鉴定,选取与轮回亲本基因型相似度高(如大于88%相似度,或2%中选率等)的植株。
5、用步骤4中选出的植株与受体亲本,如H28B,回交获得BC2F1。
6、种植BC2F1,重复步骤3和步骤4,选出gms2基因型杂合,遗传背景回复率高(如大于98%,或2%中选率等)的植株,收自交种BC2F2。
7、种植BC2F2,重复步骤3和步骤4,选出gms2基因型杂合,遗传背景纯合率最高的植株,收自交种BC2F3。BC2F3后代中分离的gms2纯合株即gms2隐性核不育系,BC2F3用于保存gms2隐性核不育系种质资源。
虽然,上文中已经用一般性说明及具体实施方案对本发明作了详尽的描述,但在本发明基础上,可以对之作一些修改或改进,这对本领域技术人员而言是显而易见的。因此,在不偏离本发明精神的基础上所做的这些修改或改进,均属于本发明要求保护的范围。
序列表
<110> 海南波莲水稻基因科技有限公司
<120> 水稻雄性育性调控基因、其应用以及利用CRISPR-Cas9调控水稻育性的方法
<130> KHP201111609.1
<160> 69
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1582
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
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ccgccgcctg ctcttcctcc tggccgcctc cctcgccgtc gcggcggtga gctcccacaa 120
catcacggac atcctcgacg gctacccgga gtactcgctg tacaacagct acctctccca 180
gaccaaggtg tgcgacgaga tcaacagccg gagcacggtc acctgcctcg tgctcaccaa 240
cggcgccatg tcctccctcg tctccaacct ctccctcgcc gacatcaaga acgcgctccg 300
cctcctcacc ctcctcgact actacgacac caagaagctg cactccctca gcgacggctc 360
cgagctcacc accacgctgt accagaccac cggcgacgcc tccggtaaca tgggccacgt 420
caacatcacc aacctgcgcg gcggcaaggt tgggttcgcc tccgccgcgc ccggctccaa 480
gttccaggcc acctacacca agtccgtcaa gcaggagccg tacaacctct ccgttcttga 540
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cctcaccgcg cttcttgaga aggccgggtg caagcagttc gcgcggctca tcgtgtcgtc 660
cggggtgatg aagatgtacc aggcggccat ggacaaggcg ctgacgctgt tcgcgcccaa 720
cgacgacgcg ttccaggcca agggcctgcc ggatctgagc aagctgacca gcgccgagct 780
ggtgacgctt ctgcagtacc acgccttgcc gcagtacgcg cccaaggcgt cgctcaagac 840
catcaagggc cacatccaga ccctggcctc caccggagcg ggtaagtacg acctctccgt 900
cgtcactaag ggcgacgacg tgtccatgga caccggcatg gacaagtccc gcgtcgcgtc 960
caccgtgctg gacgacaccc cgacggttat ccacacggtg gacagcgtgc tgctgccgcc 1020
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tgacatgtcg cccgccgacg cgcccgcggg agaagaggct gcagacaaag ccgagaagaa 1260
gaacggcgcc accgcggcgg ccacgagcgt tgcggccact gtggcctccg ccgccgctct 1320
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agctacctct cccagaccaa ggtgtgcgac gagatcaaca gccggagcac ggtcacctgc 180
ctcgtgctca ccaacggcgc catgtcctcc ctcgtctcca acctctccct cgccgacatc 240
aagaacgcgc tccgcctcct caccctcctc gactactacg acaccaagaa gctgcactcc 300
ctcagcgacg gctccgagct caccaccacg ctgtaccaga ccaccggcga cgcctccggt 360
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gcgcccggct ccaagttcca ggccacctac accaagtccg tcaagcagga gccgtacaac 480
ctctccgttc ttgaggtctc cgaccccatc accttccccg gcctcttcga ctccccgtcg 540
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gcgtcgctca agaccatcaa gggccacatc cagaccctgg cctccaccgg agcgggtaag 840
tacgacctct ccgtcgtcac taagggcgac gacgtgtcca tggacaccgg catggacaag 900
tcccgcgtcg cgtccaccgt gctggacgac accccgacgg ttatccacac ggtggacagc 960
gtgctgctgc cgccagagct cttcggtggc gcaccttccc ccgcgccggc gcccggaccg 1020
gcaagcgatg tgccagccgc ttctcccgcg ccagaaggct cctcgccggc gccctccccc 1080
aaggcggcgg gcaagaagaa aaagaagggc aagtcgcctt cccattcccc acccgcgcct 1140
ccggccgaca cgcctgacat gtcgcccgcc gacgcgcccg cgggagaaga ggctgcagac 1200
aaagccgaga agaagaacgg cgccaccgcg gcggccacga gcgttgcggc cactgtggcc 1260
tccgccgccg ctctgctcgc cgcgtcgttc ttgtga 1296
<210> 3
<211> 431
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 3
Met Ala Ala Thr Asp Arg Arg Leu Leu Phe Leu Leu Ala Ala Ser Leu
1 5 10 15
Ala Val Ala Ala Val Ser Ser His Asn Ile Thr Asp Ile Leu Asp Gly
20 25 30
Tyr Pro Glu Tyr Ser Leu Tyr Asn Ser Tyr Leu Ser Gln Thr Lys Val
35 40 45
Cys Asp Glu Ile Asn Ser Arg Ser Thr Val Thr Cys Leu Val Leu Thr
50 55 60
Asn Gly Ala Met Ser Ser Leu Val Ser Asn Leu Ser Leu Ala Asp Ile
65 70 75 80
Lys Asn Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Tyr Asp Thr Lys
85 90 95
Lys Leu His Ser Leu Ser Asp Gly Ser Glu Leu Thr Thr Thr Leu Tyr
100 105 110
Gln Thr Thr Gly Asp Ala Ser Gly Asn Met Gly His Val Asn Ile Thr
115 120 125
Asn Leu Arg Gly Gly Lys Val Gly Phe Ala Ser Ala Ala Pro Gly Ser
130 135 140
Lys Phe Gln Ala Thr Tyr Thr Lys Ser Val Lys Gln Glu Pro Tyr Asn
145 150 155 160
Leu Ser Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe
165 170 175
Asp Ser Pro Ser Ala Ala Ser Thr Asn Leu Thr Ala Leu Leu Glu Lys
180 185 190
Ala Gly Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Met
195 200 205
Lys Met Tyr Gln Ala Ala Met Asp Lys Ala Leu Thr Leu Phe Ala Pro
210 215 220
Asn Asp Asp Ala Phe Gln Ala Lys Gly Leu Pro Asp Leu Ser Lys Leu
225 230 235 240
Thr Ser Ala Glu Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Gln
245 250 255
Tyr Ala Pro Lys Ala Ser Leu Lys Thr Ile Lys Gly His Ile Gln Thr
260 265 270
Leu Ala Ser Thr Gly Ala Gly Lys Tyr Asp Leu Ser Val Val Thr Lys
275 280 285
Gly Asp Asp Val Ser Met Asp Thr Gly Met Asp Lys Ser Arg Val Ala
290 295 300
Ser Thr Val Leu Asp Asp Thr Pro Thr Val Ile His Thr Val Asp Ser
305 310 315 320
Val Leu Leu Pro Pro Glu Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro
325 330 335
Ala Pro Gly Pro Ala Ser Asp Val Pro Ala Ala Ser Pro Ala Pro Glu
340 345 350
Gly Ser Ser Pro Ala Pro Ser Pro Lys Ala Ala Gly Lys Lys Lys Lys
