CN1250730C - 提高棉纤维品质的方法 - Google Patents
提高棉纤维品质的方法 Download PDFInfo
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- CN1250730C CN1250730C CNB008155623A CN00815562A CN1250730C CN 1250730 C CN1250730 C CN 1250730C CN B008155623 A CNB008155623 A CN B008155623A CN 00815562 A CN00815562 A CN 00815562A CN 1250730 C CN1250730 C CN 1250730C
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Abstract
本发明涉及一种在植物体内控制纤维素合成,以便优化植物产品的产量水平和品质的方法。具体地讲,本发明提供了一种转基因棉花植物,这种植物具有较高的棉纤维和种子产量。本发明还提供了用于提高由棉花植物所产生的棉纤维的品质的方法。本发明还提供改变植物的纤维素与其他干重成分的比例的通用方法,用于改变细胞壁的厚度,用于提高产量和改变其他植物纤维的品质,用于提高种子产量,以及用于提高光合效率对于夜晚低温的承受能力。
Description
发明领域
本发明涉及通过改变蔗糖磷酸酯合成酶表达提高植物产品的产量或品质的方法。具体地讲,本发明提供了相对非转基因棉花植物而言具有较高含量的蔗糖磷酸酯合成酶的转基因棉花植物。还提供了用于提高由棉花植物所产生的棉纤维的产量或品质以及棉籽产量的方法。提供用于调节细胞壁厚度、用于提高产量和其他植物纤维品质、用于调节植物的纤维素与其他干重成分的比例,用于提高种子产量、以及用于提高光合效率对夜晚低温的承受能力的通用方法。
发明背景
通过温度控制纤维素生产的高比例及其调控对于农业来说是重要的,因为所有植物生长(以及所有粮食作物的生产)取决于由植物的营养部分或生殖部分进行的用于建造细胞壁的纤维素的合成。植物体的原始细胞壁中的纤维素还具有直接的商业价值,它可以作为动物饲料的可消化部分,并用于生产增稠剂、乙醇、以及其他纤维素基或纤维素衍生产品。另外,基于次级细胞壁的具有高纤维素含量的植物部分包含在经济上重要的植物产品中或构成了经济上重要的植物产品,包括棉纤维、木材、以及饲料作物中的纤维。木材和棉花,以及诸如大麻和亚麻的其他纤维作物的农业产量和产品品质在很多程度上是由纤维素的生物合成决定的。因此,了解纤维素合成的基础调控基质以及它是如何对温度胁迫做出反应的,可以通过遗传工程对作物进行有利的改变(改善产品产量和品质)。
由于棉纤维重量中超过90%是纤维素,因此棉花是一种特殊的作物,通过增加流向纤维素生产的碳可以提高产量和品质,如果能在包括会妨碍棉纤维发育的夜晚低温在内的在棉花生产大田中会遇到的各种环境条件下实现上述目的的话,将是特别有利的结果。例如,已知夜晚低温会妨碍棉纤维的季节性产量和品质(Gipson,温度对生长、发育和纤维特性的影响,参见Mauney著,棉花生理学,棉花结晶:孟菲斯,47-56页),因为夜晚低温会妨碍纤维素合成的效率(Roberts等,循环温度对培养的棉花胚珠中的纤维代谢的影响,植物生理学,100:979-986,1992)。操纵棉花产量和纤维品质参数和在不同和/或胁迫性环境条件下保持或改善上述参数的能力,使得能够通过遗传工程对作物进行有利的改变(概述产品品质)。
对生产者来说,棉纤维的产量是作物价值的最重要的决定因素。著名的棉花育种家最近已经指出,棉花生产已经达到了纤维产量的平台区,这预示着生产者将无法在经济上获得成功,因为要投入更高的成本。造成这一问题的潜在因素包括棉纤维和种子发育对环境的敏感性,商业化棉花的窄的遗传学基础,以及最近通过与转化过的陆地棉Coker312栽培种回交而导入的诸如抗除草剂和抗虫的转基因特征。Coker312(C312)是一种因为其高的再生能力而经常被用于转化的老的栽培种。通过遗传工程使得棉花作物生产对胁迫更有承受能力,增强栽培棉花的遗传学潜力,以及改善具有各种新性状的转化过的棉花的产量,将必然导致作物产量的提高。
类似地,对于棉花生产者来说种子产量是具有价值的,因为种子被出售用于油料生产和动物饲料生产。其他微量成分,每一粒种子上的短绒纤维提供了种子作物的附加的经济价值。在不损害皮棉纤维产量或品质的前提下提高种子和短绒纤维产量将有利于生产者在每英亩棉花生产中获得更高的收益。对于棉花种子来说,提高任何种子作物的产量对于农业都有重大好处。
改善了的棉纤维品质参数,如纤维细度、成熟比例、长度、长度均匀度、纤维束强度和单纤维强度是纺织工业生产高品质产品并充分利用现代纺织技术所需要的。对于棉花生产者来说,纤维品质参数也应当高到足以避免在它将棉花卖给轧棉人时使价格打折扣的程度。例如,在德克萨斯南部高原的短的生长期中,生产者经常会由于纤维细度低而在价格上遭受损失。通过育种技术能获得的越来越高的品质业已成为棉花工业的必要标准,并且市场压力可能改变,因此,生产者越来越需要通过优质棉花获得更高价格的报偿。因此,通过遗传工程稳定或提高在各种环境条件下纤维的品质可以提高棉花作物生产的收益,并提供可以被加工行业所利用的新型材料特性。
其他植物纤维,尽管通常具有不同的组织来源,但它与棉纤维具有相同的结构特征,那就是它们是具有富含纤维素的细胞壁的拉长了的细胞。与棉纤维一样,具有工业用途的其他植物纤维也必须具有由以下因素所决定的高品质,如纤维素含量和细胞壁厚度、直径、细度(或粗糙度)、长度、强度、耐用度、均匀度、弹性和延伸性。对于每一种特定的纤维来源和工业用途来说,都有一个最佳的上述参数的范围。以在造纸中进行纸浆生产以后的木材纤维为例,较长的纤维长度和较高的单纤维拉伸度都能提高纸张的抗拉强度。另外,纤维壁的厚度能提高纸浆产量和具有高的抗拉强度的吸水纸的产量。不过,较薄的纤维壁可以改善纤维的收缩,并有利于纤维间的相互结合,这有助于生产优质输血用纸。因此,有必要沿负方向或正方向控制各种纤维的细胞壁厚度和其他纤维品质参数,以便改善其产量或品质或拓宽其工业应用范围。
提高每英亩作物产量和用途是可持续农业的关键因素。提高同一种作物的多种产物的产量,如种子和纤维产量将是有用的。类似地,提高进行成功地作物收获的可能性,例如,在不损害种子作物产量的前提下通过生产具有能更好地抗倒伏的具有较坚固的茎干的植物也是有利的。
增加空气中的二氧化碳含量是受到关注的问题,因为预测它与地球温度升高有关。需要能通过将二氧化碳转化成有用产物而更好地固定二氧化碳的植物,特别是转化成在燃烧后通常不会再生二氧化碳的产物。
在白天,棉花叶片将大部分碳同化成淀粉,并在夜晚将淀粉转化成蔗糖以便转运到储存器官。正如上文所述,棉纤维不能很好地适应在寒冷的夜晚将蔗糖有效用于纤维素合成。因此,在随后的温暖的白天,寒冷的夜晚会降低棉花光合效率(Warner等,棉花碳代谢对夜晚温度的反应,农艺杂志,87:1193-1197,1995),这可能是由于在夜晚妨碍了碳水化合物的利用。所导致的叶片的碳水化合物的积累对会发出下调光合基因的信号。在寒冷的夜晚之后,保留在叶片中的过多的淀粉可能与某些负反馈基质相关,由于这种基质即使在回暖之后也会降低光合效率。有必要利用遗传工程手段减轻在随后的温暖的白天时间与寒冷的夜晚相关的对光合作用的抑制。
蔗糖磷酸酯合成酶(SPS)是参与植物碳代谢的一种关键蛋白(参见图1)。SPS能催化由UDP-葡萄糖和果糖6-磷酸酯形成蔗糖磷酸酯。在叶片中,SPS在控制还原碳在淀粉和可转运的蔗糖之间的分配方面起着重要作用(Huber等,蔗糖磷酸酯合成酶在高等植物中的作用和调控,植物生理学植物分子生物学年度综述,47:431-44,1996)。在生长中的储存细胞中,本发明的数据表明了SPS参与指导碳向纤维素的流动。其活性水平可以调节流向纤维素合成与流向呼吸的代谢流量(参见图2)。根据这一模型,纤维素储存细胞中的SPS可以在下面一种或两种情况下促进蔗糖合成提高纤维素合成的速度而增强储存强度:(a)如果从叶片中转运来的蔗糖在进入储存细胞之前或之后被分解,释放出葡萄糖和果糖的话,和/或(b)将在蔗糖合成酶的作用下释放的果糖重新用在由纤维素合成酶催化的UDP-葡萄糖和果糖通道上。较低的SPS活性水平能以类似的基质减弱储存强度,在任何情况下,储存填充是受蔗糖含量影响的。
在番茄上,业已证实SPS的超量表达有时候总的果实干重提高32%。这种提高不是由于单个果实重量的增加,而是由于果实数量增加了50%(Micallef等,在具有增强了的蔗糖合成能力的转基因番茄植物中改变了的光合作用、开花和结果,植物,196:327-334,1995)。业已证实这些番茄植物有时候每一个果实具有较高的鲜果重量和较高的可溶性固体(糖)(Laporte等,用玉米蔗糖磷酸酯合成酶转化过的番茄植物的蔗糖磷酸酯合成酶活性和产量分析,植物,203:253-259,1997)。这些报导没有提供有关种子产量的信息,因为相对番茄果实而言番茄种子的重量很小,并且没有将种子与果实分离进行称重。
应当指出的是,尽管棉铃和番茄在植物学上都被分类成果实,但这两种果实的性质以及它们所包含的种子的相对重要性有很大不同。番茄果实主要是充满了水的初级细胞壁的液囊,并以可溶性葡萄糖、果糖和蔗糖作为储存碳水化合物。这些糖在干燥时结晶,构成果实的干重。在果实中,番茄种子由于其体积小而不是主要的储存器官,并且除了用于繁殖番茄之外它们没有经济价值。在番茄上果实是主要的储存器官;它构成了几乎所有的番茄产量,并且是具有明显经济价值的唯一的番茄部分。
相反,棉花果实比较干燥并且壁也较薄。果实本身并不构成棉花的主要储存器官或者是决定棉花产量。它保护着种子直到棉铃打开,随后它就枯萎。果实没有或者少有经济价值(综合而言)。棉籽以及所附着的纤维构成了棉花作物上的重要经济价值的两个主要部分。棉纤维是棉籽壳的伸长了的表皮细胞。在植物学上将其定义为毛。因此,组织上的两个主要储存器官是:(1)种子胚胎的子叶,它储存油和蛋白;和(2)种子表皮毛(棉纤维)的次级细胞壁,它储存不可溶性纤维素。成熟的棉籽或果实中没有储存大量的可溶性糖。棉籽以及棉籽上所粘接的纤维构成了棉花作物的全部产量。因此,棉花以及其他产纤维的植物与番茄明显不同。
在番茄叶片上业已证实了能超量表达SPS的植物的营养部分的较高的总的干重。在同一个研究中,没有发现茎干重和根干重降低的变化(Galtier等,较高的蔗糖磷酸酯合成酶活性对番茄(番茄UC82B变种)光合作用、同化分配和生长的营养,植物生理学,101:535-543,1993)。番茄叶片不含有大量的主要由叶肉细胞和导管组织组成的纤维。相同的植物有时候表现出在整株植物基础上具有较高的干重(Ferrario-Mery等,在转化过的植物中控制蔗糖合成和氮同化途径以便改善光合作用和产量,Foyer著,高等植物初级代谢的分子途径,Taylor和Francis:纽约,125-153页,1997),并且在包括叶片和茎干的地上部分表现出较高的干重(Laporte等,用玉米蔗糖磷酸酯合成酶转化过的番茄植物的蔗糖磷酸酯合成酶活性和产量分析,植物,203:253-259,1997)。在超量表达SPS的马铃薯上,业已证实提高了的块茎的总的干重(Shewmaker,利用蔗糖磷酸酯合成酶编码序列改变可溶性固体,PCT国际公开号WO97/15678)。马铃薯块茎不含有大量的纤维。它主要是由具有初级细胞壁的薄壁细胞组成,这种细胞储存有大量淀粉和较少量蛋白。马铃薯块茎的主要产量成分是淀粉。以上所有报导都缺乏有关SPS超量表达对植物的细胞壁厚度、纤维素含量、以及纤维和种子产量影响的信息。不过,由于不能证实茎干重量增加,这一点与所分析过的番茄植物上纤维含量的提高矛盾。
业已证实了增强了的SPS表达能在番茄和拟南芥属上产生其他有利影响。在这两种植物上,叶片淀粉储存较少,以便优先用于蔗糖合成。在这两种植物上,光合作用的最大速度得到加强,在较高二氧化碳和饱和光照条件下增强最为明显(Galtier等,光照和大气二氧化碳富集对超量表达蔗糖磷酸酯合成酶的番茄植物叶片的光合作用和碳分配的影响,实验植物学杂志,46:1335-1344,1995;Micallef等,在具有增强了的蔗糖合成能力的转基因番茄植物上改变了的光合作用、开花和结果,植物,196:327-334,1995;和Signora等,在拟南芥中超量表达蔗糖磷酸酯合成酶,在通过二氧化碳富集延长生长之后会导致较高的叶片碳水化合物积累,实验植物学杂志,49:669-680,1998)。不过,这些报导没有提供有关寒冷的夜晚对温暖的白天的光合作用影响的信息。
因此,需要一种控制产纤维植物,特别是棉花的纤维素合成水平的方法。需要能够在各种环境条件下控制棉花的具有商业价值的纤维的产量和品质。总体上需要能够控制植物中纤维素的生物合成和细胞壁的厚度。总体上需要改善在夜晚低温下生长的植物的光合效率。能够提高作物的种子产量同样是重要的。本发明致力于解决这些要求并提供改进了的植物。
发明概述
本发明总体上涉及控制植物的纤维素合成以便优化来自该植物的产物的产量和品质的方法。
本发明包括控制具有农业或工业用途的任何植物细胞壁的纤维素含量、厚度或产量。所述细胞壁包括典型的薄的初级细胞壁,如在饲料中被消化的细胞壁以及存在于有用的农业残余物中的细胞壁,例如,在进行糖提取之后剩余的甜菜根薄壁细胞,这种细胞可以转化成增稠剂。所述细胞壁包括厚壁,如厚角组织和木质部薄壁组织的壁,这样的细胞壁决定着植物的硬度或决定饲料或其他农作物的产量和可消化性。所述细胞壁还包括二级细胞壁,如在纤维中经常见到的细胞壁。
具体地讲,本发明提供了相对非转基因棉花植物而言具有较高的蔗糖磷酸酯合成酶含量的转基因棉花植物。
本发明还提供了通过将嵌合的DNA结构导入棉花植物提高棉花植物产量的方法,所述DNA结构能改变蔗糖磷酸酯合成酶活性水平使其达到足以提高这种棉花植物的组织和纤维产量的程度。
本发明还可用于通过将嵌合的DNA结构导入棉花植物而提高由棉花植物所产生的棉纤维的品质,所述DNA结构能改变蔗糖磷酸酯合成酶活性水平使其达到足以提高这种棉花植物所产生的棉纤维的品质的程度。
本发明包括一种通过向植物中导入嵌合DNA提高光合效率对夜晚低温耐受能力的方法,所述嵌合DNA能改变蔗糖磷酸酯合成酶活性,达到增强光合效率对夜晚低温耐受能力的程度。
在另一种实施方案中,本发明提供了一种通过向植物中导入一种嵌合DNA结构调控该植物的纤维素与其他干重成分比例的方法,所述DNA结构能够改变蔗糖磷酸酯合成酶活性,达到能调控该植物的纤维素与其他干重成分比例的程度。
本发明还提供了一种通过向植物中导入嵌合DNA结构调控植物细胞壁厚度的方法,所述DNA结构能改变蔗糖磷酸酯合成酶活性,达到足以调控细胞壁厚度的程度。
在另一种实施方案中,本发明提供了一种通过向植物中导入嵌合DNA结构提高来自含纤维的植物的可收获的纤维产量的方法,所述DNA结构能够改变蔗糖磷酸酯合成酶活性,达到提高来自产纤维植物的可收获纤维产量的程度。
在另一种实施方案中,本发明提供了一种通过向植物中导入嵌合DNA结构提高来自产种子植物的可收获的种子产量的方法,所述DNA结构能够改变蔗糖磷酸酯合成酶活性,达到能提高来自产种子植物的可收获的种子产量的程度。
在另一种实施方案中,本发明提供了一种通过向植物中导入嵌合DNA结构改善来自产纤维植物的纤维品质的方法,所述DNA结构能够改变蔗糖磷酸酯合成酶活性,达到足以调控纤维品质的程度。所述改善可以表现为长度、强度和每单位长度的重量的改变。
附图简述
图1表示碳同化、淀粉合成和分解以及蔗糖合成途径。UDP-葡萄糖焦磷酸化酶催化葡萄糖1-磷酸酯(G-1-P)和UDP-葡萄糖之间的高度可逆的反应。蔗糖磷酸酯合成酶催化由UDP-葡萄糖和果糖6-磷酸形成蔗糖磷酸酯。
图2表示与纤维素生物合成有关的储存细胞中的代谢途径和酶。
图3是来自多种植物的SPS基因序列之间的氨基酸排比。
图4是菠菜叶片SPS基因序列和来自集胞藻属的同源序列之间的氨基酸排比。
图5是每粒种子纤维重量的矩形图,它表示在所有三种转基因品系中的提高(在这里以及所有随后的矩形图中,误差线是平均值的标准误差(平均值打印在每一个误差线上方)。
图6是每粒种子的去毛种子重量的矩形图。它表示所有三个转基因品系的升高。
图7是每粒种子的纤维重量的去毛种子重量的比例的矩形图。它表示这两种产量参数倾向于平行提高,在转基因品系中纤维重量的提高略占优势。
图8是每粒种子的纤维重量与每粒种子的去毛种子重量的分散曲线。它表示这两种参数以50%的水平相互依赖(在这里以及所有其他分散曲线中,R的平方由线性回归曲线计算出的决定系数。同样,将来自亲本C312的数据点标在其右侧,而来自三个转基因品系的数据点在左侧不做标记)。不过,要指出的是,C312没有表现出任何线性关系,因为在亲本品系中每粒种子的重量表现出很小的波动。因此,所有数据点之间的总体线性关系来自转基因植物。与亲本C312植物相比,转基因植物在每粒种子的去毛种子重量和每粒种子的纤维重量方面具有更大的差异和更高的水平。
图9是每粒种子的短纤维重量的矩形图。它表示在三个转基因品系中的两个上有所提高,而在一个转基因品系上降低。
图10是纤维细度的矩形图。它表示在所有三个转基因品系上有所提高。
图11是纤维细度与每粒种子纤维重量的分散曲线,表示这两种参数以60%的水平相互依赖。这一结果是合理的,因为每粒种子的纤维重量取决于三个重量:纤维的数量、纤维的长度、以及纤维壁厚度。在这三种因素中,纤维细度仅仅取决于纤维壁的厚度。要指出的是,这种线性关系也适用于C312,但转基因植物在每粒种子的纤维重量和纤维细度方面具有更高的值。
图12是断开单纤维所需粒的克数的矩形(Tb;g)。它表示在所有三种转基因品系中有所提高。
图13是拉断单纤维时伸长量的矩形图(占原始纤维长度的百分比)。它表示在所有三种转基因品系上有所提高。不过,要指出的是在转基因品系13-3a上的拉伸量最高,该品系在转基因品系中具有最低的断裂所需克数的提高。这表明这两种因素主要是由不同的纤维特性决定的,正像在理论上所预测的那样,并且通过下面的分散曲线得到证实。
图14是断开单纤维所需功的矩形图(μJ)。功是由断裂所需克数和拉伸量计算出来的综合因素,在所有转基因品系上功都有所提高。