355 360 365
Lys Gly Lys Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Thr
370 375 380
Pro Asp Met Ser Pro Ala Asp Ala Pro Ala Gly Glu Glu Ala Ala Asp
385 390 395 400
Lys Ala Glu Lys Lys Asn Gly Ala Thr Ala Ala Ala Thr Ser Val Ala
405 410 415
Ala Thr Val Ala Ser Ala Ala Ala Leu Leu Ala Ala Ser Phe Leu
420 425 430
<210> 4
<211> 1583
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
ctccccaacg tgtcacacca caccacacaa caccaccacc gccgccatgg ccgccaccga 60
ccgccgcctg ctcttcctcc tggccgcctc cctcgccgtc gcggcggtca gctcccacaa 120
catcacggac atcctcgacg gctacccgga gtactcgctg tacaacagct acctctccca 180
gaccaaggtg tgcgacgaga tcaacagccg gagcacggtc acctgcctcg tgctcaccaa 240
cggcgccatg tcctccctcg tctccaacct ctccctcgcc gacatcaaga acgcgctccg 300
cctcctcacc ctcctcgact actacgacac caagaagctg cactccctca gcgacggctc 360
cgagctcacc accacgctgt accagaccac cggcgacgcc tccggtaaca tgggccacgt 420
caacatcacc aacctgcgcg gcggcaaggt tgggttcgcc tccgccgcgc ccggctccaa 480
gttccaggcc acctacacca agtccgtcaa gcaggagccg tacaacctct ccgttcttga 540
ggtctccgac cccatcacct tccccggcct cttcgactcc ccgtcggccg cgtcgaccaa 600
cctcaccgcg cttcttgaga aggccgggtg caagcagttc gcgcggctca tcgtgtcgtc 660
cggggtgatg aagatgtacc aggcggccat ggacaaggcg ctgacgctgt tcgcgcccaa 720
cgacgacgcg ttccaggcca agggcctgcc ggatctgagc aagctgacca gcgccgagct 780
ggtgacgctt ctgcagtacc acgccttgcc gcagtacgcg cccaaggcgt cgctcaagac 840
catcaagggc cacatccaga ccctggcctc caccggagcg ggtaagtacg acctctccgt 900
cgtcactaag ggcgacgacg tgtccatgga caccggcatg gacaagtccc gcgtcgcgtc 960
caccgtgctg gacgacaccc cgacggttat ccacacggtg gacagcgtgc tgctgccgcc 1020
agagctcttc ggtggcgcac cttcccccgc gccggcgccc ggaccggcaa gcgatgtgcc 1080
agccgcttct cccgcgccag aaggctcctc gccggcgccc tcccccaagg cggcgggcaa 1140
gaagaaaaag aagggcaagt cgccttccca ttccccaccc gcgcctccgg ccgacacgcc 1200
tgacatgtcg cccgccgacg cgcccgcggg agaagaggct gcagacaaag ccgagaagaa 1260
gaacggcgcc accgcggcgg ccacgagtgt tgcggccact gtggcctccg ccgccgctct 1320
gctcgccgcg tcgttcttgt gagcgtcagg tgttcgacgt tgagctctcg ttgttccccc 1380
ctgggcatgc atggtgtgat gcagtccggt gttcgcttct gagctcgtgg gctccatgga 1440
taatctcatc ctgaagttgt gttcttctct tcctggttgg tagtactcgg tagttagata 1500
ggatttgaat gattgggatc ctcaggtgga gaacggtgat tgtgatgcct attttgttag 1560
agctcggaac catgttttgt ttt 1583
<210> 5
<211> 1583
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
ctccccaacg tgtcacacca caccacacaa caccaccacc gccgccatgg ccgccaccga 60
ccgccgcctg ctcttcctcc tggccgcctc cctcgccgtc gcggcggtca gctcccacaa 120
catcacggac atcctcgacg gctacccgga gtactcgctg tacaacagct acctctccca 180
gaccaaggtg tgcgacgaga tcaacagccg gagcacggtc acctgcctcg tgctcaccaa 240
cggcgccatg tcctccctcg tctccaacct ctccctcgcc gacatcaaga acgcgctccg 300
cctcctcacc ctcctcgact actacgacac caagaagctg cactccctca gcgacggctc 360
cgagctcacc accacgctgt accagaccac cggcgacgcc tccggtaaca tgggccacgt 420
caacatcacc aacctgcgcg gcggcaaggt tgggttcgcc tccgccgcgc ccggctccaa 480
gttccaggcc acctacacca agtccgtcaa gcaggagccg tacaacctct ccgttcttga 540
ggtctccgac cccatcacct tccccggcct cttcgactcc ccgtcggccg cgtcgaccaa 600
cctcaccgcg cttcttgaga aggccgggtg caagcagttc gcgcggctca tcgtgtcgtc 660
cggggtgatg aagatgtacc aggcggccat ggacaaggcg ctgacgctgt tcgcgcccaa 720
cgacgacgcg ttccaggcca agggcctgcc ggatctgagc aagctgacca gcgccgagct 780
ggtgacgctt ctgcagtacc acgccttgcc gcagtacgcg cccaaggcgt cgctcaagac 840
catcaagggc cacatccaga ccctggcctc caccggagcg ggtaagtacg acctctccgt 900
cgtcactaag ggcgacgacg tgtccatgga caccggcatg gacaagtccc gcgtcgcgtc 960
caccgtgctg gacgacaccc cgacggttat ccacacggtg gacagcgtgc tgctgccgcc 1020
agagctcttc ggtggcgcac cttcccccgc gccggcgccc ggaccggcaa gcgatgtgcc 1080
agccgcttct cccgcgccag aaggctcctc gccggcgccc tcccccaagg cggcgggcaa 1140
gaagaaaaag aagggcaagt cgccttccca ttccccaccc gcgcctccgg ccgacacgcc 1200
tgacatgtcg cccgccgacg cgcccgcggg agaagaggct gcagacaaag ccgagaagaa 1260
gaacggcgcc accgcggcgg ccacgagtgt tgcggccact gtggcctccg ccgccgctct 1320
gctcgccgcg tcgttcttgt gagcgtcagg tgttcgacgt tgagctctcg ttgttccccc 1380
ctgggcatgc atggtgtgat gcagtccggt gttcgcttct gagctcgtgg gctccatgga 1440
taatctcatc ctgaagttgt gttcttctct tcctggttgg tagtactcgg tagttagata 1500
ggatttgaat gattgggatc ctcaggtgga gaacggtgat tgtgatgcct attttgttag 1560
agctcggaac catgttttgt ttt 1583
<210> 6
<211> 1574
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
ctccccaacg tgtcacacca caccacacaa caccaccacc gccgccatgg ccgccaccga 60
ccgccgcctg ctcttcctcc tggccgcctc cctcgccgtc gcggcggtca gctcccacaa 120
catcacggac atcctcgacg gctacccgga gtactcgctg tacctctccc agaccaaggt 180
gtgcgacgag atcaacagcc ggagcacggt cacctgcctc gtgctcacca acggcgccat 240
gtcctccctc gtctccaacc tctccctcgc cgacatcaag aacgcgctcc gcctcctcac 300
cctcctcgac tactacgaca ccaagaagct gcactccctc agcgacggct ccgagctcac 360
caccacgctg taccagacca ccggcgacgc ctccggtaac atgggccacg tcaacatcac 420
caacctgcgc ggcggcaagg ttgggttcgc ctccgccgcg cccggctcca agttccaggc 480
cacctacacc aagtccgtca agcaggagcc gtacaacctc tccgttcttg aggtctccga 540
ccccatcacc ttccccggcc tcttcgactc cccgtcggcc gcgtcgacca acctcaccgc 600
gcttcttgag aaggccgggt gcaagcagtt cgcgcggctc atcgtgtcgt ccggggtgat 660
gaagatgtac caggcggcca tggacaaggc gctgacgctg ttcgcgccca acgacgacgc 720
gttccaggcc aagggcctgc cggatctgag caagctgacc agcgccgagc tggtgacgct 780
tctgcagtac cacgccttgc cgcagtacgc gcccaaggcg tcgctcaaga ccatcaaggg 840
ccacatccag accctggcct ccaccggagc gggtaagtac gacctctccg tcgtcactaa 900
gggcgacgac gtgtccatgg acaccggcat ggacaagtcc cgcgtcgcgt ccaccgtgct 960
ggacgacacc ccgacggtta tccacacggt ggacagcgtg ctgctgccgc cagagctctt 1020
cggtggcgca ccttcccccg cgccggcgcc cggaccggca agcgatgtgc cagccgcttc 1080
tcccgcgcca gaaggctcct cgccggcgcc ctcccccaag gcggcgggca agaagaaaaa 1140
gaagggcaag tcgccttccc attccccacc cgcgcctccg gccgacacgc ctgacatgtc 1200
gcccgccgac gcgcccgcgg gagaagaggc tgcagacaaa gccgagaaga agaacggcgc 1260
caccgcggcg gccacgagtg ttgcggccac tgtggcctcc gccgccgctc tgctcgccgc 1320
gtcgttcttg tgagcgtcag gtgttcgacg ttgagctctc gttgttcccc cctgggcatg 1380
catggtgtga tgcagtccgg tgttcgcttc tgagctcgtg ggctccatgg ataatctcat 1440
cctgaagttg tgttcttctc ttcctggttg gtagtactcg gtagttagat aggatttgaa 1500
tgattgggat cctcaggtgg agaacggtga ttgtgatgcc tattttgtta gagctcggaa 1560
ccatgttttg tttt 1574
<210> 7
<211> 1287
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
atggccgcca ccgaccgccg cctgctcttc ctcctggccg cctccctcgc cgtcgcggcg 60
gtcagctccc acaacatcac ggacatcctc gacggctacc cggagtactc gctgtacctc 120
tcccagacca aggtgtgcga cgagatcaac agccggagca cggtcacctg cctcgtgctc 180
accaacggcg ccatgtcctc cctcgtctcc aacctctccc tcgccgacat caagaacgcg 240
ctccgcctcc tcaccctcct cgactactac gacaccaaga agctgcactc cctcagcgac 300
ggctccgagc tcaccaccac gctgtaccag accaccggcg acgcctccgg taacatgggc 360
cacgtcaaca tcaccaacct gcgcggcggc aaggttgggt tcgcctccgc cgcgcccggc 420
tccaagttcc aggccaccta caccaagtcc gtcaagcagg agccgtacaa cctctccgtt 480
cttgaggtct ccgaccccat caccttcccc ggcctcttcg actccccgtc ggccgcgtcg 540
accaacctca ccgcgcttct tgagaaggcc gggtgcaagc agttcgcgcg gctcatcgtg 600
tcgtccgggg tgatgaagat gtaccaggcg gccatggaca aggcgctgac gctgttcgcg 660
cccaacgacg acgcgttcca ggccaagggc ctgccggatc tgagcaagct gaccagcgcc 720
gagctggtga cgcttctgca gtaccacgcc ttgccgcagt acgcgcccaa ggcgtcgctc 780
aagaccatca agggccacat ccagaccctg gcctccaccg gagcgggtaa gtacgacctc 840
tccgtcgtca ctaagggcga cgacgtgtcc atggacaccg gcatggacaa gtcccgcgtc 900
gcgtccaccg tgctggacga caccccgacg gttatccaca cggtggacag cgtgctgctg 960
ccgccagagc tcttcggtgg cgcaccttcc cccgcgccgg cgcccggacc ggcaagcgat 1020
gtgccagccg cttctcccgc gccagaaggc tcctcgccgg cgccctcccc caaggcggcg 1080
ggcaagaaga aaaagaaggg caagtcgcct tcccattccc cacccgcgcc tccggccgac 1140
acgcctgaca tgtcgcccgc cgacgcgccc gcgggagaag aggctgcaga caaagccgag 1200
aagaagaacg gcgccaccgc ggcggccacg agtgttgcgg ccactgtggc ctccgccgcc 1260
gctctgctcg ccgcgtcgtt cttgtga 1287
<210> 8
<211> 428
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 8
Met Ala Ala Thr Asp Arg Arg Leu Leu Phe Leu Leu Ala Ala Ser Leu
1 5 10 15
Ala Val Ala Ala Val Ser Ser His Asn Ile Thr Asp Ile Leu Asp Gly
20 25 30
Tyr Pro Glu Tyr Ser Leu Tyr Leu Ser Gln Thr Lys Val Cys Asp Glu
35 40 45
Ile Asn Ser Arg Ser Thr Val Thr Cys Leu Val Leu Thr Asn Gly Ala
50 55 60
Met Ser Ser Leu Val Ser Asn Leu Ser Leu Ala Asp Ile Lys Asn Ala
65 70 75 80
Leu Arg Leu Leu Thr Leu Leu Asp Tyr Tyr Asp Thr Lys Lys Leu His
85 90 95
Ser Leu Ser Asp Gly Ser Glu Leu Thr Thr Thr Leu Tyr Gln Thr Thr
100 105 110
Gly Asp Ala Ser Gly Asn Met Gly His Val Asn Ile Thr Asn Leu Arg
115 120 125
Gly Gly Lys Val Gly Phe Ala Ser Ala Ala Pro Gly Ser Lys Phe Gln
130 135 140
Ala Thr Tyr Thr Lys Ser Val Lys Gln Glu Pro Tyr Asn Leu Ser Val
145 150 155 160
Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe Asp Ser Pro
165 170 175
Ser Ala Ala Ser Thr Asn Leu Thr Ala Leu Leu Glu Lys Ala Gly Cys
180 185 190
Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Met Lys Met Tyr
195 200 205
Gln Ala Ala Met Asp Lys Ala Leu Thr Leu Phe Ala Pro Asn Asp Asp
210 215 220
Ala Phe Gln Ala Lys Gly Leu Pro Asp Leu Ser Lys Leu Thr Ser Ala
225 230 235 240
Glu Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Gln Tyr Ala Pro
245 250 255
Lys Ala Ser Leu Lys Thr Ile Lys Gly His Ile Gln Thr Leu Ala Ser
260 265 270
Thr Gly Ala Gly Lys Tyr Asp Leu Ser Val Val Thr Lys Gly Asp Asp
275 280 285
Val Ser Met Asp Thr Gly Met Asp Lys Ser Arg Val Ala Ser Thr Val
290 295 300
Leu Asp Asp Thr Pro Thr Val Ile His Thr Val Asp Ser Val Leu Leu
305 310 315 320
Pro Pro Glu Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro Ala Pro Gly
325 330 335
Pro Ala Ser Asp Val Pro Ala Ala Ser Pro Ala Pro Glu Gly Ser Ser
340 345 350
Pro Ala Pro Ser Pro Lys Ala Ala Gly Lys Lys Lys Lys Lys Gly Lys
355 360 365
Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Thr Pro Asp Met
370 375 380
Ser Pro Ala Asp Ala Pro Ala Gly Glu Glu Ala Ala Asp Lys Ala Glu
385 390 395 400
Lys Lys Asn Gly Ala Thr Ala Ala Ala Thr Ser Val Ala Ala Thr Val
405 410 415
Ala Ser Ala Ala Ala Leu Leu Ala Ala Ser Phe Leu
420 425
<210> 9
<211> 424
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 9