图15是断开单纤维所需粒的克数与纤维细度的分散曲线。该曲线表示这些参数在所有数据点上具有68%的相互依赖性。预计这两种参数都会随着纤维壁的增厚而有所提高。
图是断开单纤维所需粒的克数与每粒种子的纤维重量的分散曲线。这些参数以61%的水平相互依赖,它类似于纤维细度的依赖性(参见图15)。这支持了这样的假说,即每粒种子纤维重量的增加在很大程度上是由于纤维壁厚度的增加。因为可以提高每粒种子的纤维重量的其他两种参数(增加了的纤维数量和增加了的纤维长度)预计不可能提高断裂所需的克数。
图17是断开单纤维所需功与纤维细度的分散曲线。这两种参数以48%的水平相互依赖。对于该复合参数而言,与断开所需克数和拉伸各自的相互依赖水平相比(参见图19),这种中等水平的依赖性是合理的。
图18是是断开单纤维所需要的功与每粒种子的纤维重量的分散曲线。这两种参数以39%的水平相互依赖,它类似于纤维细度的依赖性(参见图17)。正如在图16中所述,这一结果支持了这样的假说,即每粒种子纤维重量的增加在很大程度上是由于纤维壁厚度的增加。
图19是断裂拉伸量与纤维细度的分散曲线。该曲线表示这两种参数不相互依赖。因此,预计SPS的超量表达能通过独立于纤维壁厚度的机制提高拉伸量,这一结论与理论吻合。
图20是生长在人工气候室中的亲本C312的光合速度与内部二氧化碳浓度关系的四条重叠的分散曲线。空心符号是生长在30/15℃下的两种植物的,而实心符号是生长30/28℃下的两种植物的。所有植物都在30℃下测定。该曲线表示对于亲本C312来说,以前的寒冷的夜晚会抑制在温暖的白天的光合速度。
图21是生长在人工气候室中的转基因品系13-3a-1的光合速度与内部二氧化碳浓度关系的四条重叠的分散曲线。空心符号是生长在30/15℃下的两种植物的,而实心符号是生长30/28℃下的两种植物的。所有植物都在30℃下测定。该曲线表示对于该转基因品系来说,先前的低温对随后的温暖的白天的光合速度没有影响。
图22是生长在人工气候室中的转基因品系225-17a的光合速度与内部二氧化碳浓度关系的四条重叠的分散曲线。空心符号是生长在30/15℃下的两种植物的,而实心符号是生长30/28℃下的两种植物的。所有植物都在30℃下测定。该曲线表示对于该转基因品系来说,先前的低温对随后的温暖的白天的光合速度没有影响。
发明详述
本发明涉及一种控制植物中纤维素合成以便优化来自该植物的产品,尤其是纤维的产量水平和品质的方法。
术语“纤维”通常被用于统一植物细胞类型的不同的组。所述细胞的共同特征是具有拉长了的性状以及在厚的细胞壁中含有丰富的纤维素,所述细胞壁经常(但不是总是)被描述为次级细胞壁。所述细胞壁可以是或不是木质素化的,并且所述细胞原生质体在成熟时可以是或者不是活的。所述纤维具有很多工业用途,例如,用于木材和生产的木制品、纸张、植物、包装材料、绳索、刷子和扫帚、填充和添塞材料、嵌缝、加固其他材料以及生产纤维素衍生物。在某些行业中,术语“纤维”通常包括厚壁导管细胞,如维管和管胞以及许多单纤维细胞的纤维状聚合物。在本文中,术语“纤维”以其最广泛的含义使用,例如,包括(a)木质部的后壁导管和非导管细胞;(b)来自木质部以外的纤维,包括来自韧皮部、表皮、地下组织和表皮的纤维;和(c)来自茎、叶、根、种子和花或花序(如用于制造刷子和扫帚的高粱花序)的纤维。除了来自树木的木材、棉花和牧草作物的之外,本发明适用于所有纤维,包括,但不排除农业残余物,如玉米、甘蔗、和水稻茎干,这些残余物可用于制造纸浆,还包括亚麻、大麻、苎麻黄麻、洋麻、木棉、椰子皮纤维、竹子、spanish moss、麻蕉、和菠萝麻。
在一种优选实施方案中,本发明提供了一种转基因棉花植物,其中,所述转基因棉花植物相对非转基因棉花植物而言具有较高水平的蔗糖磷酸酯合成酶。表1表示来自未转化过的C312植物和四种转化过的植物品系的SPS活性水平。所有转化过的植物品系在叶片和纤维中的SPS活性都表现出明显的提高。
蔗糖磷酸酯合成酶在植物细胞内的碳代谢流中起着关键作用。业已从多种植物中分离了编码蔗糖磷酸酯合成酶的基因并进行了测序。[菠菜:Salvucci等,植物生理学102:529-536,1993;Sonnewald等,植物,189(2):1740181,1993;水稻:Valdez-Alarcon等,基因,170(2):217-222,1996;Craterostigma plantaqineum:Ingram等,植物生理学,115(1):113-121,1997;蚕豆:Heim等,基因,178(1-2):201-203,1996;马铃薯:WMBL保藏号X73477;温州蜜柑:Akra等,分子基因遗传学252:346-351,1996;甘蔗:Sugiharto等,植物细胞生理学38:961-965,1997;甜菜:Hesse等,分子基因遗传学247(4):515-520,1995;玉米Worrell等,植物细胞3:1121-1130,1991;拟南芥:Bevan等,NCBI保藏号AL049487;集胞藻:Kaneko等,DNA研究2(4):153-166,1995;Kaneko等,DNA研究3(3):109-136,1996;和未知生物:Van Assche等,US5665892-A,以上文献被收作本丈参考]。在图3中提供了来自高等植物的若干种现有SPS基因序列的比较。在图4中提供了集胞藻SPS(Kaneko等,DNA研究2(4):153-166,1996,收作本文参考)与菠菜SPS的比较;来自兰细菌的这种蛋白与菠菜SPS以及所有高等植物蛋白具有很高的同源性。优选的蔗糖磷酸酯合成酶包括从菠菜、拟南芥属甜菜、菜豆、柠檬、玉米、苔藓、马铃薯、水稻、甘蔗和集胞藻属中分离的基因。最优选的蔗糖磷酸酯合成酶基因是菠菜蔗糖磷酸酯合成酶。
除了蔗糖磷酸酯合成酶的已知序列之外,所述已知序列的修饰形式也属于本发明的范围。所述序列的变化包括可以对蔗糖磷酸酯合成酶的已知序列进行的取代、插入和缺失。对所有现有的序列进行比较发现了哪些氨基酸是高度保守的以及哪些氨基酸是可变的。利用这些信息,有可能选择仍然能产生有功能的蛋白的变异。
可以通过披露于以下文献中的方法根据由果糖-6-P和UDP-葡萄糖形成蔗糖-6-P(+蔗糖)通过比色方法测定(蔗糖磷酸酯合成酶的最大活性:Copeland,蔗糖代谢酶,植物生物化学方法,3:73-83,1990,该文献被收作本文参考)。在液氮中将冷冻的叶片或纤维组织粉碎,并在50mM HEPES(pH7.4)、10mM氯化镁、1mMEDTA、1mMEGTA、10%甘油和0.1%Triton-X100中研磨。在每一个SPS测定中使用每一种上清液的28微升的样品,并对每一种提取物进行三次测试。70微升的测试混合物中含有50mM HEPES(pH7.4)、10mMUDPG、6mM果糖-6-P、20mM葡萄糖-6-P(SPS激活剂)、10mM氯化镁、1mMEDTA、0.40mMEGTA、4.0%甘油和0.04%Triton-X100。该测定在32-34℃下进行10分钟(在最大活性平台上),然后添加70微升1N氢氧化钠终止测定。
通过将试管浸在煮沸的水浴中10分钟破坏未反应的己糖磷酸酯。在冷却到室温之后,添加250微升溶解在乙醇中的01%间苯二酚和750微升浓盐酸,然后在80℃下培养8分钟。将试管迅速冷冻到温室,用分光光度计测定A520nm,参照蔗糖标准曲线确定植物提取物中的蔗糖含量。对每一种提取物进行三次重复的对照测定,以便将每一种提取物中蔗糖的可能的不同的内在含量进行归一化。对于对照来说,在添加植物提取物之前将氢氧化钠添加到测试试管中;然后对该试管做与上文所述相同的处理,所不同的是通过氢氧化钠终止测试的步骤已经完成。还要通过Bradford蛋白测定方法分析植物提取物中的蛋白含量,并通过其吸收值分析叶片提取物的叶绿素含量,以便对不同样品之间的SPS活性进行比较。另外,还可以根据尿苷5’-二磷酸的释放通过分光光度计测定蔗糖磷酸酯合成酶的活性,所述释放是通过同样披露于以下文献中的丙酮酸激酶偶联酶反应检测的:*Kaneko等,DNA研究2(4):153-166,1996,收作本文参考。
为了在植物中表达蔗糖磷酸酯合成酶,通过用能表达蔗糖磷酸酯合成酶的嵌合DNA结构转化植物生产携带编码蔗糖磷酸酯合成酶基因的转基因植物。
为了表达来自所述嵌合DNA的蔗糖磷酸酯合成酶基因,该结构应当包括植物专一性启动子。该启动子应当确保外源基因在植物中表达。可以对启动子进行选择,以便所述表达只发生在特定组织中,发生在植物发育的特定时间点上或者发生在由外部因素所决定的时间点上。所述启动子可以与所述植物同源或异源。例如,合适的启动子包括披露于US5034322(收作本文参考)中的RUBISCO小亚基启动子、纤维专一性启动子、花椰菜花叶病毒的35S RNA启动子,披露于US5106739(收作本文参考)中的强化35S启动子,披露US5378619(收作本文参考)中的复式S35启动子、来自的玄参花叶病毒的FMV启动子,披露于US5466792(收作本文参考)中的RI T-DNA启动子,披露于US5428147(收作本文参考)中的章鱼氨酸T-DNA启动子,醇脱氢酶1启动子(Callis等,基因发育1(10):1183-1200,1987,(收作本文参考),马铃薯蛋白启动子B33(Rocha-Sosa等,EMBO杂志,8:23-29,1989,收作本文参考),E8启动子(Deikman等,,EMBO杂志,7(11):3315-3320,1988,收作本文参考),β-conglycin启动子(Tierney等,植物,172:356-363,1987,收作本文参考),酸性壳多糖酶启动子(Samac等,植物生理学,93:907-914,1990,收作本文参考),披露于US5491288(收作本丈参考)中的拟南芥属组蛋白H4启动子,或披露于US5290924(收作本文参考)中的用于在单子叶植物中表达基因的重组启动子。
优选的启动子包括RUBISCO小亚基启动子、35S启动子、纤维强化启动子、维管细胞强化启动子、干细胞强化启动子、或种子强化启动子。所述启动子能确保以组织专一性或组织增强形式表达,但也可以在其他细胞类型中表达。例如,它可以确保在光合作用活性组织中强化表达(RUBISCO(Worrell等,植物细胞,3:1121-1130,1991,收作本文参考)或其他叶肉细胞专一性启动子(Datta等,理论应用遗传学,97:20-30,1998,收作本文参考)或纤维(棉纤维、木质部纤维、或外木质部纤维专一性或强化启动子))。可以使用能确保仅在特定器官中表达的其他启动子,所述器官如叶片、根、块茎、种子、茎、花或特殊细胞类型,如薄壁细胞、表皮或维管细胞。组织专一性启动子的一个例子是RB7启动子,它是根专一性的(US5459252,收作本文参考)。所述启动子可以单独使用或组合使用,以便优化在最理想的一组组织或器官中的超量表达。
优选的棉纤维强化启动子包括以下棉纤维表达基因的启动子:E6(John等,植物分子生物学,30:297-306,1996和John等,美国科学院院报,53:12768-12773,1996,收作本文参考),H6(John等,植物生理学,108:669-676,1995,收作本文参考),FbL2A(Rinehart等,植物生理学,112:1331-1341,1996和John等,美国科学院院报,93:12768-12773,1996,收作本文参考),rac(Delmer等,分子基因遗传学,248:43-51,1995,收作本文参考);Cela(Pear等,美国科学院院报,63:12637-16642,1996,收作本文参考);CAP(Kawai等,植物细胞生理学39:1380-1383,1998);ACP(Song等,生物化学生物物理学学报,1351:305-312,1997);和LTP(Ma等,生物化学生物物理学学报,1344:111-114,1997)。
能增强在维管组织中表达的优选启动子包括CAD2启动子(Samaj等,植物,204:437-443,1998,收作本文参考),Pt4C11启动子(Hu等,美国科学院院报,95:5407-5412,1998,收作本文参考),C4H启动子(Meyer等,美国科学院院报,95:6619-6623,1998,收作本文参考),PtX3H6和PtX14A9启动子(Loopstra等,植物分子生物学,27:277-291,1995收作本文参考),RolC启动子(Graham,植物分子生物学,33:729-735,1997,收作本文参考),Hvhsp17启动子(Raho等,实验植物学杂志,47:1587-1594,1996,收作本文参考),和COMT启动子(Capellades等,植物分子生物学,31:307-322,1996,收作本文参考)。
能增强在茎组织中表达的优选启动子包括木髓启动子(Datta,理论应用遗传学,97:20-30,1998和Ohta等,分子基因遗传学,225:369-378,1991,收作本文参考)和阴离子过氧化物酶启动子(Klotz等,植物分子生物学,36:509-520,1998,收作本文参考)。能在韧皮部、皮层和软木组织中增强表达但不能在木质部或木髓中增强表达的优选启动子包括Psam-1启动子(Mijnsbrugge等,植物和细胞生理学,37:1108-1115,1996,收作本文参考)。
能增强在种子中表达的优选启动子包括phas启动子(Geest等,植物分子生物学32:579-588,1996);GluB-1启动子(Takaiwa等,植物分子生物学30:1207-1221,1996);γ-玉米醇溶蛋白启动子(Torrent等,植物分子生物学34:139-149,1997);和油蛋白启动子(Sarmiento等,植物杂志,11:783-796,1997)。
还可以使用包括能产生组织强化表达的特殊核苷酸片段的截短的和/或合成的启动子,例如,鉴定能产生木质部强化表达的较大启动子上的调控因子(Seguin等,植物分子生物学,35:281-291,1997;Dorres-Schumann等,植物杂志,9:283-296,1996;和Leyva等,植物细胞,4:263-271,1992,收作本文参考)。
在本发明的一种实施方案中,所述嵌合DNA结构被稳定地整合到棉花植物的基因组上。当植物是通过农杆菌介导的转化进行转化的时,Ti质粒的一部分整合到植物基因组上,并稳定地传给后代植物细胞。
现有多种转化植物细胞的方法。优选方法包括电穿孔、农杆菌属介导的转化、粒子轰击基因转化、化学介导的转化或显微注射。
上述载体可以利用微量移液管直接显微镜注射到植物细胞中,以机械方式转移重组DNA(Crossway,分子基因遗传学,202:179-185,1985,收作本文参考)。还可用聚乙二醇将遗传材料转入植物细胞(Krens等,自然,296:72-74,1982,收作本文参考)。
用能提高纤维和种子产量以及纤维品质的基因转化植物细胞的另一种方法是宿主细胞进行粒子轰击(又被称为粒子轰击转化)。这一目的可以通过若干种方式实现。第一种方式涉及推进细胞上的惰性或生物活性粒子。这种技术披露于Sanford等所拥有的US494505,US5036006和US5100792中,以上专利被收作本文参考。一般,该方法包括在能有效穿透细胞外表面并整合到细胞内部的条件下推进细胞上的惰性或生物活性粒子。当利用惰性粒子时,可以通过用含有异源DNA的载体对所述粒子进行包衣而将该载体导入细胞。或者,可以用所述载体包围靶细胞,以便由粒子将载体带入细胞。还可以将生物学活性粒子(例如,含有载体和异源DNA的干燥的细菌细胞)推进到植物细胞中。
导入的另一种方法是让原生质体与其他实体,如小细胞、细胞、溶酶体或其他可融合的脂表面体融合(Fraley等,美国科学院院报,79:1859-63,,1992,收作本丈参考)。
还可以通过电穿孔将DNA分子导入植物细胞(Fromm等,美国科学院院报,82:5824,1985,收作本文参考)。在该技术中,在有含有表达框的质粒的条件下对植物原生质体进行电穿孔。高场强电脉冲可逆地透化生物膜以便导入质粒。电穿孔的植物原生质体重新形成细胞壁、分裂、并再生。
将DNA分子导入植物细胞的另一种方法是用事先用基因转化过的根癌农杆菌或毛根农杆菌感染植物细胞。在本领域技术人员所公知的合适条件下,让转化过的植物细胞生长形成幼苗或根,并进一步发育成植株。一般,这一过程包括用细菌悬浮液接种植物组织,并在25-28℃下在不含抗生素的再生培养基上培养该组织48-72小时。
农杆菌属是革兰氏阴性科根瘤菌科的一个代表性的属。其种决定着冠瘿病(根癌农杆菌)和毛根病(毛根农杆菌)的发病。冠瘿瘤和毛根中的植物细胞被诱导产生被称为冠瘿碱的氨基酸衍生物,这一过程仅仅是由所述细菌催化的。决定冠瘿碱表达的细菌基因是嵌合表达框的控制因子的方便来源。另外,测定冠瘿碱的存在可用于鉴定转化过的组织。
通过根癌农杆菌的Ti质粒或毛根农杆菌Ri质粒可以将异源遗传学导入合适的植物细胞。Ti或Ri质粒是通过农杆菌属感染传递到植物细胞中的,并且稳定整合到植物基因组上(Schell,科学,237,1176-83,1987,收作本文参考)。在转化之后,可以收获完整的转化植物。如果直接产生了转化过的种子的话,可以通过在筛选培养基上发芽并长成植株进行筛选(Glouzh等,植物杂志16:735-743,1998,收作本文参考)。如果直接产生了转化过的花粉,可将这种花粉用于活体授粉,然后筛选转化过的种子(Touraev等,植物杂志,12:949-956,1997,收作本文参考)。如果分生组织是转化过的,可以让分生组织在培养基中长成植株,然后转移到土壤中(Gould,J等,植物细胞报导10:12-16,1991,收作本文参考)。
如果转化的是原生质体或外殖体,可以再生植株。由培养的原生质体再生的植株披露于被收作本文参考的以下文献中:Evans等,植物细胞培养手册,第1卷,纽约,纽约:MacMillan出版公司,1983;和Vasil著,植物细胞培养和体细胞遗传学,Orlando:学术出版社,第1卷,1984,和第3卷,1986。用于再生的方法根据植物种的不同而变化,但一般首先要提供转化过的原生质体的悬浮液或含有转化过的外殖体的培养皿。产生愈伤组织,并可以从愈伤组织诱导长芽,然后长根。另外,可以在愈伤组织上诱导胚胎的形成。这些胚胎向天然胚胎一样发芽形成植株。培养基通常含有各种氨基酸和激素,如生长素和细胞激动素。尤其是对诸如玉米和苜蓿这样的物种来说向培养基中添加谷氨酸和脯氨酸也是有利的。有效的再生取决于培养基、基因型和培养基的历史。如果控制上述三种变量,再生通常就是可以再现的和可以重复的。