Met Ala Val Ser Arg Ala Phe Ser Leu Phe Ala Phe Thr Leu Ser Leu
1 5 10 15
Leu Ala Val Ala Tyr Thr Val Ser Gly His Asn Ile Thr Gln Ile Leu
20 25 30
Ser Asp Thr Pro Glu Tyr Ser Ser Phe Asn Asn Tyr Leu Ser Gln Thr
35 40 45
Lys Leu Ala Asp Glu Ile Asn Ser Arg Thr Thr Ile Thr Val Leu Val
50 55 60
Leu Asn Asn Gly Ala Met Ser Ser Leu Ala Gly Lys His Pro Leu Ser
65 70 75 80
Val Val Lys Asn Ala Leu Ser Leu Leu Val Leu Leu Asp Tyr Tyr Asp
85 90 95
Pro Leu Lys Leu His Gln Leu Ala Lys Gly Ser Thr Leu Thr Thr Thr
100 105 110
Leu Tyr Gln Thr Thr Gly His Ala Pro Gly Asn Leu Gly Phe Val Asn
115 120 125
Val Thr Asp Leu Lys Gly Gly Lys Val Gly Phe Gly Ser Ala Ala Pro
130 135 140
Gly Ser Lys Leu Asp Ser Ser Tyr Thr Lys Ser Val Lys Gln Ile Pro
145 150 155 160
Tyr Asn Ile Ser Val Leu Glu Ile Asn Ala Pro Ile Ile Ala Pro Gly
165 170 175
Ile Leu Thr Ala Ala Ala Pro Ser Ser Gly Gly Val Asn Asn Leu Thr
180 185 190
Gly Leu Leu Glu Lys Ala Gly Cys Lys Thr Phe Ala Asn Leu Leu Val
195 200 205
Ser Ser Gly Val Leu Lys Thr Tyr Glu Ser Thr Val Glu Lys Gly Leu
210 215 220
Thr Val Phe Ala Pro Ser Asp Glu Ala Phe Lys Ala Lys Gly Val Pro
225 230 235 240
Asp Leu Thr Asn Leu Thr Gln Ala Glu Val Val Ser Leu Leu Glu Tyr
245 250 255
His Ala Leu Ala Glu Tyr Lys Pro Lys Gly Ser Leu Lys Thr Asn Lys
260 265 270
Asp Ala Ile Ser Thr Leu Ala Thr Asn Gly Ala Gly Lys Tyr Asp Leu
275 280 285
Thr Thr Ser Thr Ser Gly Asp Glu Val Ile Leu His Thr Gly Val Gly
290 295 300
Pro Ser Arg Leu Ala Asp Thr Val Val Asp Glu Thr Pro Val Val Ile
305 310 315 320
Phe Thr Val Asp Asn Val Leu Leu Pro Thr Glu Leu Phe Gly Lys Ser
325 330 335
Pro Ser Pro Ala Pro Ala Pro Ala Pro Glu Pro Val Ser Ala Pro Thr
340 345 350
Pro Ser Pro Ala Asn Ala Pro Ser Pro Val Glu Ala Pro Ser Pro Thr
355 360 365
Ala Ala Ser Pro Pro Ala Pro Pro Val Asp Glu Ser Ser Pro Glu Gly
370 375 380
Ala Pro Ser Asp Ser Pro Thr Ser Ser Glu Asn Ser Asn Ala Lys Asn
385 390 395 400
Ala Ala Leu His Val Thr Ala Pro Ala Leu Phe Thr Ala Leu Val Thr
405 410 415
Leu Ala Ala Thr Ser Leu Leu Ser
420
<210> 10
<211> 420
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 10
Met Ile Ser Ser Val Leu Leu Leu Leu Val Leu Leu Cys Gly Pro Trp
1 5 10 15
Ala Gly Cys Arg Ala His Asn Ile Thr Ala Ile Leu Glu Arg Tyr Pro
20 25 30
Glu Tyr Thr Leu Tyr Asn Ser Tyr Leu Thr Arg Thr Lys Val Cys Asp
35 40 45
Glu Ile Asn Ala His Glu Thr Val Thr Cys Leu Val Leu Asp Asp Gly
50 55 60
Ala Met Ser Thr Leu Ala Ala Lys Arg Pro Leu Ala Ala Ile Lys Asn
65 70 75 80
Ala Leu Arg Leu Leu Ala Leu Leu Asp Tyr Phe Asp Pro Pro Lys Leu
85 90 95
His Ala Leu Ser Ser Gly Thr Thr Leu Thr Thr Thr Leu Leu Gln Thr
100 105 110
Thr Gly Asn Glu Ala Gly Asn Leu Gly Phe Val Asn Ile Thr Asn Leu
115 120 125
Arg Gly Gly Arg Val Gly Phe Ala Ser Thr Ala Pro Gly Ser Lys Phe
130 135 140
Asp Ser Thr Tyr Thr Lys Ser Ile Glu Gln Ile Pro Tyr Asn Leu Ser
145 150 155 160
Val Leu Ala Val Ser Ala Pro Ile Val Phe Pro Gly Leu Leu Asp Thr
165 170 175
Pro Thr Ala Ala Ser Ser Asn Leu Thr Ala Leu Leu Glu Lys Ala Gly
180 185 190
Cys Lys Thr Phe Ala Arg Leu Leu Thr Thr Ser Gly Val Leu Lys Val
195 200 205
Phe Gln Asp Ala Met Ala Lys Gly Leu Thr Leu Phe Ala Pro Asn Asp
210 215 220
Glu Ala Phe Lys Ala Thr Asp Ala Pro Asp Leu Asn Ser Leu Ser Ser
225 230 235 240
Ala Glu Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Ser Tyr Thr
245 250 255
Pro Lys Ala Ser Leu Lys Ser Val Gly Gly Arg Leu Pro Thr Met Ala
260 265 270
Ser Ser Ala Ala Gly Lys Tyr Asp Leu Ser Val Val Ser Arg Gly Asp
275 280 285
Asp Val Ser Leu Asp Thr Gly Val Asp Thr Ser Arg Val Ala Ser Thr
290 295 300
Val Leu Asp Asp Thr Pro Val Cys Ile Leu Thr Val Asp Asn Leu Leu
305 310 315 320
Leu Pro Ile Glu Leu Phe Gly Ala Ala Pro Ser Pro Ala Pro Thr Pro
325 330 335
Ser Pro Ser Thr Ser Pro Val Glu Ala Pro Ala Pro Thr Pro Val Ala
340 345 350
Lys Ala Pro Ser Pro Lys Ser His Lys Lys His His Ser Pro Pro Ala
355 360 365
Pro Pro Met Ala Ser Pro Glu Ser Ala Pro Ser Asp Ala Pro Ala Ala
370 375 380
Ala Ala Asp Lys Ala Asp Val Lys Ser Ala Val Gly Val Ala Thr Pro
385 390 395 400
Ile Gly Thr Leu Ala Thr Val Ala Val Val Leu Ala Thr Leu Ala Met
405 410 415
Ala Ser Leu Pro
420
<210> 11
<211> 431
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 11
Met Ala Ala Thr Asp Arg Arg Leu Leu Phe Leu Leu Ala Ala Ser Leu
1 5 10 15
Ala Val Ala Ala Val Ser Ser His Asn Ile Thr Asp Ile Leu Asp Gly
20 25 30
Tyr Pro Glu Tyr Ser Leu Tyr Asn Ser Tyr Leu Ser Gln Thr Lys Val
35 40 45
Cys Asp Glu Ile Asn Ser Arg Ser Thr Val Thr Cys Leu Val Leu Thr
50 55 60
Asn Gly Ala Met Ser Ser Leu Val Ser Asn Leu Ser Leu Ala Asp Ile
65 70 75 80
Lys Asn Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Tyr Asp Thr Lys
85 90 95
Lys Leu His Ser Leu Ser Asp Gly Ser Glu Leu Thr Thr Thr Leu Tyr
100 105 110
Gln Thr Thr Gly Asp Ala Ser Gly Asn Met Gly His Val Asn Ile Thr
115 120 125
Asn Leu Arg Gly Gly Lys Val Gly Phe Ala Ser Ala Ala Pro Gly Ser
130 135 140
Lys Phe Gln Ala Thr Tyr Thr Lys Ser Val Lys Gln Glu Pro Tyr Asn
145 150 155 160
Leu Ser Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe
165 170 175
Asp Ser Pro Ser Ala Ala Ser Thr Asn Leu Thr Ala Leu Leu Glu Lys
180 185 190
Ala Gly Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Met
195 200 205
Lys Met Tyr Gln Ala Ala Met Asp Lys Ala Leu Thr Leu Phe Ala Pro
210 215 220
Asn Asp Asp Ala Phe Gln Ala Lys Gly Leu Pro Asp Leu Ser Lys Leu
225 230 235 240
Thr Ser Ala Glu Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Gln
245 250 255
Tyr Ala Pro Lys Ala Ser Leu Lys Thr Ile Lys Gly His Ile Gln Thr
260 265 270
Leu Ala Ser Thr Gly Ala Gly Lys Tyr Asp Leu Ser Val Val Thr Lys
275 280 285
Gly Asp Asp Val Ser Met Asp Thr Gly Met Asp Lys Ser Arg Val Ala
290 295 300
Ser Thr Val Leu Asp Asp Thr Pro Thr Val Ile His Thr Val Asp Ser
305 310 315 320
Val Leu Leu Pro Pro Glu Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro
325 330 335
Ala Pro Gly Pro Ala Ser Asp Val Pro Ala Ala Ser Pro Ala Pro Glu
340 345 350
Gly Ser Ser Pro Ala Pro Ser Pro Lys Ala Ala Gly Lys Lys Lys Lys
355 360 365
Lys Gly Lys Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Thr
370 375 380
Pro Asp Met Ser Pro Ala Asp Ala Pro Ala Gly Glu Glu Ala Ala Asp
385 390 395 400
Lys Ala Glu Lys Lys Asn Gly Ala Thr Ala Ala Ala Thr Ser Val Ala
405 410 415
Ala Thr Val Ala Ser Ala Ala Ala Leu Leu Ala Ala Ser Phe Leu
420 425 430
<210> 12
<211> 431
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 12
Met Ala Ala Pro Asp Arg Arg Leu Leu Phe Leu Leu Ala Val Ser Leu
1 5 10 15
Ala Val Ala Ala Val Ser Ser His Asn Ile Thr Asp Ile Leu Asp Gly
20 25 30
Tyr Pro Glu Tyr Ser Leu Tyr Asn Ser Tyr Leu Ser Gln Thr Lys Val
35 40 45
Cys Asp Glu Ile Asn Ser Arg Ser Thr Val Thr Cys Leu Val Leu Thr
50 55 60
Asn Gly Ala Met Ser Ser Leu Val Ser Asn Leu Ser Leu Ala Asp Ile
65 70 75 80
Lys Asn Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Tyr Asp Thr Lys
85 90 95
Lys Leu His Ser Leu Ser Asp Gly Ser Glu Leu Thr Thr Thr Leu Tyr
100 105 110
Gln Thr Thr Gly Asp Ala Ser Gly Asn Met Gly His Val Asn Ile Thr
115 120 125
Asn Leu Arg Gly Gly Lys Val Gly Phe Ala Ser Ala Ala Pro Gly Ser
130 135 140
Lys Phe Gln Ala Thr Tyr Thr Lys Ser Val Lys Gln Glu Pro Tyr Asn
145 150 155 160
Leu Ser Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe
165 170 175
Asn Ser Pro Ser Ala Ala Ser Thr Asn Leu Thr Ala Leu Leu Glu Lys
180 185 190
Ala Gly Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Ile
195 200 205
Lys Met Tyr Gln Ala Ala Met Asp Lys Gly Leu Thr Leu Phe Ala Pro
210 215 220
Asn Asp Asp Ala Phe His Ala Lys Asp Leu Pro Asp Leu Ser Lys Leu
225 230 235 240
Thr Ser Ala Glu Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Gln
245 250 255
Tyr Ala Pro Lys Ala Ser Leu Lys Thr Ile Lys Gly Asn Leu Gln Thr
260 265 270
Leu Ala Ser Thr Gly Ala Gly Lys Tyr Asp Leu Ser Val Val Ala Lys
275 280 285
Gly Asp Asp Val Ser Met Asp Thr Gly Val Asp Lys Ser Arg Val Ala
290 295 300
Ser Thr Val Leu Asp Asp Thr Pro Thr Val Ile His Thr Val Asp Ser
305 310 315 320
Val Leu Leu Pro Arg Glu Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro
325 330 335
Ala Ala Gly Pro Ala Ser Asp Val Pro Ala Ala Ser Pro Ala Pro Glu
340 345 350
Gly Ser Ser Pro Ala Pro Ser Pro Lys Ala Ala Gly Lys Lys Lys Lys
355 360 365
Lys Gly Lys Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Thr
370 375 380
Pro Asp Met Ser Pro Ala Asp Ala Pro Gln Gly Glu Glu Ala Ala Asp
385 390 395 400
Lys Ala Val Lys Lys Asn Gly Ala Thr Ala Val Ala Thr Ser Val Ala
405 410 415
Thr Thr Val Ala Ser Val Ala Val Leu Leu Ala Ala Ser Phe Leu
420 425 430
<210> 13
<211> 429
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 13
Met Ala Ala Gly Arg Arg Leu Leu Val Leu Leu Ala Val Ser Leu Ala
1 5 10 15
Ala Val Ala Ala Thr Arg Gly His Asn Ile Thr Glu Ile Leu Asp Gly
20 25 30
Tyr Ser Glu Tyr Ser Leu Tyr Asn Asn Tyr Leu Ser Gln Thr Lys Val
35 40 45
Cys Asp Glu Ile Asn Ser Arg Ser Thr Val Thr Ser Leu Val Leu Thr
50 55 60
Asn Gly Ala Met Ser Ser Leu Val Ala Asn Leu Ser Leu Ala Asp Val
65 70 75 80
Lys Asn Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Tyr Asp Pro Lys
85 90 95
Lys Leu His Ser Leu His Gly Gly Ser Glu Leu Thr Thr Thr Leu Tyr
100 105 110
Gln Thr Thr Gly Asp Ala Ser Gly Asp Met Gly His Val Asn Ile Thr
115 120 125
Ser Leu Arg Gly Gly Lys Val Gly Phe Ala Ser Ala Glu Pro Gly Ser
130 135 140
Lys Phe Gln Ala Thr Tyr Thr Lys Ser Ile Lys Glu Glu Pro Tyr Asn
145 150 155 160
Leu Ser Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe
165 170 175
Ser Ser Pro Ser Ala Ala Ser Thr Asn Leu Thr Ala Leu Leu Glu Lys
180 185 190
Ala Gly Cys Lys His Phe Ala Arg Leu Ile Val Ser Ser Gly Val Ile
195 200 205
Lys Thr Tyr Gln Ala Ala Met Asp Lys Gly Leu Thr Leu Phe Ala Pro
210 215 220
Asn Asp Asp Ala Phe Gln Ala Lys Gly Leu Pro Asp Leu Ser Lys Leu
225 230 235 240
Ser Ser Ala Asp Leu Val Ala Leu Leu Glu Tyr His Ala Leu Pro Gln
245 250 255
Tyr Ala Pro Lys Ala Ser Leu Lys Thr Met Lys Gly Gly Ile Pro Thr
260 265 270
Leu Ala Ser Thr Gly Lys Gly Lys Tyr Asp Leu Ser Val Val Ala Lys
275 280 285
Gly Asp Asp Val Ser Met Asp Thr Gly Met Asp Lys Ser Arg Val Ala