众所周知的是,实际上所有植物都可以从培养的细胞或组织再生,这些植物包括,但不限于甘蔗、甜菜、棉花、树木、牧草作物和产纤维作物。产种子的作物也是可以再生的,包括,但不限于玉米、水稻、小麦、大豆、油菜、向日葵和花生。
在表达框稳定地整合到转基因植物中之后,它就可以通过有性杂交的方式转移给其他植物。根据要杂交的物种,可以使用多种标准育种技术中的任一种。
一旦生产出了这样类型的转基因植物,就可以按照常规方法栽培这种植物本身,在存在编码蔗糖磷酸酯合成酶的基因的条件下,会导致提高了的种子产量和/或提高了的纤维产量和/或提高了的纤维品质。或者,从所述转基因植物上收获转基因种子。然后可以将这些种子播种到土壤中,并用常规方法栽培,以便生产转基因植物。
本发明还提供了由具有增强了的蔗糖磷酸酯合成酶的合成的转基因植物所生产的种子。
在另一种实施方案中,本发明提供了一种通过将嵌合DNA结构导入棉花植物提高棉花植物产量的方法,所述DNA结构能改变蔗糖磷酸酯合成酶活性,达到足以提高棉花植物产量的程度。可将嵌合基因导入植物细胞或组织中。转化过的细胞通常是利用选择标记筛选的。然后利用转化过的细胞产生转化过的植物(Fraley等,美国科学院院报79:1859-1863,1982,收作本文参考)。
优选的植物是棉花植物。所述转化过的植物可以具有提高了的棉籽或棉纤维产量。
本发明还提供了一种通过将嵌合DNA结构导入棉花植物提高由棉花植物所产生的棉纤维的品质的方法,该DNA结构能改变蔗糖磷酸酯合成酶活性,达到足以提高由该棉花植物所产生的棉纤维的品质的程度。
蔗糖磷酸酯合成酶的含量可以通过能提高该基因表达水平的表达因素得到加强。所述因素可作用于控制表达的调控位点,该位点通常位于靠近蔗糖磷酸酯合成酶基因处,或用于控制位于靠近嵌合结构中所述基因的异源调控位点。另外,蔗糖磷酸酯合成酶的含量可以通过导入能直接表达蔗糖磷酸酯合成酶的嵌合DNA结构而提高。
一般,本发明可用于通过将嵌合DNA结构导入植物改变纤维素与整株植物的干重的比例或纤维素与植物成分的干重的比例,所述DNA结构能改变蔗糖磷酸酯合成酶活性,达到足以改变纤维素与全植株的干重的比例或纤维素与植物成分的比例的程度。纤维素的改变体现在与植物总重量的关系上或与植物的分离的部分的关系上,这些部分包括,但不是排他性的,淀粉、总细胞壁、纤维细胞壁、特定器官,如茎,或细胞壁成分,如果胶、半纤维素、蛋白、提取物和木质素。当单独考虑或以任何组合形式考虑所述分离部分时,可以发现纤维素与植物分离部分的比例的改变。
在本发明中所声称的品质的改变是指与缺乏转基因的植物相比至少有10%的变化。例如,细胞壁中纤维素的比例可以从20%改变到18%或更低或22%或更高。与亲本水平相比的这种变化可用于于所有细胞壁或植物的任何细胞壁部分。
在一种优选实施方案中,可以提高纤维素的干重量,以便它与其他干重成分的比例超过40%。这种超过40%的提高可应用于木材、纤维、和其他富含纤维素的细胞壁,如厚角细胞和增厚了的木质部薄壁细胞。
为了实现某些改变,可以通过能减弱所述基因表达水平的表达因素降低蔗糖磷酸酯合成酶的含量。所述因素可作用于控制表达的调控位点,所述位点通常位于靠近蔗糖磷酸酯合成酶基因处或作用于位点靠近嵌合结构中的该基因的异源调控位点。另外,在反义技术中,蔗糖磷酸酯合成酶的含量可以通过导入含有蔗糖磷酸酯合成酶的互补cDNA的嵌合DNA结构而降低(Arndt等,基因组,40:785-797,1997,收作本文参考)。另外,通过同源性依赖基因沉默(Wassenegger等,植物分子生物学37:349-362,1998,收作本文参考),病毒诱导的基因沉默(Baulcombe,植物生物学当代观点2:109-113,1999,收作本文参考),嵌合RNA/DNA寡核苷酸(Zhu等,美国科学院院报15:8768-8773,1999,收作本丈参考),或同源重组(Shalev等,美国科学院院报96:7398-7402,1999,收作本文参考)诱导SPS活性降低。
在另一种实施方案中,本发明提供了一种通过将嵌合DNA结构导入植物提高光合效率对夜晚低温的耐受性的方法,所述DNA结构能改变蔗糖磷酸酯合成酶活性,达到足以提高光合效率对夜晚低温耐受性的程度。
本发明可用于通过将嵌合DNA结构导入植物调控植物细胞壁的厚度,所述DNA结构能改变蔗糖磷酸酯合成酶活性。具体地讲,所述方法可用于提高来自任何产纤维植物的可收获的纤维产量。
在一种优选实施方案中,所述植物是产纤维植物。更优选的产纤维植物是甘蔗、甜菜、树木、牧草作物、产纤维植物、和产种子植物。
在另一种实施方案中,本发明可用于提高植物纤维的可收获产量。本发明还可用于改变从植物中分离的纤维的品质。蔗糖磷酸酯合成酶的改变可以改变纤维强度、纤维长度、或单位长度的重量。所述改变还可以是提高或降低强度、长度或单位长度的重量。
本发明可用于通过将嵌合DNA结构导入植物提高从产种子植物上收获的种子产量,所述DNA结构能提高蔗糖磷酸酯合成酶活性。
本发明的方法可以广泛应用,并可用于多种植物,包括棉花、树木、牧草作物、甜菜、亚麻、大麻、黄麻和其他产纤维植物。还可将其用于产种子植物,包括棉花、亚麻、小麦、水稻、玉米、大豆、油菜、向日葵、红花、花生、棕榈和其他产种子植物。在下面的实施例中将对本发明作进一步说明。
实施例
实施例1-材料和方法
所披露的大多数植物是生长在杜克大学人工气候室的生长箱中:360ppm(正常)二氧化碳;30℃/15-19℃昼/夜周期;白天14小时/夜晚10小时;光照1200mM m-2s-1(金属卤化物);每天用1/2强度的Hoagland’s溶液浇水2次;在装有沙砾和沙子混合物4加仑花盆中盆栽。在生长大约4个月之后将30/19℃改变成30/15℃,这一时间点相当于C312和所有转基因品系的首批棉铃成熟的中点时间。这种温度条件随后被简单地称之为30-15℃。强调该生长箱是因为其温度和二氧化碳条件代表了棉花作物在大田中有可能遇到的条件,例如,但不排除其他可能,在德克萨斯南部高原上所遇到的条件。
其他植物生长在所述杜克大学人工气候室的3个其他的生长箱中,所不同的是进行以下改变:(a)360ppm二氧化碳,30℃/28℃昼/夜周期;(b)700ppm(提高了的)二氧化碳,30℃/15-19℃昼/夜周期;和(c)700ppm二氧化碳,30℃/28℃昼/夜周期。
其他植物在德克萨斯技术大学温室中生长:正常二氧化碳和光照;大约32℃/22℃昼/夜周期;2加仑花盆;每天浇水2-3次;混在土壤中的缓式和每周使用1次的可溶性肥料。
从每一个植株上收获所有开花的棉铃,用这些棉铃评估种子和纤维参数。通过人工剥离除去种子上的皮棉纤维。棉花种子被皮棉纤维(用于纺织的长纤维)和短纤维(用于各种工业用途的短纤维)所覆盖。通过称重测定来自每一个植株的(皮棉)纤维重量和短纤维种子重量。在下丈中,‘纤维’是指皮棉纤维,在必要时特指短纤维。通过计数测定每个植株的种子数量(发育不全的“微粒”的种子和纤维不包括在内)。将纤维送到Cottoa Incorproated,Raleigh,NC进行HVI、AFIS和Mantis纤维品质分析。然后将来自30/15℃生长箱的种子进行酸去绒、风干并称重。来自该生长箱的每粒种子的短纤维重量是通过扣除有绒和去绒种子的重量之后测定的。
对于测定茎干重量的植物来说,去掉所有未开放的棉铃和叶片及叶柄。将地上部分的茎干烘干并称重。
所使用的植物品系是C312野生型(未转化过的亲本)和四种转基因品系。已知每一种转基因植物品系代表不同的转化事件,被命名为13-3a、225-17a、40-4b和40-6a。T0、T1或T2分别表示原始转化体和一代和二代转化体。所测试的所有转基因植物都是卡那霉素抗性的,这是通过在含有卡那霉素的琼脂上能形成发芽幼苗的侧根而确定的。在卡那霉素上发芽的种子的比例被表达为抗性/敏感比例(表1)。在7-14天之后测定比例,以便包括大多数缓慢发芽的种子。
生长在人工气候室中用于产生每一种参数的平均数据的单个植株的数量(40-6a-4除外)表示为人工气候室植株(n)(表2)。品系40-6a-4尽管总体上与其他品系表现一致,但还是将其从纤维品质平均数中去掉,因为它仅代表生长在30-15℃,360ppm二氧化碳生长箱中的一个植株。将来自品系40-4b的2个T2的品系的值加以平均,因为T1#1和T1#4是能产生类似的T2后代的类似的姊妹株(除了分离比例不同之外)。
通过对叶片中外源SPS的mRNA进行北方分析测定叶片和纤维RNA含量,将其归类成阳性或阴性(表1)。将可提取的SPS活性(蔗糖产量)标准化为叶片活性的微摩尔蔗糖/毫克叶绿素/小时或标准化为纤维活性的微摩尔/毫克蛋白/小时(表1)。
每个植株的棉铃数是每一种植株上没有脱落的棉铃的数量。
每粒种子的去绒种子重量(g)和每粒种子的(皮棉)纤维重量(g)(表2)是来自每一个植株在该实验结束时的所有开放的棉铃的数量。在30-28℃下,所有棉铃都开放,但在30-15℃下,在实验结束时某些未开放的棉铃留在各自的植株上。每一种数据点表示129-487个种子产生24.5-48.5克皮棉纤维。
通过自动化HVI和AFIS测试测定的纤维束特性归纳在表3和4中。纤维细度(通过HVI)测定是均匀度指标,它取决于纤维成熟度(由次级壁纤维素含量所决定的壁厚)和纤维直径。
纤维束强度(通过HVI测定)是以cN/特为单位表示的。这是纤维束的比强度,其中,单纤维的细度(特)是由所述细度值计算的。
纤维细度(通过AFIS测定)是以毫特为单位表示的。它表示1千米纤维的重量毫克数。重量为1毫克的1千米长的纤维=1毫特。
纤维成熟度比例(通过AFIS测定)表达的是细胞壁增厚的程度(取决于次级细胞壁纤维素沉积)它是圆整度比例为0.5(或更高)的纤维除以圆整度比例为0.25(或更低)的纤维量的比例(壁较厚的纤维在干燥时收缩的可能性较低更有可能保持圆形)。成熟度比例越高,纤维就越成熟,并且该纤维在用于染色时越好。
成熟纤维含量(IFC%,通过AFIS测定)是具有低于0.25成熟度的纤维的百分比。IFC%越低,这种纤维就越适合染色。
有若干种不同的单位被用作纤维长度的指标。表3示出了本文所披露的三种指标的值。上半部平均值(UHM,通过HVI测定)是最长的一半纤维的平均长度(重量偏差)。纤维均匀度指数(UI,通过HIV测定)表示平均值(平均长度)与上半部平均长度的比例。它是纤维长度在该群体内分散的指标;如果所有纤维的长度相同,UI=100%。短纤维含量(IFC%,通过HVI测定)是以重量为基础的长度低于1/2”的纤维的百分比。HVI被认为只是通过遗传学测定来测定短纤维含量,因为该测定不构成额外的潜在纤维断裂应力。
在本文中要讨论的其他纤维长度指标如下。重量基础长度(L(w)[in],通过AFIS测定)是以重量为基础计算的纤维长度。以数量为基础的长度(L(n)[in],通过AFIS测定)是用数量计算的纤维平均长度。长度“L%(n)”[in](通过AFIS测定)是5%跨越长度,或5%纤维所跨越的长度,此时这些纤维是平行的并且是随机分布的。长度“L%(n)”[in](通过AFIS测定)是2.5%跨越长度,或2.5%纤维所跨越的长度,此时这些纤维是平行的并且是随机分布的。“UQL(w)”[in](通过AFIS测定)是上1/4纤维长度的重量,或超过纤维25%长度的重量。最后,“SFC(c)”[in]和“SFC(w)”[in](通过AFIS测定)是分别以数量和重量为基础的长度低于0.50英寸的纤维的百分比。与HVI相反,AFIS在进行上述测定之前要抽打纤维,这有可能导致纤维断裂。因此,AFISSFC值是经过正常加工以后的纤维特征的良好的指标。
在表5中归纳了通过Mantis测定所得到的单纤维强度和拉伸参数。Tb[克]是断开单纤维所需力的克数。拉伸量[%]是在断裂之前纤维相对原始长度的拉伸百分比。功[μJ]是Tb和拉伸量的总和,表示断开单纤维所耗费的功。
在下面的实施例中包括特定实验的详细方法。
实施例2-证实具有较高SPS活性的转基因植物提高了的纤维和种子产量的结果的概述
通过北方分析证实,具有受组成型启动子控制的菠菜SPS的转基因棉花植物在叶片和纤维中表现出外源基因表达。在T1/T2代,与亲本C312相比,在叶片和纤维中分别表现出SPS酶活性平均提高了3.3倍和2.3倍(表1)。在该表以及下面所有的表中,表示优于亲本C312的转基因植物的特征的值都用黑体表示。
表1
在转基因植物中鉴定菠菜SPS基因表达和总SPS活性
植物品系 | 分离比例 | 叶片RNA | 纤维RNA | 叶片SPS活性(纤维素) | 正规化叶片SPS活性 | 纤维SPS活性(蛋白) | 正规化纤维SPS活性 |
C312-wt | na | - | - | 23.53a | 1.0 | 39.91 | 1.0 |
31.30b | 1.0 | ||||||
13-3a | |||||||
T0 | + | 119.2 | 5.1 | ||||
T1 | 22∶6 | ||||||
T1#1@T2 | 66∶0 | + | 127.2 | 4.0 | 103.39 | 2.6 | |
225-17a | |||||||
T0 | + | 118.5 | 5.0 | ||||
T1 | 25∶12 | + | 121.8 | 3.9 | 93.71 | 2.4 | |
40-4b | |||||||
T0 | + | 107.3 | 4.6 | ||||
T1 | 11∶4 | ||||||
T1#1@T2 | 51∶16 | 60.3 | 1.9 | 91.67 | 2.3 | ||
T1#4@T2 | 10∶0 | + | 66.4 | 2.1 | 76.00 | 1.9 | |
40-6a | |||||||
T0 | + | 89.3 | 3.8 | ||||
T1 | 6∶5 | ||||||
T1#4@T2 | 9∶2 | 57.6 | 1.8 | 74.12 | 1.9 | ||
T1/T2c转基因平均值 | 103.9 | 3.3 | 85.4 | 2.3 |
a所测定的用于T0比较的值。
b所测定的T1和T2比较的值。
C品系40-6a的排除值,并利用品系40-4b的复合平均值,与用于纤维品质数据分析的方法平行处理。
在30/15℃,360ppm二氧化碳人工气候室中生长的头9周,测定植株高度和叶片数量,转基因品系与亲本C312相同生长。转基因植物的平均高度为0.90×亲本C312的高度值。转基因植物的平均叶片数是1.02C×亲本C312的叶片数。
在30/15℃,360ppm二氧化碳人工气候室中,被上调的SPS基因表达导致纤维和种子作物的产量成分的提高(表2)。
表2
SPS转基因植物的产量成分与亲本C312的比较(30/15℃和360ppm
二氧化碳)
植物品系 | 人工气候室植物(n) | 每株的棉铃数 | 正规化棉铃数 | 每粒种子的去绒种子重量(克) | 正规化每粒种子的种子重量(克) | 每粒种子的纤维重量(克) | 正规化每粒种子的纤维重量 |
C312-wt | 4 | 22.8 | 1.0 | 0.090 | 1.0 | 0.047 | 1.0 |
13-3a | |||||||
T1#1@T2 | 4 | 26.5 | 1.16 | 0.107 | 1.19 | 0.058 | 1.23 |
225-17a | |||||||
T1 | 4 | 26.0 | 1.14 | 0.110 | 1.22 | 0.063 | 1.34 |
40-4b | |||||||
T1#1@T2 | 5 | 28.2 | 1.24 | 0.100 | 1.11 | 0.057 | 1.21 |
40-6a | |||||||
T1#4@T2 | 1 | 28.0 | 1.23 | 0.105 | 1.17 | 0.054 | 1.15 |
T1/T2a转基因平均值 | 26.9 | 1.18 | 0.106 | 1.18 | 0.059 | 1.25 |
a平均值中去掉了品系40-6a,因为重复很少。
棉纤维和棉籽都是有价值的作物,皮棉纤维用于纺织和其他用途,而种子作为油料和种子粉的来源。另外,短纤维(又称为短绒)是作为化学纤维素的来源收获的,除此之外还有其他用途。观察到了每个植株棉铃数、每粒种子的种子重量、每粒种子的纤维重量和每粒种子的短纤维重量的增加。每个植株的棉铃数量表示生产具有所述纤维的总体能力。另外,种子和每粒种子纤维重量的增加会导致产量的提高。能超量表达SPS的转基因植物能同时获得两种类型作物的产量提高,种子产量主要是基于蛋白和油类的储存,而纤维产量基础纤维素的储存。因此,可以预测超量表达SPS的植物就作物产量而言在每英亩面积上可以为棉花生产者产生更多的收益。能超量表达SPS的C312植株也可以在将来的转化中使用,以便克服由于在遗传工程中使用这种老的栽培种所产生的任何潜在的产量下降。在其他作物上可以同时提高种子和纤维产量,并可以产生更坚固的茎干以便抗倒伏而又不会损害种子产量。
每个植株增加的棉铃数:
与亲本C312相比,在30/15℃,360ppm二氧化碳生长箱中测试的3个转基因品系以良好的重复性表现出每个植株的棉铃数增加14-24%,平均增加了18%。还在30/15℃,700ppm二氧化碳和30/28℃,700ppm二氧化碳生长箱中观察到了所有转基因品系的棉铃数的增加。
提高了的每粒种子的纤维重量:
与亲本C312相比,在30/15℃,360ppm二氧化碳生长箱中测试的3个转基因品系的每粒种子的纤维重量提高了21-34%,平均提高了25%(表2,图5)。这种效果在其他生长箱中并没有一致地出现。每粒种子的纤维重量是纤维数量、纤维长度和纤维壁厚度的综合指标。由于平均纤维细度(表示增加了的壁厚度)和其他相关因素在所有生长箱中的所有转基因品系都有所提高(参见下文),人们有理由认为,未测定的因素,如改变纤维数量可能影响在几乎稳定的温度条件或较高的二氧化碳条件下每粒种子的纤维重量。
在实验室产量分析中有时候要用的一种指标是皮棉%=(皮棉纤维重量)/(和种子和皮棉纤维总重量)。三个转基因品系都比亲本C312 31.14%的值提高了1.8-2.7%(转基因植物平均提高了2.1%)。这一值预示着转基因品系纤维产量的提高,因为种子重量也提高了(参见下文)。
提高了的每粒种子的种子重量:
与亲本C312相比,在在30/15℃,360ppm二氧化碳生长箱中测试的3个转基因品系每粒种子的去绒种子重量提高11/22%,平均提高了18%(表2,图6),仅对来自其他生长室中的有绒种子进行了称重,不过,比较来自在30/15℃,360ppm二氧化碳生长箱的有绒和去绒值可以发现,有绒种子值体现了种子产量的趋势。在生长在其他三个生长箱中的转基因品系上有绒种子表现出每粒种子种子重量的提高,唯一的例外是在在30/28℃,700ppm二氧化碳生长室中225-17a表现出每粒种子的种子重量与亲本C312相同。
在3个转基因品系上在30/15℃,360ppm二氧化碳生长箱中每粒种子的纤维重量与每粒种子的去绒种子重量的比例平均提高了9.0%(图7)。每粒种子的纤维重量与每粒种子的去绒种子重量的分散曲线表明,转基因植物通过这两种产量成分的提高而偏离亲本C312(图8)。不过,在SPS转基因植物上,纤维重量的提高比种子重量的提高有优势。
提高了的每粒种子的短纤维重量:
每粒种子的短纤维重量是通过从来自30/15℃,360ppm二氧化碳生长室的有绒种子的单位种子重量中扣除去绒种子的单位种子重量之后获得的(图9)。由两个转基因品系(225-17a和40-4b)表现出提高(与亲本C312相比平均提高了19%),而一个转基因品系(13-3a)表现出降低(与亲本C312相比平均降低了19%)。在去绒之前,品系13-3a的种子看上去颜色暗一些,这表明一开始种子上的短纤维就少于亲本C312或其他两个转基因品系种子上的短纤维。因此,转基因品系在短纤维数量的启动方面表现出某些差异,但一旦开始,超量表达的SPS会以类似于提高皮棉纤维的方式提高其产量。
实施例3-通过对总体样品进行自动化HVI和AFIS分析证实提高了的纤维品质的结果的概述
棉花的很多纺丝特性取决于它的作为整体样品的特性。HVI和AFIS是分析这些特性的自动化系统,能得到互补的信息。上述分析表明,由SPS转基因植物生产的纤维的品质参数作为整体移动到优质范围内。来自SPS转基因植物的纤维更长更结实并且更成熟-所有这些特征在棉花加工和纺织工业目前都是有价值的,因为可以生产出优质织物。即使是在德克萨斯南部高原上通常遇到的胁迫性的30-15-19℃温度周期下,来自SPS转基因植物的纤维的品质也类似于在加利弗尼亚传统种植的优良棉花的品质。因此,来自SPS转基因的棉纤维可以满足成品市场的扩大,并卖出好价钱。种植SPS转基因棉花的生产者还可以避免由于诸如细度低的不良品质而导致的价格下跌,这种不良品质可能会在德克萨斯南部高原上种植的传统棉花上出现。因此,SPS转基因棉花应当能够根据改善了的纤维品质而稳定或提高棉花生产者在每一亩上获得的收益。
在30/15℃,360ppm二氧化碳下的改进:
在表3中示出了来自生长在30/15℃,360ppm二氧化碳生长箱中的纤维的通过HVI和AFIS分析的重要的总体纤维品质参数。在表4中示出了转基因品系相对亲本C312提高的倍数。
表3
SPS转基因植物与亲本C312的纤维品质参数(30/15℃和360ppm二
氧化碳)
植物品系 | 人工气候室植物(n) | 纤维细度 | 纤维束强度(cN/特) | 纤维细度(毫特) | 纤维成熟度比例 | 成熟纤维含量(%) | 纤维长度(UHM)(in) | 纤维均匀度(UI,%) | 短纤维含量(%,通过HVI测定) |
C312-wt | 4 | 3.68 | 27.1 | 167 | 0.89 | 7.45 | 1.04 | 83.1 | 7.5 |
13-3a | |||||||||
T1#1@T2 | 4 | 4.55 | 28.8 | 170 | 0.92 | 6.85 | 1.15 | 88.9 | 5.9 |
225-17a | |||||||||
T1 | 4 | 5.12 | 31.0 | 189 | 0.99 | 4.35 | 1.14 | 87.9 | 2.9 |
40-4b | |||||||||
T1#1@T2 | 5 | 4.50 | 31.1 | 180 | 0.95 | 5.64 | 1.12 | 84.8 | 5.9 |
40-6a | |||||||||
T1#4@T2 | 1 | 5.30 | 29.6 | 177 | 0.96 | 5.20 | 1.08 | 86.1 | 11.3 |
T1/T2M转基因平均值 | 4.72 | 30.3 | 180 | 0.95 | 5.61 | 1.14 | 87.2 | 4.9 |
a平均值中去掉了品系40-6a,因为重复很少。
表4
SPS转基因植物的纤维品质参数的改变:(30/15℃和360ppm二氧化碳)所表示的值是相对被设定为1.0的C312-wt值正规化的或者是相对亲本
C312值的改变百分比
植物品系 | 人工气候室植物(n) | 正规化纤维细度 | 正规化纤维束强度(cN/特) | 正规化纤维细度(毫特) | 正规化纤维成熟性比例 | 未成熟纤维含量变化(%) | 正规化纤维长度(UHM) | 纤维均匀性变化(UI,%) | 短纤维含量变化(%通过HVI测定) |
C312-wt | 4 | 1.00 | 1.00 | 1.00 | 1.00 | 7.45% | 1.00 | 83.1% | 7.5% |
13-3a | |||||||||
T1#1@T2 | 4 | 1.23 | 1.06 | 1.02 | 1.03 | -0.60% | 1.11 | +5.8% | -1.6% |
225-17a | |||||||||
T1 | 4 | 1.39 | 1.14 | 1.13 | 1.11 | -3.10% | 1.09 | +4.8% | -4.6% |
40-4b | |||||||||
T1#1@T2 | 5 | 1.22 | 1.15 | 1.08 | 1.07 | -1.81% | 1.07 | +1.7% | -1.6% |
40-6a | |||||||||
T1#4@T2 | 1 | 1.44 | 1.09 | 1.08 | 1.08 | -2.25% | 1.04 | +3.0% | +3.8% |
T1/T2a转基因平均变化值 | 1.28 | 1.12 | 1.08 | 1.07 | -1.84% | 1.10 | +4.1% | -2.6% |
a平均值中去掉了品系40-6a,因为重复很少。
细度。三个转基因品系表现出平均提高了28%,达到平均细度为4.72(图10),细度取决于次级壁厚度和纤维直径。理想的是,细度提高的出现是由于次级壁厚度的提高,而不是由于纤维直径的增加。纤维直径是通过纤维细度和纤维成熟度比例(表3)之间的标准化关系估算的,并且发现在转基因品系上有很小的变化。亲本C312和转基因品系估计的纤维直径为16.5-17.0微米。另外,细度与每粒种子的纤维重量的曲线表现出59%的相互依赖性(图11),表明在SPS转基因植物的纤维中存在较厚的壁。有关纤维强度、成熟性比例和未成熟纤维含量的其他数据(参见下文)也表明了来自SPS转基因植物纤维的壁厚度的提高。有超过90%的棉纤维壁的厚度是由于在次级细胞壁中几乎纯的纤维素的沉积而造成的。因此,SPS的超量表达提高了棉纤维的纤维素含量。
纤维束强度。三个转基因品系表现出平均提高了12%,达到平均束强度为30.3cN/特。
纤维细度。三个转基因品系表现出平均提高了8%,达到平均细度为180。较高的纤维细度传统上是不理想的,因为它通常会造成较大的纤维直径。不过,由于SPS转基因植物的纤维具有大体上与亲本C312大体上相同的直径(见上文),细度的增加可能是由于纤维壁厚度的增加,导致了每单位长度更大的重量。因此,预计提高来自SPS转基因植物的纤维细度是一种中性的或有利的纤维品质因素。
纤维成熟性比例。三个转基因品系表现出平均提高了7%,达到平均成熟比例为0.95,这一比例落在“超过平均”范围内(0.95-1.00)。这一结果优于亲本C312,亲本C312的平均值为0.89,“成熟”范围内(0.85-0.95)。
未成熟纤维含量。三个转基因品系表现出平均降低了1.84%,达到平均为5.61%的未成熟纤维。转基因纤维好于亲本C312的纤维,亲本纤维含有平均为7.45的未成熟纤维。
纤维长度。三个转基因品系表现出在上半部分平均长度平均提高了10%,达到平均UHM为1.14英寸。这三个品系还具有更均匀的纤维长度,平均均匀性指数提高了4.1%,达到平均UI为84.2%。这三个品系还具有较少的短纤维,通过HVI测定的平均短纤维含量降低了2.6%,达到平均SFC%为49%。除了在表3和4中归纳的数据以外,其他AFIS参数支持在SPS转基因植物的纤维中纤维长度增加的结果。对于三个转基因品系的平均值来说,L(w)提高了7%,达到1.06英寸,L(n)提高了9%,达到0.96英寸,UQL(w)提高了6%,达到1.19英寸,L5%(n)[in]提高了6%,达到1.34英寸,L2.5%(n)提高了5%,达到1.46英寸。类似的,AFIS证实平均而言,三个转基因品系具有降低了的短纤维含量,SFC%(w)降低了1.0%-31%,而SFC%(n)降低了2.0%-10.6%(上述AFIS SFC%平均值去掉了来自品系40-4b的一个植株的值,因为它的值偏离平均值太多,会使平均值极大地偏离该品系中其他四个植株的值)。由于AFIS在进行测定之前要抽打纤维,这些降低了的SFC%值是在正常纤维加工过程中来自SPS转基因植物的纤维的改善了的均匀性的良好指标。
在不同环境条件下的改善:
在30/15℃,360ppm二氧化碳生长室中,与亲本C312相比大多数转基因品系的很多纤维品质参数得到提高,上述条件是测试的棉花的唯一的典型生长条件。不过,生长在其他人工气候室中的转基因植物的纤维品质也得到了保持或提高,其中温度在30/15℃至30/28℃之间变化,和/或二氧化碳在360ppm-700ppm之间变化。这一结果得到了转基因值的证实,并且得到了种植在其他三个生长室中的三个转基因品系的纤维品质数据与相对C312的值发生变化的证实,不包括30/15℃,360ppm生长室,该生长室单独归纳。SPS的超量表达能保持对纤维细度和平均纤维长度L(n)特别强的影响,与对UI和SFC的影响一致。
细度。4.65;是C312平均值的1.13倍。
纤维束强度。30cN/特;1.02倍。
纤维成熟度比例。0.92;1.03倍。
未成熟纤维含量。0.69%;降低了1.1%。
长度(n)。0.95英寸;1.08倍。
上部1/4长度。1.21英寸;1.03倍。
纤维均匀性指数。87.7%;提高了1.3%。
通过HIV测定的短纤维含量(w)。3.77%;降低了1%。通过AFIS测定的短纤维含量(w)。3.95%;降低了1.75%。
当30/15℃改变成30/28℃(360ppm二氧化碳)或360ppm二氧化碳改变成700ppm二氧化碳(30/15℃)时,每一个植物品系内的变化与纤维细度、UHM、UI、纤维束强度、SFC%、UQL、L(n)、IFC%和成熟度比例的品质参数的平均值相当。以上计算表明,在转基因品系中SPS的超量表达能促进纤维品质有几乎最大的提高,即使是在最受限制的30/15℃,360ppm二氧化碳条件下也是如此。相反,提高最低温度或二氧化碳含量能明显提高亲本C312的纤维细度、UHM、UI和纤维束强度。因此,SPS转基因植物的高纤维品质更独立于环境。
实施例4-通过Mantis单纤维测试分析的证实提高了的纤维品质的结果的概述
纺织工业非常看中具有较高单纤维强度的棉花纤维,因为这种纤维在加工期间较少断裂。因此,在加工过程中可以较高的值保持平均纤维长度,并能生产出具有较少缺陷的优质织物。提高单纤维强度是棉花行业的一个主要目标。
为了测试单纤维强度而进行的Mantis测试是用来自每个植物品系的至少4个植株的100个纤维(分成各5根纤维的2个独立的组进行的)。因此,表5中的数据是来自植物品系的至少一共400根纤维的平均值。
表5
SPS转基因植物与亲本C312单纤维强度的比较(30/15℃,360ppm
二氧化碳)
植物品系 | 纤维# | Tb(克) | 正规化Tb | TbS.D. | TbS.D.% | 拉伸量(%) | 拉伸量变化% | 功(μJ) | 功变化 | 功S.D. | 功S.D.% |
C312-wt | 400 | 5.30 | 1.00 | 2.45 | 46.2 | 15.05 | 13.21 | 1.00 | 8.98 | 68.0 | |
13-3a | |||||||||||
T1#1@T2 | 400 | 5.90 | 1.11 | 2.55 | 43.2 | 17.40 | +2.35 | 15.99 | 1.21 | 8.62 | 53.9 |
225-17a | |||||||||||
T1 | 400 | 7.18 | 1.35 | 2.85 | 39.7 | 16.67 | +1.62 | 18.09 | 1.37 | 9.55 | 52.8 |
40-4b | |||||||||||
T1#1#4@T2 | 500 | 6.60 | 1.24 | 2.71 | 41.1 | 16.89 | +1.84 | 17.22 | 1.30 | 9.21 | 53.5 |
转基因平均值 | 6.56 | 1.24 | 2.70 | 41.2 | 16.99 | +1.94 | 17.10 | 1.29 | 9.13 | 53.4 |
Tb:断开单纤维所需要力的克数
拉伸量%:在断裂之前单纤维的拉伸量,以原始长度的百分比表示功∶Tb和拉伸量的综合=断裂单纤维所消耗的功
XXS.D:有关值的标准误差
XXS.D%:实际值的百分比,由标准误差值表示
表5表示与亲本C312相比,在所有三个转基因品系中,单纤维强度得到了提高,表现在Tb、拉伸量和功都一致地得到改善上。在三个转基因品系中平均来说,Tb提高24%,达到6.56克(图12),拉伸量提高了1.94%,达到16.99%(图13),而功提高了29%,达到了17.10μJ(图14)。(与亲本C312相比转基因品系的HVI未表现出伸长量%的任何提高,因为基于纤维束HVI的测试仅仅反映的是该纤维束中最弱的纤维的拉伸量)。另外,标准误差是转基因单纤维强度值的较低的百分比(功平均低1.46%),表明单纤维强度具有改善了的均匀性(预计Mantis单纤维测试的结果具有高的标准误差)。
图15-19中的分散曲线表示来自30/15℃,360ppm二氧化碳生长箱的单纤维强度参数和细度或每粒种子的纤维重量之间的相关性。这些结果表明了Tb和功和纤维细度和每粒种子纤维重量之间的正相关性(图15-18)。相反,在拉伸量和纤维细度(图19)或每粒种子的纤维重量之间没有发现正相关性。测定系数表明,通过纤维细度和每粒种子纤维重量的提高确定了Tb和功提高了39-68%。这些正相关性主要是通过来自SPS转基因植物的独立的数据点的组确定的。该点在表6中标明,示出了分别考虑的每一个植物品系的测定系数(R2)。与转基因品系不同,亲本C312没有表现出显著的正的R2值。因此,与亲本C312相比,SPS的超量表达导致转基因纤维细度值的提高,这种提高与单纤维强度值的提高相关。
表6
来自对纤维细度和每粒种子的纤维重量作图的单一植物品系的单纤
维强度参数的线性回归曲线的测定系数(R2)
Y轴 | 功 | Tb | 拉伸量 | |||
X轴 | 纤维细度 | 每粒种子的纤维重量 | 纤维细度 | 每粒种子的纤维重量 | 纤维细度 | 每粒种子的纤维重量 |
植物品系 | ||||||
C312 | -0.10 | -0.10 | 0.16 | 0.15 | -0.29 | -0.29 |
13-3a | 0.50 | 0.06 | 0.37 | 0.00 | 0.56 | 0.30 |
225-17a | 0.40 | 0.67 | 0.95 | 0.99 | -0.57 | -0.31 |
40-4b | 0.34 | 0.83 | 0.83 | 0.54 | 0.10 | 0.83 |
纤维细度(在三个转基因品系中)和每粒种子的纤维重量(在二个转基因品系上)与Tb和功的显著的正相关性支持这样一种事实,即每粒种子纤维重量和纤维细度的提高是由于纤维壁中纤维数沉积的增加。由于纤维数量的增加而导致的每粒种子纤维重量的增加或由于纤维直径的增加而导致的纤维细度的增加不会导致单纤维强度的提高(注意不能确定每粒种子的纤维数量,而这些数据使得人们可以用标准方法推测纤维直径没有改变)。不过,在单纤维强度值和纤维细度和每粒种子的纤维重量之间缺乏完整相关性这一事实表明,SPS的超量表达还独立地造成了单纤维强度的提高。功的值提高了52%-61%可以用除了壁厚度增加以外的其他因素来解释。另外,正如所预料的,转基因纤维拉伸量的提高趋势不依赖于纤维壁的纤维束含量的提高(拉伸量高度取决于纤维壁内的纤维束微纤维的方向)。通过比较品系13-3a和其他转基因品系可以看出这一点。
实施例5-在夜晚低温下的光合效率
SPS在叶片中的超量表达能提高对寒冷夜晚的耐受能力,这一目的是通过在经过15℃的夜晚之后的温暖的白天保持光合速度等于温暖生长的植物的光合速度而实现的。相反,未转化过的棉花在经过寒冷的夜晚之后的温暖的白天表现出光合速度的降低。
在7-14周龄时测定生长在人工气候室中的转基因植物和亲本C312植物的光合效率。从顶上摘下第一片完全展开的叶子(通过深绿的颜色、形状、和大小判断-第三或第四叶下面),并用ADCLCA-4分析仪在不同的内部二氧化碳浓度下测定光合效率。在30/28℃下生长的植物在7-10周龄测定,在30/15℃下生长的植物在10-14周龄测定。在最初的情况下,所述植物要接触环境条件大约4周。所述植物在30℃下进行测定并在4小时进入光周期,它表示在从28℃或15℃完全回温到30℃之后的3小时。对每个生长箱中的每个品系的2个植株进行测定。
曲线表现的是亲本C312(图21)和2个转基因品系13-3a-1(图22)和225-17a(图23)在各种内部二氧化碳浓度下的光合速度。正常的大气二氧化碳浓度相当于大约270μL/L的二氧化碳浓度。每一个曲线图是4个分散曲线的组合,每一个分散曲线是测定相应品系的每一个植株得到的。要比较品系之间空心符号(30/15℃条件)和实心符号(30/28℃条件)的相对位置。比较在低于500微升/升的内部二氧化碳浓度下的光合速度,所测定的2个转基因品系的所有4个植株当在30/15℃的周期下生长时保持与植株在30/28℃的周期下生长的在温暖的白天期间相同的光合速度。相反,亲本C312表现出预期的光合速度的由夜晚低温诱导的减弱,即使该测定总是在温暖的白天进行。对于测试过的4个转基因植株中的3个来说,这种差异在所测试的所有内部二氧化碳浓度下都得到保持。