290 295 300
Ser Thr Val Leu Asp Asp Thr Pro Val Thr Val His Thr Val Asp Ser
305 310 315 320
Val Leu Leu Pro Pro Glu Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro
325 330 335
Gly Ala Ser Ala Asp Ser Pro Ala Ser Ala Pro Ala Pro Glu Thr Ser
340 345 350
Ala Pro Ala Pro Ser Pro Lys His Asp Lys Lys Lys Pro Lys Gly Lys
355 360 365
Ser Pro Ala His Ser Pro Pro Ala Pro Pro Ala Asp Ser Pro Asp Asn
370 375 380
Ala Pro Ala Asp Ala Pro Asp Gly Glu Gly Asp Glu Glu Ala Asp Lys
385 390 395 400
Ala Asp Ser Lys Asn Gly Ala Thr Ala Val Gly Met Ser Ile Ala Ala
405 410 415
Met Val Ala Ser Val Ala Leu Val Gly Ala Ser Leu Leu
420 425
<210> 14
<211> 429
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 14
Met Ala Ala Ser His His Leu Ile Leu Leu Val Leu Cys Leu Thr Ala
1 5 10 15
Ala Ala Ala Ser Ala His Asn Ile Thr Ala Ile Leu Asp Gly Arg Ser
20 25 30
Glu Tyr Thr Leu Tyr Asn Ser Tyr Leu Ser Gln Thr Lys Val Cys Asp
35 40 45
Glu Ile Asn Ser Arg Ser Ser Val Thr Val Leu Val Leu Thr Asn Gly
50 55 60
Ala Met Ser Ser Leu Val Ala Asn Leu Ser Leu Ala Asp Ile Lys Asn
65 70 75 80
Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Phe Asp Glu Lys Lys Leu
85 90 95
His Ser Ile Gly Ser Ser Ser Gln Leu Thr Thr Ser Leu Tyr Gln Thr
100 105 110
Thr Gly Gln Ala Ala Gly Asp Met Gly His Val Asn Ile Thr Asp Leu
115 120 125
Arg Gly Gly Lys Val Ala Phe Ala Ser Ala Ala Pro Gly Ala Lys Phe
130 135 140
Gln Ser Thr Tyr Thr Lys Arg Val Ala Asp Phe Pro Ser Asn Leu Ser
145 150 155 160
Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe Gly Ser
165 170 175
Pro Ser Ala Ser Ser Ala Asn Ile Thr Asp Leu Leu Glu Lys Ala Gly
180 185 190
Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Val Lys Thr
195 200 205
Tyr Gln Ala Ala Met Asp Lys Gly Leu Thr Leu Phe Ala Pro Asn Asp
210 215 220
Asp Ala Phe Lys Ala Lys Asp Leu Pro Asp Leu Ser Lys Leu Thr Ser
225 230 235 240
Ala Asp Leu Val Ala Leu Leu Gln Tyr His Ala Leu Pro Gln Tyr Ala
245 250 255
Pro Lys Ala Ser Leu Lys Val Ala Ser Gly Arg Ile Pro Thr Leu Ala
260 265 270
Ser Thr Gly Ala Gly Lys Tyr Asp Leu Thr Val Ala Ser Ser Gly Asp
275 280 285
Glu Val Ser Leu Asp Thr Gly Val Asp Lys Ser Arg Val Ala Ser Thr
290 295 300
Val Leu Asp Asp Pro Pro Thr Val Ile Leu Thr Val Asp Ser Val Leu
305 310 315 320
Leu Pro His Val Ile Phe Gly Gly Ala Pro Ser Pro Ala Pro Ala Pro
325 330 335
Gly Pro Ala Ala Asp Val Pro Ala Ser Ala Pro Ala Pro Glu Gly Ser
340 345 350
Ala Pro Ala Pro Thr Pro Lys Ala Ala Gly Lys Lys Lys Lys Lys Lys
355 360 365
Ala Lys Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Ser Pro
370 375 380
Asp Leu Ala Pro Ala Asp Ala Pro Asp Asp Asp Ala Ala Asp Lys Val
385 390 395 400
Glu Ser Lys Lys Asn Gly Ala Ala Ala Ala Ala Val Ser Phe Val Ala
405 410 415
Ser Val Ala Ser Ala Gly Leu Ala Val Ala Leu Leu Leu
420 425
<210> 15
<211> 429
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 15
Met Ala Ala Ser His Arg Leu Ile Leu Leu Val Leu Cys Leu Thr Ala
1 5 10 15
Thr Ala Ala Ser Ala His Asn Ile Thr Ala Ile Leu Asp Gly Arg Ser
20 25 30
Glu Tyr Thr Leu Tyr Asn Ser Tyr Leu Ser Glu Thr Lys Val Cys Asp
35 40 45
Glu Ile Asn Ser Arg Ser Ser Val Thr Val Leu Val Leu Thr Asn Gly
50 55 60
Ala Met Ser Ser Leu Val Ala Asn Leu Ser Leu Ala Asp Ile Lys Asn
65 70 75 80
Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Phe Asp Glu Lys Lys Leu
85 90 95
His Ser Ile Gly Ser Ser Ser Gln Leu Thr Thr Ser Leu Tyr Gln Thr
100 105 110
Thr Gly Gln Ala Ala Gly Asp Met Gly His Val Asn Ile Thr Asp Leu
115 120 125
Arg Gly Gly Lys Val Ala Phe Ala Ser Ala Ala Pro Gly Ala Lys Phe
130 135 140
Gln Ser Thr Tyr Thr Lys Arg Val Ala Asp Phe Pro Ser Asn Leu Ser
145 150 155 160
Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe Gly Ala
165 170 175
Pro Ser Ala Ser Ser Ala Asn Ile Thr Asp Leu Leu Glu Lys Ala Gly
180 185 190
Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Val Lys Thr
195 200 205
Tyr Gln Ala Ala Met Asp Lys Ala Leu Thr Leu Phe Ala Pro Asn Asp
210 215 220
Asp Ala Phe Lys Ala Lys Asp Leu Pro Asp Leu Ser Lys Leu Thr Ser
225 230 235 240
Ala Asp Leu Val Ala Leu Leu Gln Tyr His Ala Leu Pro Gln Tyr Ala
245 250 255
Pro Lys Ala Ser Leu Lys Val Ala Ser Gly Arg Ile Pro Thr Leu Ala
260 265 270
Ser Thr Ala Ala Gly Lys Tyr Asp Leu Ala Val Ala Ser Ser Gly Asp
275 280 285
Glu Val Thr Leu Asp Thr Gly Val Asp Lys Ser Arg Ile Ala Ser Thr
290 295 300
Val Leu Asp Asp Pro Pro Thr Val Ile Leu Thr Val Asp Ser Val Leu
305 310 315 320
Leu Pro His Val Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro Ala Pro
325 330 335
Gly Pro Ala Ala Asp Val Pro Ala Ser Ala Pro Ala Pro Glu Gly Ser
340 345 350
Ala Pro Ala Pro Ala Pro Lys Ala Ala Gly Lys Arg Lys Lys Lys Lys
355 360 365
Ala Arg Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ala Asp Ser Pro
370 375 380
Asp Met Ala Pro Ala Asp Ala Pro Thr Glu Asp Ala Ala Asp Lys Val
385 390 395 400
Glu Ser Lys Lys Asn Gly Gly Ala Ala Ala Ala Val Ser Phe Ala Ala
405 410 415
Ser Val Ala Ser Val Ala Leu Ala Val Ala Tyr Leu Leu
420 425
<210> 16
<211> 430
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 16