在实验时30/15℃和30/28℃生长箱中植物年龄的差异意味着温度周期之间的比较应当被视为实验性的。不过,使用来自每一种情况下活跃生长的植株的相同类型的叶片支持了它的有效性。
尚不了解超量表达SPS的植物为什么没有像亲本C312那样对冷害作出适应性的光合作用。将来对叶片碳水化合物含量所作的分析将表明在转基因植物叶片中在温暖的白天是否合成了更多的蔗糖,这种现象与较高的光合速度相关,有可能导致与亲本C312相比有更多的碳水化合物从叶片中输出以便形成纤维。这种机制可以导致超量表达SPS植物的种子和纤维产量的提高和纤维品质的提高。与亲本C312相比还发现超量表达SPS的转基因植物在其下胚轴中储存较少的淀粉。这表明外部碳水化合物的另一种来源有助于提高种子和纤维产量以及提高纤维品质。
实施例6-在储存细胞中代谢流向纤维素偏移
表2和3表示取决于纤维素含量的纤维特性,包括纤维重量/种子、纤维细度、和纤维成熟度比例,当SPS活性在叶片和纤维中都有所提高时以上参数在转基因植物中有所提高。因此,通过全植株分析,人们无法判断上述改进是由于从叶片到纤维的蔗糖输出的增加所造成的还是由于在纤维(储存)细胞中蔗糖合成的增强造成的,或者是由于这两种原因。由于一直认为纤维素合成是用蔗糖作为专用底物,在蔗糖合成酶的作用下由它产生UDP-葡萄糖,储存细胞中的SPS可以通过两种机制中的一种或两种促进代谢流朝向纤维素。SPS能够在储存细胞中再合成蔗糖,因为被转运的蔗糖在进入该细胞之前或进入后不久即被分解,和/或SPS能够再利用通过蔗糖合成酶的作用所释放的果糖来合成更多的蔗糖(图2)。
从体外培养的具有结合的发育中的纤维的棉花胚珠所获得的超量表达SPS纤维素储存细胞(以棉花纤维为代表)加强了代谢流朝向纤维素合成的证据。培养的胚珠/纤维是一种非光合系统,它利用植物组织培养基中的外部葡萄糖作为碳源维持种子和纤维成熟所需要的代谢。认可了蔗糖是纤维素合成的专用底物,SPS在仅添加葡萄糖的组织培养的胚珠/纤维中合成蔗糖。SPS还能再利用在蔗糖合成酶的作用下所释放的果糖合成更多的果糖。在该系统中所观察到的SPS超量表达的正面影响不一定独立于光合作用。不过,在该组织培养系统中底物的供应是稳定的,这意味着不可能排除增强了的蔗糖供应是由于叶片中增强了的SPS表达或者由于减弱了的下胚轴中淀粉的储存,后者在全植株上所观察到的改进方面起着重要作用。
让产生表7所示结果的植物在7月-12月之间在温室中开花。将胚珠从花上解剖下来,并在34℃下在1DPA上培养。来自一朵花的胚珠分别在30℃和15℃下进行比较。在一朵花内部进行的比较能更好地控制波动性,这种波动性是在21DPA的来自相同植物品系的不同花的培养物的纤维素合成速度上出现的。在每一种温度下的每一项测试包括分成三个重复的培养皿的12-18个胚珠。在18DPA将培养物从恒定的34℃转移到34-15℃12-12小时周期,此时次级壁沉积已经开始。在34℃和15℃下,在21DPA用14C-葡萄糖标记发育中的胚珠和纤维。因此,该培养物在调整到15℃之前有3天时间,并在21DPA在转移到15℃之后进行4小时的15℃测定。以相同方式处理的亲本C312培养物几乎总是与转基因植物品系平行测定的。
在这两种温度下测定呼吸速度(14C二氧化碳净化)和结晶纤维素合成速度(在乙酸/硝酸试剂中煮沸之后保留的不溶性14C纤维素)。将胚珠(组织)和棉纤维的代谢活性综合在所得到的数据中。不过,在测定后分离胚珠和纤维的前期工作完全证实了在34℃/15℃条件下,82%的总纤维素dpm(胚珠+纤维)是由于纤维本身产生的。
根据14C二氧化碳和14C纤维素数据计算出了每一个植物品系的4个值:(1)R%-由吸附在培养箱中氢氧化钾浸泡的滤纸上的dpm14C二氧化碳的15℃/34℃比例得到的百分比;(2)C%-由在乙酸/硝酸试剂中煮沸以后保持不溶的dpm14C二氧化碳的15℃/34℃比例得到的百分比;(3)C/R15-15℃下dpm14C纤维素和dpm14C二氧化碳的比例;和(4)C/R34-34℃下dpm14C纤维素和dpm14C二氧化碳的比例。R%和C%分别表示34℃下呼吸或纤维素合成在15℃可以被保持的比例。C/R15和C/R34分别表示在15℃或34℃下代谢流朝向纤维素合成与呼吸的比例。在表7中示出了来自亲本C312和7个测试过的转基因品系的能良好重复的平行进行的实验的结果,被认为高于亲本C312的值用黑体表示。
表7
由在体外培养物中在34℃和15℃下纤维素合成和呼吸速度计算出的
数据
植物品系 | 测试数量 | R% | C% | C/R34 | C/R15 |
C312-wt | 12 | 17.2 | 21.5 | 2.8 | 3.5 |
13-3a* | 6@T2 | 15.3 | 21.8 | 1.8 | 3.0 |
38-4a | 7@T2 | 13.0 | 25.7 | 1.9 | 3.9 |
40-4b* | 5@T2 | 13.1 | 25.4 | 1.9 | 3.7 |
40-6a* | 6@T2 | 15.4 | 20.4 | 2.8 | 3.7 |
58-3a | 4@T1 | 14.3 | 25.9 | 3.4 | 6.2 |
225-17a* | 4@T1 | 20.9 | 22.6 | 2.8 | 3.1 |
619-1a | 7@T1 | 15.9 | 24.9 | 2.9 | 4.6 |
*表示在人工气候室中被证实具有改善了的纤维品质的品系。
表7中的数据表明,SPS的超量表达能导致平行测试的7个转基因品系中有6个与亲本C312相比R%降低。与此同时在测试过的7个转基因品系中有5个的C%提高,表明大多数SPS转基因品系在15℃能比亲本C312更有效的合成纤维素。相应的,在测试过的7个转基因品系中有5个在15℃下的纤维素合成速度与呼吸速度的比例(C/R15)提高。一个转基因品系在C/R34方面表现出提高。在人工气候室中表现出具有改善了的纤维品质的转基因品系13-3a在该测定中除了R%降低之外未表现出改善。这也许是由于在体外次级壁的产生不如在植物体内进行的活跃。
实施例7-在初级和次级壁沉积期间储存细胞(棉纤维)重量增加的较高速度
所述体外胚珠/纤维培养系统业已提供了SPS在储存细胞中的超量表达通过独立于光合作用的机制可导致在温暖的温度和寒冷温度下纤维重量增加较高的速度的直接证据。
转基因和对照C312的胚珠在34℃的恒温或在34℃/15℃的循环温度下进行体外培养。在纤维成熟期间(12-45DPA),以一定间隔从平行培养物(包括来自至少3个植株的相同的代表性5-8朵花)上收获胚珠/纤维(每个数据点8-10个)。剥去胚珠上的纤维,烘干,并称重。将纤维重量对时间作图,并在两种温度方案下测定在高速次级壁纤维素合成期间重量增加的斜率。还要计算一个植物品系内的34℃/15℃、34℃比例的斜率,该斜率要规范特定品系培乔物的纤维重量增加速度的任何固有的差异。对于测试过的大多数植物品系来说,在不同时间进行该实验的若干重复,以便比较平均斜率。在第二压缩时间间隔期间的第二个实验包括生长在人工气候室中的转基因植物品系和品系38-4a-1的纤维重量增加的三个完全时间过程的重复。有关该第二种实验的结果在表中用独立的斜体形式表示,它表示的是该实验的可重复性。在表8中明显大于亲本C312亲本系的值用黑体突出。
表8
在34℃或34℃/15℃下体外培乔的纤维中纤维素沉积的速度
植物品系 | 34℃斜率 | 34/15℃斜率 | 34/15℃∶34℃斜率比例 |
C312-wt | 0.54 | 0.33 | 0.61 |
C312-wt | 0.52 | 0.31 | 0.60 |
13-3a-1* | 0.37 | 0.31 | 0.84 |
13-3a-1* | 0.45 | 0.39 | 0.87 |
38-4a-1 | 0.45 | 0.25 | 0.56 |
40-4b-1* | 0.55 | 0.19 | 0.34 |
40-4b-1* | 0.46 | 0.24 | 0.52 |
-2 | 0.36 | 0.25 | 0.69 |
-2KS** | 0.38 | 0.26 | 0.68 |
40-6a-1 | 0.38 | 0.30 | 0.78 |
-4* | 0.22 | 0.10 | 0.45 |
40-17a-6 | 0.34 | 0.28 | 0.82 |
58-3a | 0.42 | 0.41 | 0.98 |
178-1a | 0.49 | 0.20 | 0.41 |
225-17a* | 0.46 | 0.24 | 0.52 |
225-17a* | 0.58 | 0.26 | 0.45 |
414-1a | 0.63 | 0.39 | 0.62 |
619-1a | 0.60 | 0.37 | 0.62 |
*在人工气候室中测试;表示改善了的纤维品质。
KS**;上丈所述卡那霉素抗性植物的卡那霉素敏感性姊妹株;预计来自分离的种子群体的卡那霉素敏感性姊妹株不具有外源基因的拷贝。注意,40-4b-2的卡那霉素敏感性和卡那霉素敏抗性姊妹株的斜率几乎相同,并且上述斜率与亲本C312之间的斜率差异与该外源基因表达无关。
品系40-6a和40-17a被列在一起并视为一个品系,因为它有可能代表相同的转化事件,这些基于它们来自相同的亲本愈伤组织并在T1具有相同的分离比例。
在34℃下两个转基因品系(414-1a和619-1a)的纤维重量增加速度高于亲本C312,并且高于非SPS表达转基因品系40-4b-2-KS。在34℃/15℃下四个转基因品系(13-3a、58-3a、414-1a和619-1a)的纤维重量增加速度高于亲本C312。三个转基因品系(13-3a-1、40-6a-1=40-17a-6、58-3a)的34℃/15℃∶34℃斜率的比例高于亲本C312和非SPS表达转基因品系40-4b-2-KS。在斜率比例分析中没有出现品系414-1a和619-1a,因为在34℃和34℃/15℃下具有较大的斜率,但它们是将来进行纤维品质分析的有希望的品系。在该实验中,在人工气候室中的测试的某些品系具有优于亲本C312的纤维品质。缺乏完全的一致性可能是由于这样一种事实,在体外次级细胞壁的产生进程不如在植物体内活跃。
在纤维重量增加的重复的时间进程中,比较了生长在人工气候室中的转基因植物品系和品系38-4a-1在15DPA(初级壁沉积结束)和30DPA(延伸的次级壁沉积之后)纤维干重量的绝对值。每一个数据点是三次实验的平均值,包括来自每个品系的4-6个植株的15-24朵花的共24-30个胚珠的纤维。结果如表9所示。
表9
来自体外培养物的纤维重量(毫升/胚珠)
15DPA | 30DPA | |||||
植物品系 | 34℃ | 34/15℃ | 34/15℃∶34℃重量比 | 34℃ | 34/15℃ | 34/15℃∶34℃重量比 |
C312-wt | 1.75 | 0.46 | 0.263 | 8.89 | 3.88 | 0.436 |
13-3a-1* | 1.94 | 0.60 | 0.309 | 7.33 | 4.64 | 0.633 |
38-4a-1 | 1.68 | 0.67 | 0.399 | 8.68 | 3.68 | 0.424 |
40-4b-1* | 2.18 | 0.64 | 0.294 | 7.36 | 3.48 | 0.473 |
225-17a* | 1.84 | 0.59 | 0.320 | 8.80 | 3.72 | 0.423 |
*在人工气候室测试;表示改善了的纤维品质。
在15DPA,在34℃/15℃下,四个转基因品系依次地表现出高于亲本C312重量增加,并且这四个转基因品系中有三个在34℃的恒温下表现出更高的重量增加。在所有四个转基因品系中,34℃/15℃与34℃的重量比都较高,表明了SPS转基因植物在独立的低温下通过独立于光合作用的机制改善了纤维的生产。在15DPA,纤维干重量主要是由初级壁构成,并且较高的纤维重量可能是由于较大的纤维长度或较大的初级壁厚度或由于这两者。
在30DPA,在34℃/15℃下,一个转基因品系表现出高于亲本C312的纤维重量增加。有二个转基因品系表现出较大的34℃/15℃与34℃的重量比。在30DPA时的纤维干重量主要是由纤维素。因此,在转基因纤维中SPS的超量表达能促进纤维素沉积,包括其在独立的低温下的沉积。转基因品系在30DPA时的结果的不一致性可以解释为由于这样的事实,在体外次级壁的沉积比纤维的增长受到更多的妨碍。不过,在人工气候室中测试的并表现出改善了的纤维品质的所有转基因品系在该体外实验中都表现出某种改进。
实施例8-转基因棉花植物的增加了的茎干重量
SPS超量表达对棉花纤维中纤维素合成所产生的有利影响可以扩展到其他纤维。纤维构成了一年生或多年生坚固茎干的重量的大部分,如在成熟棉花植物上所见到的茎干。因此测定了生长在人工气候室中和生长在德克萨斯Tech温室中的棉花植物的茎干重量(表10)。德克萨斯Tech温室的条件最接近人工气候室30℃/15℃,360ppm二氧化碳的条件。
表10
茎干重量、直径和高度的规划值(将转基因植物的平均值相对被设
定为1.00的C312野生型亲本的相应值进行正规化)
人工气候室试验 | 温室试验 | ||||||||
植物品系 | 每个生长室中的人工气候室植株数,按顺序排列(n) | 茎重30/15℃CO2=360 | 茎重30/15℃CO2=700 | 茎重30/28℃CO2=360 | 茎重30/28℃CO2=700 | 温室植物(n) | 茎重 | 茎直径 | 茎高 |
C312-wt | 4,4,4,4 | 1.00 | 1.00 | 1.00 | 1.00 | 6 | 1.00 | 1.00 | 1.00 |
13-3a | |||||||||
T1#1@T2 | 4,4,4,4 | 1.12 | 1.20 | 1.03 | 1.11 | ||||
225-17a | |||||||||
T1 | 4,4,4,4 | 0.95 | 1.11 | 1.28 | 1.07 | ||||
40-4b | |||||||||
T1#1@T2 | 5,5,7,5 | 0.81 | 1.12 | 1.22 | 1.13 | ||||
40-6a | |||||||||
T1#4@T2 | 1,1,2,0 | 1.33 | 1.30 | 1.82 | - | ||||
T2-4-3@T3 | 5 | 1.27 | 1.11 | 1.06 | |||||
357-6a | |||||||||
T1#1@T2 | 6 | 0.92 | 0.93 | 0.94 |
在人工气候室中,对于30℃/28℃的生长室来说,不同植物品系的茎干重量测定时间略有不同,因为每一个植株是在所有的棉铃都开花以后不久收获的。对于30℃/15℃条件来说,植物生长是同时结束的,此时有一些未成熟的棉铃还保留在所有植株上。在收获时所有植株为6-7月龄。在德克萨斯Tech温室中,亲本和转基因植物随机分布在两个相邻的台子上,并生长30周,然后同时收获。还要测定温室植物的主茎直径和高度。
在人工气候室中,在15个植株中有11个转基因植物的茎干重量比亲本C312提高了10%或更高(表示植物品系×实验生长室的乘积)。在品系40-6a-4的三个生长室中所述提高特别突出和一致,不过该品系在人工气候室中仅有很少的重复植株。因此,在下一代(T3)在德克萨斯Tech温室中对品系40-6a-4-3用更多的重复与亲本C312和另一种转基因品系357-6a-1的T2代同时进行测试。品系40-6a-4-3再次表现出茎干重量的平均提高,变化幅度与在人工气候室中在30℃/15℃和360与700ppm二氧化碳条件下观察到的变化幅度相似。另外,与亲本C312和转基因品系357-6a-1(它小于C312)相比,品系40-6a-4-3表现出茎干重量和茎干直径的平均提高。因此,转基因品系并不总是表现出茎干重量的增加,这可能是由于组织专一性基因表达的差异。就植株主茎而言(排除分支,同样对分支进行称重),作为直角圆锥体其体积=πR2h/3,品系40-6a-4-3具有较大的体积,为亲本C312的1.31倍。上述结果与所观察到的重量增加1.27倍的结果的相似性表明,大多数重量增加是与含有丰富纤维的主茎的体积的增加相关。理论预计和实际观察之间4%的差异可能是由于分支程度或尚未测定的茎干密度改变的差异所致。
实施例9-在多个转基因棉花品系上茎干直径的提高
除了品系40-6a之外,生长在温室中的转基因棉花植物的某些茎干似乎比其他的要大一些。不过,这些植株具有不同的年龄。为了试图对该观测进行定量,于98年9月23日用电子量规测定生长在温室中的所有植物大约在土壤平面以上2英寸处的茎干直径(不包括在前面的研究中所提到的感兴趣的所有植物)。还要记录每一个测定过的植物的种植数据。通过分析生长在温室中的具有不同年龄的植株的C312亲本和转基因品系58-3a(2)(T1个体,数量1-7)的值,估算了每天茎干直径增加速度的如下近似值。随着时间的发展增加速度降低,因为在用于种植的2加仑的花盆中,在大约5个月时,亲本C312植物的茎干直径增加明显变慢或停止。
株龄
茎干直径增加速度
<150日 0.13毫米/日
160-200日 0.10毫米/日
>210日 0.06毫米/日
在分析过的独立的12个转基因品系中(各自有若干个重复的花盆),有6个的平均值大于所确定的亲本C312的标准值(位于其范围的上限)(表11)。未表现出茎干直径增加速度提高的转基因品系,可能在其茎干中表达菠菜SPS的能力不强。
表11
在温室中具有提高了的茎干直径增加速度的转基因植物品系
植物品系 | 株龄(日) | 茎杆直径增长速度(mm/天) |
40-4b-2-7 | 216 | 0.076 |
40-6a-4-2 | 180 | 0.124 |
-3,4 | 215 | 0.107 |
58-3a-3 | 214 | 0.078 |
414-1a-1,2 | 193 | 0.086 |
530-1a-2,3 | 197 | 0.095 |
619-1a-6 | 153 | 0.140 |
注意,表10通过第二个实验证实了品系40-6a-4-3茎干直径增加速度的提高。茎干直径的增加取决于茎干内含有较多纤维素的纤维。在生长期结束时较大的茎干直径可以解释为直径增加的速度较快或者在一个生长季节中直径增加的持续时间较长。在任一种情况下都会导致有更多可收获的茎干纤维。
实施例10-有更多的大气二氧化碳转化成可收获的作物,优选为以纤维素为基础的纤维
如表12所示,比较360和700ppm二氧化碳的30℃/15℃人工气候室的数据发现,SPS转基因植物能更有效地将正常含量的二氧化碳转化成以纤维素为基础的棉纤维。在正常二氧化碳浓度下,SPS转基因植物能够更洁净地达到其最大的纤维生产潜力(通过比较每粒种子的皮棉纤维重量证实)。因此,将二氧化碳浓度提高到700ppm,其纤维壁厚度的增加低于亲本C312(通过比较纤维细度的变化证实)。不过,当茎干重量被认为是所有类型纤维生产潜力的指标时,在30℃/15℃下甚至是在较高二氧化碳浓度下转基因植物仍然优于亲本C312。相反,在30℃/15℃下提高二氧化碳浓度倾向于降低转基因植物和亲本C312的种子重量(不过,转基因种子重量总是高于亲本C312的种子重量-参见例2)。
因此,SPS的超量表达对棉纤维生产的优势影响可能是由于提高了以这种以纤维素为基础的储存器官的储存需求。正如以前所证实,在纤维中SPS的超量表达能优先增加朝向纤维素和纤维重量增加的代谢流。在表12中示出了支持上述结论的数据,该表示出了在人工气候室中在30℃/15℃下当二氧化碳从300提高到700ppm时各种参数值的变化百分比。
表12
在30℃/15℃下当当二氧化碳从300提高到700ppm时各种作物相关
参数的改变百分比
植物品系 | 纤维细度 | 每粒种子的皮棉纤维重量 | 每粒种子的短纤维重量 | 纤维与有绒种子的重量比 | 茎重 |
C312-wt | +9% | +35% | -8% | +48% | +22% |
13-3a-1@T2 | +2% | +10% | -6% | +18% | +31% |
225-17a@T1 | -18% | -5% | -14% | +12% | +42% |
40-4b-1,4@T2 | +7% | +25% | 0% | +24% | +71% |
转基因植物平均值 | -3% | +10% | -7% | +18% | +48% |
能超量表达SPS的纤维作物可以将正常的二氧化碳更有效地转化成有经济价值的纤维。作为作物广泛种植的上述植物有助于应付大气中二氧化碳含量的提高,因为它们能在正常二氧化碳含量下以更高的效率将二氧化碳固定成纤维的纤维素,并且这种生产效率在较高二氧化碳水平下可以保持(对棉纤维而言)或增加(对茎干纤维而言)。
尽管本丈对优选实施方案作了讲解和说明,本领域技术人员显而易见的是,在不超出本发明构思的前提下可以进行各种改进、增加、替换等,因此,这种改变被认为属于由下面的权利要求所限定的本发明的范围。
序列表
<110>得克萨斯技术大学(Texas Tech University)
<120>增强了蔗糖磷酸酯合成酶表达的转基因产纤维植物
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Val Leu Ile Ser Leu His Gly Leu Ile Arg Gly Glu Asn Met Glu Leu
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His Tyr Ala Asp Ala Gly Asp Ser Ala Ala Leu Leu Ser Gly Ala Leu
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Asn Val Pro Met Val Phe Thr Gly His Ser Leu Gly Arg Asp Lys Leu
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Asp Gln Leu Leu Lys Gln Gly Arg Leu Ser Arg Glu Glu Val Asp Ala
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Thr Tyr Lys Ile Met Arg Arg Ile Glu Ala Glu Glu Leu Cys Leu Asp
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Val Gly Pro Gly Leu Asp Asp Ala Lys Ser Ser Leu Leu Leu Arg Glu
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Leu Ser Arg Asp Glu Ile Asn Ser Thr Tyr Lys Ile Met Arg Arg Ile
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Glu Ala Glu Glu Leu Ser Leu Asp Ala Ser Glu Met Val Ile Thr Ser
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Thr Arg Gln Glu Ile Glu Glu Gln Trp Arg Leu Tyr Asp Gly Phe Asp
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Pro Ile Leu Glu Arg Lys Leu Arg Ala Arg Ile Lys Arg Asn Val Ser
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Cys Tyr Gly Arg Phe Met Pro Arg Met Met Val Ile Pro Pro Gly Met
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Glu Phe His His Ile Val Pro His Asp Gly Asp Leu Asp Ala Glu Pro
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Glu Phe Asn Glu Asp Ser Lys Ser Pro Asp Pro His Ile Trp Thr Glu
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Ile Met Arg Phe Phe Ser Asn Pro Arg Lys Pro Met Ile Leu Ala Leu
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Ala Arg Pro Asp Pro Lys Lys Asn Leu Thr Thr Leu Val Lys Ala Phe
485 490 495
Gly Glu Cys Lys Pro Leu Arg Glu Leu Ala Asn Leu Thr Leu Ile Met
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Gly Asn Arg Asp Asn Ile Asp Glu Met Ser Gly Thr Asn Ala Ser Val
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Leu Leu Ser Ile Leu Lys Met Ile Asp Lys Tyr Asp Leu Tyr Gly Leu
530 535 540
Val Ala Tyr Pro Lys His His Lys Gln Ser Asp Val Pro Asp Ile Tyr
545 550 555 560
Arg Leu Ala Ala Lys Thr Lys Gly Val Phe Ile Asn Pro Ala Phe Ile
565 570 575
Glu Pro Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala His Gly Leu Pro
580 585 590
Ile Val Ala Thr Lys Asn Gly Gly Pro Val Asp Ile His Arg Val Leu
595 600 605
Asp Asn Gly Ile Leu Val Asp Pro His Asn Gln Glu Ser Ile Ala Asp
610 615 620
Ala Leu Leu Lys Leu Val Ala Glu Lys His Leu Trp Ala Lys Cys Arg
625 630 635 640
Ala Asn Gly Leu Lys Asn Ile His Leu Phe Ser Trp Pro Glu His Cys
645 650 655
Lys Ser Tyr Leu Ser Lys Leu Ala Ser Cys Lys Pro Arg Gln Pro Arg
660 665 670
Trp Leu Arg Asn Glu Glu Asp Asp Asp Glu Asn Ser Glu Ser Asp Ser
675 680 685
Pro Ser Asp Ser Leu Arg Asp Ile Gln Asp Ile Ser Leu Asn Leu Lys
690 695 700
Phe Ser Phe Asp Gly Asp Lys Asn Glu Ser Arg Glu Lys Gly Gly Gly
705 710 715 720
Ser His Pro Asp Asp Arg Ala Ser Lys Ile Glu Asn Ala Val Leu Glu
725 730 735
Trp Ser Lys Gly Val Ala Lys Gly Pro Gln Arg Ser Met Ser Ile Glu
740 745 750
Lys Gly Glu His Asn Ser Asn Ala Gly Lys Phe Pro Ala Leu Arg Arg
755 760 765
Arg Lys Ile Met Phe Val Ile Ala Val Asp Cys Lys Pro Ser Ala Gly
770 775 780
Leu Ser Glu Ser Val Arg Lys Val Phe Ala Ala Val Glu Asn Glu Arg
785 790 795 800
Ala Glu Gly Ser Val Gly Phe Ile Leu Ala Thr Ser Phe Asn Ile Ser
805 810 815
Glu Ile Arg His Phe Leu Val Ser Glu Lys Leu Asn Pro Thr Asp Phe
820 825 830
Asp Ala Phe Ile Cys Asn Ser Gly Gly Asp Leu Tyr Tyr Ser Ser His
835 840 845
His Ser Glu Asp Asn Pro Phe Val Val Asp Leu Tyr Tyr His Ser Gln
850 855 860
Ile Glu Tyr Arg Trp Gly Gly Glu Gly Leu Arg Lys Thr Leu Val Arg
865 870 875 880
Trp Ala Ala Ser Ile Thr Asp Lys Lys Gly Glu Lys Glu Glu His Val
885 890 895
Tle Ile Glu Asp Glu Glu Thr Ser Ala Asp Tyr Cys Tyr Ser Phe Lys
900 905 910
Val Gln Lys Pro Asn Val Val Pro Pro Val Lys Glu Ala Arg Lys Val
915 920 925
Met Arg Ile Gln Ala Leu Arg Cys His Val Val Tyr Cys Gln Asn Gly
930 935 940
Asn Lys Ile Asn Val Ile Pro Val Leu Ala Ser Arg Ala Gln Ala Leu
945 950 955 960
Arg Tyr Leu Tyr Leu Arg Trp Gly Met Glu Leu Ser Lys Thr Val Val
965 970 975
Val Val Gly Glu Ser Gly Asp Thr Asp Tyr Glu Glu Met Leu Gly Gly
980 985 990
Val His Lys Thr Val Val Leu Ser Gly Val Cys Thr Thr Ala Thr Asn
995 1000 1005
Leu Leu His Ala Asn Arg Ser Tyr Pro Leu Ala Asp Val Val Cys Phe
1010 1015 1020
Asp Asp Leu Asn Ile Phe Lys Thr His Asn Glu Glu Cys Ser Ser Thr
1025 1030 1035 1040
Asp Leu Arg Ala Leu Leu Glu Glu His Gly Ala Phe Lys Ala
1045 1050
<210>4
<211>1081
<212>PRT
<213>Craterostigma plantagineum
<400>4
Met Ala Gly Asn Glu Trp Ile Asn Gly Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Thr Gly Ala Ser Ala Ile Asp Glu Asn Ser Gly Gly Gly Lys Thr Ala
20 25 30
Ala Ala Gln Lys Gly Arg His His Asp His His Phe Asn Pro Thr Lys
35 40 45
Tyr Phe Val Glu Glu Val Val Ser Gly Val Asp Glu Ser Asp Leu His
50 55 60
Arg Thr Trp Ile Lys Val Val Ala Thr Arg Asn Thr Arg Glu Arg Ser
65 70 75 80
Ser Arg Leu Glu Asn Met Cys Trp Arg Ile Trp His Leu Thr Arg Lys
85 90 95
Lys Lys Gln Leu Glu Trp Glu Asp Leu Gln Arg Leu Ala Ala Arg Lys
100 105 110
Trp Glu Arg Glu Gln Gly Arg Lys Asp Val Thr Glu Asp Met Ser Glu
115 120 125
Asp Leu Ser Glu Gly Glu Lys Gly Asp Val Met Gly Glu Thr Pro Val
130 135 140
Ala Leu Asp Ser Pro Arg Gly Asn Lys Lys Tyr His Arg Asn Phe Ser
145 150 155 160
Asn Leu Glu Val Trp Ser Asp Ser Asn Lys Glu Lys Lys Leu Tyr Ile
165 170 175
Val Leu Ile Ser Leu His Gly Leu Val Arg Gly Glu Asn Met Glu Leu
180 185 190
Gly Arg Asp Ser Asp Thr Gly Gly Gln Ile Lys Tyr Val Val Glu Val
195 200 205
Ala Arg Ala Leu Ala Lys Met Pro Gly Val Tyr Arg Val Asp Leu Phe
210 215 220
Thr Arg Gln Ile Ser Ser Pro Glu Val Asp Trp Ser Tyr Ala Glu Pro
225 230 235 240
Thr Glu Met Leu Ser Ser Ser Ser Thr Thr Ala Gly Glu Ala His Glu
245 250 255
Pro Glu Glu Glu Glu Glu Glu Glu Asp Leu Gly Glu Gly Ser Gly Ala
260 265 270
Tyr Ile Ile Arg Ile Pro Phe Gly Pro Arg Asp Lys Tyr Leu Arg Lys
275 280 285
Glu Leu Leu Trp Pro His Ile Gln Glu Phe Val Asp Gly Ala Leu Ser
290 295 300
His Ile Val Asn Met Ser Lys Ala Leu Gly Asp Gln Ile Gly Gly Gly
305 310 315 320
Gln Pro Val Trp Pro Tyr Val Ile His Gly His Tyr Ala Asp Ala Gly
325 330 335
Asp Ser Ala Ala Leu Leu Ser Gly Ala Leu Asn Val Pro Met Val Leu
340 345 350
Thr Gly His Ser Leu Gly Arg Asn Lys Leu Glu Gln Leu Leu Lys Gln
355 360 365
Gly Arg Gln Thr Lys Glu Asp Ile Asn Ser Met Tyr Arg Ile Met Arg