Met Ala Ala Ser His His Gly His Phe Phe Leu Leu Ala Leu Ser Ala
1 5 10 15
Ala Ala Asp Ser Ser Ser Thr His Asn Ile Thr Thr Val Leu Asp Gly
20 25 30
Arg Ser Glu Tyr Thr Leu Tyr Asn Ser Tyr Leu Ser Glu Thr Lys Val
35 40 45
Cys Asp Glu Ile Asn Ser Glu His Thr Val Thr Val Leu Val Leu Thr
50 55 60
Asn Gly Ala Met Ser Ser Leu Val Ala Asn Leu Ser Leu Ala Asp Ile
65 70 75 80
Lys Asn Ala Leu Arg Leu Leu Thr Leu Leu Asp Tyr Phe Asp Glu Lys
85 90 95
Lys Leu His Ser Leu Asp Ser Gly Ser Glu Leu Thr Thr Ser Leu Tyr
100 105 110
Gln Lys Thr Gly Gln Ala Ala Gly Asn Met Gly His Val Asn Ile Thr
115 120 125
Asp Leu Arg Gly Gly Lys Val Gly Phe Ala Pro Ala Ala Pro Gly Ala
130 135 140
Lys Phe Gln Ser Thr Tyr Thr Lys Arg Val Asp Glu Glu Pro Ser Thr
145 150 155 160
Leu Ser Val Leu Glu Val Ser Asp Pro Ile Thr Phe Pro Gly Leu Phe
165 170 175
Gly Ser Pro Ser Ala Ser Ser Val Asn Leu Thr Asp Leu Leu Glu Lys
180 185 190
Ala Gly Cys Lys Gln Phe Ala Arg Leu Ile Val Ser Ser Gly Val Val
195 200 205
Lys Met Tyr Gln Ala Ala Met Asp Lys Ala Leu Thr Leu Leu Ala Pro
210 215 220
Asn Asp Asp Ala Phe Lys Ala Lys Asp Leu Pro Asp Leu Ser Lys Leu
225 230 235 240
Ser Ser Ala Asp Leu Val Thr Leu Leu Gln Tyr His Ala Leu Pro Gln
245 250 255
Tyr Thr Pro Lys Ser Ser Leu Lys Val Ala Lys Gly Asp Ile Pro Thr
260 265 270
Leu Ala Ser Thr Gly Ala Gly Lys Tyr Asp Leu Ser Val Val Ser Ser
275 280 285
Gly Asp Asp Val Ser Leu Asp Thr Gly Lys Asp Lys Ser Arg Val Ala
290 295 300
Ser Thr Val Leu Asp Asp Thr Pro Thr Val Ile Leu Thr Val Asp Lys
305 310 315 320
Val Leu Leu Pro Pro Gly Leu Phe Gly Gly Ala Pro Ser Pro Ala Pro
325 330 335
Ala Pro Gly Pro Ala Ala Asp Val Pro Ala Ser Ala Pro Ala Pro Glu
340 345 350
Thr Ser Ala Pro Ala Pro Ser Pro Lys Ala Ala Gly Lys Lys Lys Lys
355 360 365
Lys Ala Lys Ser Pro Ser His Ser Pro Pro Ala Pro Pro Ser Asp Ser
370 375 380
Pro Asp Met Ala Pro Ala Asp Ala Pro Glu Gly Asp Ala Ala Asp Lys
385 390 395 400
Val Glu Ser Lys Lys Asn Gly Ala Ala Ala Ala Ala Ala Ser Phe Ala
405 410 415
Ala Thr Gly Ala Cys Val Ala Leu Ala Val Ala Ser Phe Leu
420 425 430
<210> 17
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 17
gcggtcggtg gcggccatgg 20
<210> 18
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 18
cgcctccctc gccgtcgcgg 20
<210> 19
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 19
gctccggctg ttgatct 17
<210> 20
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 20
gcctgctctt cctcctg 17
<210> 21
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 21
aaacagaaag ccccaatg 18
<210> 22
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 22
tgccgcagta cgcgcccaag 20
<210> 23
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 23
ttgtccatgc cggtgtccat 20
<210> 24
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 24
ggtcacggca caaactca 18
<210> 25
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 25
gaatggcttt ccgtgtt 17
<210> 26
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 26
caaggtcctc ctcaacg 17
<210> 27
<211> 75
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 27
tgtcacacca caccacacaa caccaccacc gccgccagcc gccaccgacc gccgcctgct 60
cttcctcctg gccgc 75
<210> 28
<211> 156
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 28
ccgccgcctg ctcttcctcc tggccgcctc cctcgccgtc gtcggcggtg agctcccaca 60
acatcacgga catcctcgac ggctacccgg agtactcgct gtacaacagc tacctctccc 120
agaccaaggt gtgcgacgag atcaacagcc ggagca 156
<210> 29
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 29
cccgacatag atgcaataac ttc 23
<210> 30
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 30
gcgcggtgtc atctatgtta ct 22
<210> 31
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 31
aaacccacgc ccagaaa 17
<210> 32
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 32
gccaggagga agagcag 17
<210> 33
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 33
gcctgctctt cctcctg 17
<210> 34
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 34
gtgctccggc tgttgat 17
<210> 35
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 35
cggggtacca tggccgccac cgac 24
<210> 36
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 36
cgcggatcct cacaagaacg acgc 24
<210> 37
<211> 33
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 37
cgcgtttcga aattttgatt tcttcatcgc act 33
<210> 38
<211> 34
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 38
gtcgcgatcg catgcacaac atggtgcaac agtg 34
<210> 39
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 39
cggtacatca cggtatcaaa tcg 23
<210> 40
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 40
taaatgctgg agcgatgcta acc 23
<210> 41
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 41
gtgctctcca tgtcggatta tgc 23
<210> 42
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 42
caaggcaaca agattggtta gtgg 24
<210> 43
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 43
ataaaggagg agggcctcag atgg 24
<210> 44
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 44
cacggtttgg aggttggaag c 21
<210> 45