370 375 380
Arg Ile Glu Ala Glu Glu Leu Ser Leu Asp Ala Ala Glu Leu Val Ile
385 390 395 400
Thr Ser Thr Lys Gln Glu Ile Glu Glu Gln Trp Gly Leu Tyr Asp Gly
405 410 415
Phe Asp Val Lys Leu Glu Arg Val Leu Arg Ala Arg Ala Arg Arg Gly
420 425 430
Val Asn Cys His Gly Arg Phe Met Pro Arg Met Ala Val Ile Pro Pro
435 440 445
Gly Met Asp Phe Ser Asn Val Val Val Pro Glu Asp Gly Ser Glu Gly
450 455 460
Asp Gly Asp Leu Ala Thr Leu Thr Glu Ala Thr Ser Pro Arg Ser Val
465 470 475 480
Pro Ala Ile Trp Ala Asp Val Met Arg Phe Leu Thr Asn Pro His Lys
485 490 495
Pro Met Ile Leu Ala Leu Ser Arg Pro Asp Pro Lys Lys Asn Ile Thr
500 505 510
Thr Leu Val Lys Ala Phe Gly Glu Cys Arg Pro Leu Arg Glu Leu Ala
515 520 525
Asn Leu Thr Leu Ile Met Gly Asn Arg Asp Asp Ile Asp Glu Met Ser
530 535 540
Gly Gly Asn Ala Ser Val Leu Thr Thr Val Leu Lys Leu Ile Asp Arg
545 550 555 560
Tyr Asp Leu Tyr Gly Gln Val Ala Phe Pro Lys His His Lys Gln Ser
565 570 575
Asp Val Pro Glu Ile Tyr Arg Leu Ala Ser Lys Thr Lys Gly Val Phe
580 585 590
Ile Asn Pro Ala Phe Ile Glu Pro Phe Gly Leu Thr Leu Ile Glu Ala
595 600 605
Ala Ala His Gly Leu Pro Met Val Ala Thr Lys Asn Gly Gly Pro Val
610 615 620
Asp Ile His Arg Ala Leu Asn Asn Gly Leu Leu Val Asp Pro His Asp
625 630 635 640
Gln Asp Ala Ile Ala Asn Ala Leu Leu Lys Leu Val Ser Glu Lys Asn
645 650 655
Leu Trp Asn Glu Cys Arg Lys Asn Gly Leu Lys Asn Ile His Leu Phe
660 665 670
Ser Trp Pro Glu His Cys Arg Thr Tyr Leu Thr Arg Val Ala Ala Cys
675 680 685
Arg Met Arg His Pro Gln Trp Lys Thr Asp Thr Pro Leu Asp Glu Thr
690 695 700
Ala Ile Asp Asp Ser Leu Asn Asp Ser Leu Lys Asp Val Leu Asp Met
705 710 715 720
Ser Leu Arg Leu Ser Val Asp Gly Glu Lys Met Ser Val Asn Glu Ser
725 730 735
Ser Ser Val Glu Leu Pro Gly Gly Glu Ala Ala Glu Leu Pro Asp Gln
740 745 750
Val Arg Arg Val Leu Asn Lys Ile Lys Arg Gln Asp Ser Gly Pro Ala
755 760 765
Gln Arg Glu Ala Glu Gly Lys Ala Gly Asp Val Pro Gly Lys Tyr Pro
770 775 780
Met Leu Arg Arg Arg Arg Lys Leu Phe Val Ile Ala Leu Asp Cys Tyr
785 790 795 800
Asp Leu Lys Gly Asn Pro Asp Lys Lys Met Ile Leu Ser Ile Gln Glu
805 810 815
Ile Val Arg Ala Val Arg Leu Asp Pro Gln Met Ser Arg Phe Ser Gly
820 825 830
Phe Ala Leu Ser Thr Ala Met Pro Val Ala Glu Leu Ala Asp Phe Leu
835 840 845
Lys Ala Gly Asp Val Lys Val Asn Asp Phe Asp Ala Leu Ile Cys Ser
850 855 860
Ser Gly Ser Glu Val Tyr Tyr Pro Gly Thr Tyr Gly Glu Glu Ser Gly
865 870 875 880
Lys Leu Tyr Leu Asp Pro Asp Tyr Thr Ser His Ile Glu Tyr Arg Trp
885 890 895
Gly Gly Asp Gly Leu Lys Lys Thr Ile Ser Lys Leu Met Asn Thr Ala
900 905 910
Glu Asp Gly Lys Ser Ser Val Ala Ser Ser Pro Ile Glu Leu Val Ala
915 920 925
Lys Ser Ser Asn Ser His Cys Leu Ser Tyr Ala Ile Lys Asp Pro Ser
930 935 940
Lys Ala Lys Lys Val Asp Asp Met Arg Gln Lys Leu Arg Met Arg Gly
945 950 955 960
Leu Arg Cys His Leu Met Tyr Cys Arg Asn Ser Thr Ser Met Gln Val
965 970 975
Val Pro Leu Leu Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Phe Val
980 985 990
Arg Trp Arg Leu Ser Val Ala Asn Met Tyr Val Ile Leu Gly Glu Thr
995 1000 1005
Gly Asp Thr Asp Tyr Glu Glu Leu Ile Ser Gly Thr His Lys Thr Leu
1010 1015 1020
Ile Met Arg Gly Val Val Glu Lys Gly Ser Glu Glu Leu Leu Arg Thr
1025 1030 1035 1040
Ala Gly Ser Tyr Leu Arg Asp Asp Val Ile Pro Gln Asp Thr Pro Leu
1045 1050 1055
Ile Ala Tyr Ala Asp Lys Gly Ala Lys Ala Glu His Ile Val Glu Thr
1060 1065 1070
Phe Arg Gln Leu Ser Lys Ala Gly Met
1075 1080
<210>5
<211>1059
<212>PRT
<213>蚕豆(Vicia faba)
<400>5
Met Ala Gly Asn Asp Trp Leu Asn Ser Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Val Gly Pro Gly Leu Asp Asp Ala Lys Ser Ser Leu Leu Leu Arg Glu
20 25 30
Arg Gly Arg Phe Ser Pro Thr Arg Tyr Phe Val Glu Glu Val Ile Gly
35 40 45
Phe Asp Glu Thr Asp Leu Tyr Arg Ser Trp Val Arg Ala Ser Ser Ser
50 55 60
Arg Ser Pro Gln Glu Arg Asn Thr Arg Leu Glu Asn Met Cys Trp Arg
65 70 75 80
Ile Trp Asn Leu Ala Arg Gln Lys Lys Gln Leu Glu Ser Glu Ala Val
85 90 95
Gln Arg Val Asn Lys Arg Arg Leu Glu Arg Glu Arg Gly Arg Arg Glu
100 105 110
Ala Thr Ala Asp Met Ser Glu Asp Leu Ser Glu Gly Glu Arg Gly Asp
115 120 125
Pro Val Ser Asp Val Ser Thr His Gly Gly Gly Asp Ser Val Lys Ser
130 135 140
Arg Leu Pro Arg Ile Ser Ser Ala Asp Ala Met Glu Thr Trp Val Asn
145 150 155 160
Ser Gln Lys Gly Lys Lys Leu Tyr Ile Val Leu Ile Ser Ile His Gly
165 170 175
Leu Ile Arg Gly Glu Asn Met Glu Leu Gly Arg Asp Ser Asp Thr Gly
180 185 190
Gly Gln Val Lys Tyr Val Val Glu Leu Ala Arg Ala Leu Gly Ser Met
195 200 205
Pro Gly Val Tyr Arg Val Asp Leu Leu Thr Arg Gln Val Ser Ser Pro
210 215 220
Asp Val Asp Trp Ser Tyr Gly Glu Pro Thr Glu Met Leu Ala Pro Arg
225 230 235 240
Asn Thr Asp Glu Phe Gly Asp Asp Met Gly Glu Ser Ser Gly Ala Tyr
245 250 255
Ile Ile Arg Ile Pro Phe Gly Pro Arg Asn Lys Tyr Ile Pro Lys Glu
260 265 270
Glu Leu Trp Pro Tyr Ile Pro Glu Phe Val Asp Gly Ala Met Gly His
275 280 285
Ile Ile Gln Met Ser Lys Ala Leu Gly Glu Gln Ile Gly Ser Gly His
290 295 300
Ala Val Trp Pro Val Ala Ile His Gly His Tyr Ala Asp Ala Gly Asp
305 310 315 320
Ser Ala Ala Leu Leu Ser Gly Ala Leu Asn Val Pro Met Ile Phe Thr
325 330 335
Gly His Ser Leu Gly Arg Asp Lys Leu Glu Gln Leu Leu Lys Gln Gly
340 345 350
Arg Leu Ser Thr Asp Glu Ile Asn Ser Thr Tyr Lys Ile Met Arg Arg
355 360 365
Ile Glu Ala Glu Glu Leu Ala Leu Asp Gly Thr Glu Ile Val Ile Thr
370 375 380
Ser Thr Arg Gln Glu Ile Glu Glu Gln Trp Arg Leu Tyr Asn Gly Phe
385 390 395 400
Asp Pro Val Leu Glu Arg Lys Ile Arg Ala Arg Ile Arg Arg Asn Val
405 410 415
Ser Cys Tyr Gly Arg Tyr Met Pro Arg Met Ser Val Ile Pro Pro Gly
420 425 430
Met Glu Phe His His Ile Ala Pro Leu Asp Gly Asp Ile Glu Thr Glu
435 440 445
Pro Glu Gly Ile Leu Asp His Pro Ala Pro Gln Asp Pro Pro Ile Trp
450 455 460
Ser Glu Ile Met Arg Phe Phe Ser Asn Pro Arg Lys Pro Val Ile Leu
465 470 475 480
Ala Leu Ala Arg Pro Asp Pro Lys Lys Asn Ile Thr Thr Leu Val Lys
485 490 495
Ala Phe Gly Glu Cys Arg Pro Leu Arg Glu Leu Ala Asn Leu Thr Leu
500 505 510
Ile Met Gly Asn Arg Asp Gly Ile Asp Glu Met Ser Ser Thr Ser Ser
515 520 525
Ser Val Leu Leu Ser Val Leu Lys Leu Ile Asp Lys Tyr Asp Leu Tyr
530 535 540
Gly Gln Val Ala Tyr Pro Lys His His Lys Gln Ser Asp Val Pro Asp
545 550 555 560
Ile Tyr Arg Leu Ala Ala Lys Thr Lys Gly Val Phe Ile Asn Pro Ala
565 570 575
Phe Ile Glu Pro Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala Tyr Gly
580 585 590
Leu Pro Met Val Ala Thr Lys Asn Gly Gly Pro Val Asp Ile His Arg
595 600 605
Val Leu Asp Asn Gly Leu Leu Ile Asp Pro His Asp Glu Lys Ser Ile
610 615 620
Ala Asp Ala Leu Leu Lys Leu Val Ser Asn Lys Gln Leu Trp Ala Lys
625 630 635 640
Cys Arg Gln Asn Gly Leu Lys Asn Ile His Leu Phe Ser Trp Pro Glu
645 650 655
His Cys Lys Thr Tyr Leu Ser Lys Ile Ala Thr Cys Lys Pro Arg His
660 665 670
Pro Gln Trp Gln Arg Ser Glu Asp Gly Gly Glu Ser Ser Glu Ser Glu
675 680 685
Glu Ser Pro Gly Asp Ser Leu Arg Asp Ile Gln Asp Leu Ser Leu Asn
690 695 700
Leu Lys Phe Ser Leu Asp Gly Glu Arg Ser Gly Asp Ser Gly Asn Asp
705 710 715 720
Asn Ser Leu Asp Pro Asp Gly Asn Ala Thr Asp Arg Thr Thr Lys Leu
725 730 735
Glu Asn Ala Val Leu Ser Trp Ser Lys Gly Ile Ser Lys Asp Thr Arg
740 745 750
Arg Gly Gly Ala Thr Glu Lys Ser Gly Gln Asn Ser Asn Ala Ser Lys
755 760 765
Phe Pro Pro Leu Arg Ser Arg Asn Arg Leu Phe Val Ile Ala Val Asp
770 775 780
Cys Asp Thr Thr Ser Gly Leu Leu Glu Met Ile Lys Leu Ile Phe Glu
785 790 795 800
Ala Ala Gly Glu Glu Arg Ala Glu Gly Ser Val Gly Phe Ile Leu Ser
805 810 815
Thr Ser Leu Thr Ile Ser Glu Ile Gln Ser Phe Leu Ile Ser Gly Gly
820 825 830
Leu Ser Pro Asn Asp Phe Asp Ala Tyr Ile Cys Asn Ser Gly Ser Asp
835 840 845
Leu Tyr Tyr Pro Ser Leu Asn Ser Glu Asp Arg Leu Phe Val Gly Asp
850 855 860
Leu Tyr Phe His Ser His Ile Glu Tyr Arg Trp Gly Gly Glu Gly Leu
865 870 875 880
Arg Lys Thr Leu Ile Arg Trp Ala Ser Ser Ile Thr Asp Lys Lys Ser
885 890 895