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 45
gcttggcatc tgcttctgtt gttgg 25
<210> 46
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 46
ctcgctgctg atcgaggtgt cg 22
<210> 47
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 47
atcgatgtag gtagagggac acc 23
<210> 48
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 48
cagatctagt cgacatggtt gg 22
<210> 49
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 49
acaccatctc tcttcttttt ctat 24
<210> 50
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 50
atatgggtag gtttggatat tcg 23
<210> 51
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 51
gtgtgtgtga gtaaaatcct agtgca 26
<210> 52
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 52
atttgtactc ctatgtttag aatagc 26
<210> 53
<211> 31
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 53
aaaaagtgtg tgtgagtaaa atcctagagc c 31
<210> 54
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 54
acaaatatat agcaaaatcg gtgacc 26
<210> 55
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 55
gtggttttgt ggatgttttg taact 25
<210> 56
<211> 31
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 56
aaaaaacaaa tatatagcaa aatcggttac g 31
<210> 57
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 57
aagtatttgt aatgcactat gtaaaggt 28
<210> 58
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 58
ttaagagcac acacttccaa taatatgt 28
<210> 59
<211> 33
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 59
aaaaaaagta tttgtaatgc actatgtaat ggc 33
<210> 60
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 60
ctgggcgcgg tgcggcgggc gaggc 25
<210> 61
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 61
ccgcctcagc gccaccgcca agctga 26
<210> 62
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 62
aaaaactggg cgcggtgcgg cgggcgtggt 30
<210> 63
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 63
aagttgtgtt tagcactatg ttattacg 28
<210> 64
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 64
tttagcataa taactactat tcatcatt 28
<210> 65
<211> 33
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 65
aaaaaaagtt gtgtttagca ctatgttatg aca 33
<210> 66
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 66
gcaggagaca cttggtgccg cctctc 26
<210> 67
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 67
gcagattatt ttcggtgggt cccgtctc 28
<210> 68
<211> 31
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 68
aaaaagcagg agacacttgg tgccgccact t 31
<210> 69
<211> 1296
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 69
atggccgcca ccgaccgccg cctgctcttc ctcctggccg cctccctcgc cgtcgcggcg 60
gtcagctccc acaacatcac ggacatcctc gacggctacc cggagtactc gctgtacaac 120
agctacctct cccagaccaa ggtgtgcgac gagatcaaca gccggagcac ggtcacctgc 180
ctcgtgctca ccaacggcgc catgtcctcc ctcgtctcca acctctccct cgccgacatc 240
aagaacgcgc tccgcctcct caccctcctc gactactacg acaccaagaa gctgcactcc 300
ctcagcgacg gctccgagct caccaccacg ctgtaccaga ccaccggcga cgcctccggt 360
aacatgggcc acgtcaacat caccaacctg cgcggcggca aggttgggtt cgcctccgcc 420
gcgcccggct ccaagttcca ggccacctac accaagtccg tcaagcagga gccgtacaac 480
ctctccgttc ttgaggtctc cgaccccatc accttccccg gcctcttcga ctccccgtcg 540
gccgcgtcga ccaacctcac cgcgcttctt gagaaggccg ggtgcaagca gttcgcgcgg 600
ctcatcgtgt cgtccggggt gatgaagatg taccaggcgg ccatggacaa ggcgctgacg 660
ctgttcgcgc ccaacgacga cgcgttccag gccaagggcc tgccggatct gagcaagctg 720
accagcgccg agctggtgac gcttctgcag taccacgcct tgccgcagta cgcgcccaag 780
gcgtcgctca agaccatcaa gggccacatc cagaccctgg cctccaccgg agcgggtaag 840
tacgacctct ccgtcgtcac taagggcgac gacgtgtcca tggacaccgg catggacaag 900
tcccgcgtcg cgtccaccgt gctggacgac accccgacgg ttatccacac ggtggacagc 960
gtgctgctgc cgccagagct cttcggtggc gcaccttccc ccgcgccggc gcccggaccg 1020
gcaagcgatg tgccagccgc ttctcccgcg ccagaaggct cctcgccggc gccctccccc 1080
aaggcggcgg gcaagaagaa aaagaagggc aagtcgcctt cccattcccc acccgcgcct 1140
ccggccgaca cgcctgacat gtcgcccgcc gacgcgcccg cgggagaaga ggctgcagac 1200
aaagccgaga agaagaacgg cgccaccgcg gcggccacga gtgttgcggc cactgtggcc 1260
tccgccgccg ctctgctcgc cgcgtcgttc ttgtga 1296
Claims (6)
1.植物雄性育性相关蛋白或编码所述植物雄性育性相关蛋白的核酸或含有所述核酸的生物材料在恢复由核苷酸序列如SEQ ID NO:1或SEQ ID NO:2所示核酸的突变导致的水稻隐性核不育的雄性育性中的应用;
所述植物雄性育性相关蛋白的氨基酸序列如SEQ ID NO:3所示。
2.根据权利要求1所述的应用,其特征在于,所述核酸的核苷酸序列如SEQ ID NO:1或2所示。
3.根据权利要求1所述的应用,其特征在于,所述生物材料为表达盒、载体、宿主细胞、转基因细胞系或转基因植物。
4.水稻、水稻组织或水稻细胞的如下任一种应用:
(1)在制备雄性不育水稻中的应用;
(2)在水稻杂交育种中的应用;
(3)在水稻种质资源改良中的应用;
所述水稻、水稻组织或水稻细胞表现为雄性不育性状,由核苷酸序列如SEQ ID NO:1或2所示核酸的突变所造成,所述突变导致氨基酸序列如SEQ ID NO:3所示的植物雄性育性相关蛋白的表达量降低、不表达或失活;
所述水稻、水稻组织或水稻细胞为采用CRISPR-Cas9方法以GCGGTCGGTGGCGGCCATGG和CGCCTCCCTCGCCGTCGCGG为靶位点得到的在靶位点或靶位点相邻区域发生突变的水稻、水稻组织或水稻细胞。
5.水稻基因组靶位点或靶向所述靶位点的sgRNA或含有所述sgRNA的CRISPR-Cas9打靶载体的如下任一种应用:
(1)在调控水稻雄性育性中的应用;
(2)在制备雄性不育水稻中的应用;
(3)在水稻杂交育种中的应用;
(4)在水稻种质资源改良中的应用;
所述靶位点为GCGGTCGGTGGCGGCCATGG和/或CGCCTCCCTCGCCGTCGCGG。
6.一种制备雄性不育水稻的方法,其特征在于,使水稻中氨基酸序列如SEQ ID NO:3所示的植物雄性育性相关蛋白的表达量降低、不表达或失活;
所述氨基酸序列如SEQ ID NO:3所示的植物雄性育性相关蛋白的表达量降低、不表达或失活为通过CRISPR-Cas9方法以GCGGTCGGTGGCGGCCATGG和CGCCTCCCTCGCCGTCGCGG为靶位点并在靶位点或靶位点相邻区域发生突变实现。
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