Glu Asn Asn Glu Gln Ile Val Ser Pro Ala Glu Gln Leu Ser Thr Asp
900 905 910
Tyr Cys Tyr Ala Phe Asn Val Arg Lys Ala Gly Met Ala Pro Pro Leu
915 920 925
Lys Glu Leu Arg Lys Leu Met Arg Ile Gln Ala Leu Arg Cys His Pro
930 935 940
Ile Tyr Cys Gln Asn Gly Thr Arg Leu Asn Val Ile Pro Val Leu Ala
945 950 955 960
Ser Arg Ser Gln Ala Leu Arg Tyr Leu Tyr Val Arg Trp Gly Phe Glu
965 970 975
Leu Ser Lys Met Val Val Phe Val Gly Glu Cys Gly Asp Thr Asp Tyr
980 985 990
Glu Gly Leu Val Gly Gly Leu His Lys Ser Val Ile Leu Lys Gly Val
995 1000 1005
Gly Ser Arg Ala Ile Ser Gln Leu His Asn Asn Arg Asn Tyr Pro Leu
1010 1015 1020
Ser Asp Val Met Pro Leu Asp Ser Pro Asn Ile Val Gln Ala Thr Glu
1025 1030 1035 1040
Gly Ser Ser Ser Ala Asp Ile Gln Ala Leu Leu Glu Lys Val Gly Tyr
1045 1050 1055
His Lys Gly
<210>6
<211>1053
<212>PRT
<213>马铃薯(Solanum tuberosum)
<400>6
Met Ala Gly Asn Asp Trp Ile Asn Ser Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Val Gly Pro Gly Leu Asp Asp Lys Lys Ser Ser Leu Leu Leu Arg Glu
20 25 30
Arg Gly Arg Phe Ser Pro Thr Arg Tyr Phe Val Glu Glu Val Ile Thr
35 40 45
Gly Phe Asp Glu Thr Asp Leu His Arg Ser Trp Ile Arg Ala Gln Ala
50 55 60
Thr Arg Ser Pro Gln Arg Arg Asn Thr Arg Leu Glu Asn Met Cys Trp
65 70 75 80
Arg Ile Trp Asn Leu Ala Arg Gln Lys Lys Gln Leu Glu Gly Glu Gln
85 90 95
Ala Gln Trp Met Ala Lys Arg Arg Gln Glu Arg Glu Arg Gly Arg Arg
100 105 110
Glu Ala Val Ala Asp Met Ser Glu Asp Leu Ser Glu Gly Glu Lys Gly
115 120 125
Asp Ile Val Ala Asp Met Ser Ser His Gly Glu Ser Thr Arg Gly Arg
130 135 140
Leu Pro Arg Ile Ser Ser Val Glu Thr Met Glu Ala Trp Val Ser Gln
145 150 155 160
Gln Arg Gly Lys Lys Leu Tyr Ile Val Leu Ile Ser Leu His Gly Leu
165 170 175
Ile Arg Gly Glu Asn Met Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly
180 185 190
Gln Val Lys Tyr Val Val Glu Leu Ala Arg Ala Leu Gly Ser Met Pro
195 200 205
Gly Val Tyr Arg Val Asp Leu Leu Thr Arg Gln Val Ser Ser Pro Glu
210 215 220
Val Asp Trp Ser Tyr Gly Glu Pro Thr Glu Leu Ala Pro Ile Ser Thr
225 230 235 240
Asp Gly Leu Met Thr Glu Met Gly Glu Ser Ser Gly Ala Tyr Ile Ile
245 250 255
Arg Ile Pro Phe Gly Pro Arg Glu Lys Tyr Ile Pro Lys Glu Gln Leu
260 265 270
Trp Pro Tyr Ile Pro Glu Phe Val Asp Gly Ala Leu Asn His Ile Ile
275 280 285
Gln Met Ser Lys Val Leu Gly Glu Gln Ile Gly Ser Gly Tyr Pro Val
290 295 300
Trp Pro Val Ala Ile His Gly His Tyr Ala Asp Ala Gly Asp Ser Ala
305 310 315 320
Ala Leu Leu Ser Gly Ala Leu Asn Val Pro Met Leu Phe Thr Gly His
325 330 335
Ser Leu Gly Arg Asp Lys Leu Glu Gln Leu Leu Ala Gln Gly Arg Lys
340 345 350
Ser Lys Asp Glu Ile Asn Ser Thr Tyr Lys Ile Met Arg Arg Ile Glu
355 360 365
Ala Glu Glu Leu Thr Leu Asp Ala Ser Glu Ile Val Ile Thr Ser Thr
370 375 380
Arg Gln Glu Ile Asp Glu Gln Trp Arg Leu Tyr Asp Gly Phe Asp Pro
385 390 395 400
Ile Leu Glu Arg Lys Leu Arg Ala Arg Ile Lys Arg Asn Val Ser Cys
405 410 415
Tyr Gly Arg Phe Met Pro Arg Met Ala Val Ile Pro Pro Gly Met Glu
420 425 430
Phe His His Ile Val Pro His Glu Gly Asp Met Asp Gly Glu Thr Glu
435 440 445
Gly Ser Glu Asp Gly Lys Thr Pro Asp Pro Pro Ile Trp Ala Glu Ile
450 455 460
Met Arg Phe Phe Ser Asn Pro Arg Lys Pro Met Ile Leu Ala Leu Ala
465 470 475 480
Arg Pro Asp Pro Lys Lys Asn Leu Thr Thr Leu Val Lys Ala Phe Gly
485 490 495
Glu Cys Arg Pro Leu Arg Asp Leu Ala Asn Leu Thr Leu Ile Met Gly
500 505 510
Asn Arg Asp Asn Ile Asp Glu Met Ser Ser Thr Asn Ser Ala Leu Leu
515 520 525
Leu Ser Ile Leu Lys Met Ile Asp Lys Tyr Asp Leu Tyr Gly Gln Val
530 535 540
Ala Tyr Pro Lys His His Lys Gln Ser Asp Val Pro Asp Ile Tyr Arg
545 550 555 560
Leu Ala Ala Lys Thr Lys Gly Val Phe Ile Asn Pro Ala Phe Ile Glu
565 570 575
Pro Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala Tyr Gly Leu Pro Met
580 585 590
Val Ala Thr Lys Asn Gly Gly Pro Val Asp Ile His Arg Val Leu Asp
595 600 605
Asn Gly Leu Leu Val Asp Pro His Asp Gln Gln Ala Ile Ala Asp Ala
610 615 620
Leu Leu Lys Leu Val Ala Asp Lys Gln Leu Trp Ala Lys Cys Arg Ala
625 630 635 640
Asn Gly Leu Lys Asn Ile His Leu Phe Ser Trp Pro Glu His Cys Lys
645 650 655
Thr Tyr Leu Ser Arg Ile Ala Ser Cys Lys Pro Arg Gln Pro Arg Trp
660 665 670
Leu Arg Ser Ile Asp Asp Asp Asp Glu Asn Ser Glu Thr Asp Ser Pro
675 680 685
Ser Asp Ser Leu Arg Asp Ile His Asp Ile Ser Leu Asn Leu Arg Phe
690 695 700
Ser Leu Asp Gly Glu Lys Asn Asp Asn Lys Glu Asn Ala Asp Asn Thr
705 710 715 720
Leu Asp Pro Glu Val Arg Arg Ser Lys Leu Glu Asn Ala Val Leu Ser
725 730 735
Leu Ser Lys Gly Ala Leu Lys Ser Thr Ser Lys Ser Trp Ser Ser Asp
740 745 750
Lys Ala Asp Gln Asn Pro Gly Ala Gly Lys Phe Pro Ala Ile Arg Arg
755 760 765
Arg Arg His Ile Phe Val Ile Ala Val Asp Cys Asp Ala Ser Ser Gly
770 775 780
Leu Ser Gly Ser Val Lys Lys Ile Phe Glu Ala Val Glu Lys Glu Arg
785 790 795 800
Ala Glu Gly Ser Ile Gly Phe Ile Leu Ala Thr Ser Phe Asn Ile Ser
805 810 815
Glu Val Gln Ser Phe Leu Leu Ser Glu Gly Met Asn Pro Thr Asp Phe
820 825 830
Asp Ala Tyr Ile Cys Asn Ser Gly Gly Asp Leu Tyr Tyr Ser Ser Phe
835 840 845
His Ser Glu Gln Asn Pro Phe Val Val Asp Leu Tyr Tyr His Ser His
850 855 860
Ile Glu Tyr Arg Trp Gly Gly Glu Gly Leu Arg Lys Thr Leu Val Arg
865 870 875 880
Trp Ala Ala Ser Ile Ile Asp Lys Asn Gly Glu Asn Gly Asp His Ile
885 890 895
Val Val Glu Asp Glu Asp Asn Ser Ala Asp Tyr Cys Tyr Thr Phe Lys
900 905 910
Val Cys Lys Pro Gly Thr Val Pro Pro Ser Lys Glu Leu Arg Lys Val
915 920 925
Met Arg Ile Gln Ala Leu Arg Cys His Ala Val Tyr Cys Gln Asn Gly
930 93 5940
Ser Arg Ile Asn Val Ile Pro Val Leu Ala Ser Arg Ser Gln Ala Leu
945 950 955 960
Arg Tyr Leu Tyr Leu Arg Trp Gly Met Asp Leu Ser Lys Leu Val Val
965 970 975
Phe Val Gly Glu Ser Gly Asp Thr Asp Tyr Glu Gly Leu Ile Gly Gly
980 985 990
Leu Arg Lys Ala Val Ile Met Lys Gly Leu Cys Thr Asn Ala Ser Ser
995 1000 1005
Leu Ile His Gly Asn Arg Asn Tyr Pro Leu Ser Asp Val Leu Pro Phe
1010 1015 1020
Asp Ser Pro Asn Val Ile Gln Ala Asp Glu Glu Cys Ser Ser Thr Glu
1025 1030 1035 1040
Ile Arg Cys Leu Leu Glu Lys Leu Ala Val Leu Lys Gly
1045 1050
<210>7
<211>1045
<212>PRT
<213>甜菜(Beta vulgaris)
<400>7
Met Ala Gly Asn Asp Trp Ile Asn Ser Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Val Gly Pro Gly Leu Asp Asp Ala Lys Ser Ser Leu Leu Leu Arg Glu
20 25 30
Arg Gly Arg Phe Ser Pro Thr Arg Tyr Phe Val Glu Glu Val Ile Thr
35 40 45
Gly Phe Asp Glu Thr Asp Leu His Arg Ser Trp Val Arg Ala Gln Ala
50 55 60
Thr Arg Ser Pro Gln Glu Arg Asn Thr Arg Leu Glu Asn Met Cys Trp
65 70 75 80
Arg Ile Trp Asn Leu Ala Arg Gln Lys Lys Gln Leu Glu Asn Glu Glu
85 90 95
Ala Gln Arg Lys Thr Lys Arg Arg Met Glu Leu Glu Arg Gly Arg Arg
100 105 110
Glu Ala Thr Ala Asp Met Ser Glu Asp Leu Ser Glu Gly Glu Lys Asp
115 120 125
Ile Ser Ala His Gly Asp Ser Thr Arg Pro Arg Leu Pro Arg Ile Asn
130 135 140
Ser Leu Asp Ala Met Glu Thr Trp Ile Ser Gln Gln Lys Glu Lys Lys
145 150 155 160
Leu Tyr Leu Val Leu Ile Ser Leu His Gly Leu Ile Arg Gly Glu Asn
165 170 175
Met Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr Val
180 185 190
Val Glu Leu Ala Arg Ala Leu Gly Ser Met Pro Gly Val Tyr Arg Val
195 200 205
Asp Leu Leu Thr Arg Gln Val Ser Ser Pro Asp Val Asp Trp Ser Tyr
210 215 220
Gly Glu Pro Thr Glu Met Leu Asn Pro Arg Asp Ser Asn Gly Phe Asp
225 230 235 240
Asp Asp Asp Asp Glu Met Gly Glu Ser Ser Gly Ala Tyr Ile Val Arg
245 250 255
Ile Pro Phe Gly Pro Arg Asp Lys Tyr Ile Ala Lys Glu Glu Leu Trp
260 265 270
Pro Tyr Ile Pro Glu Phe Val Asp Gly Ala Leu Asn His Ile Val Gln
275 280 285
Met Ser Lys Val Leu Gly Glu Gln Ile Gly Ser Gly Glu Thr Val Trp
290 295 300
Pro Val Ala Ile His Gly His Tyr Ala Asp Ala Gly Asp Ser Ala Ala
305 310 315 320
Leu Leu Ser Gly Gly Leu Asn Val Pro Met Leu Leu Thr Gly His Ser
325 330 335
Leu Gly Arg Asp Lys Leu Glu Gln Leu Leu Lys Gln Gly Arg Met Ser
340 345 350
Lys Asp Asp Ile Asn Asn Thr Tyr Lys Ile Met Arg Arg Ile Glu Ala
355 360 365
Glu Glu Leu Ser Leu Asp Ala Ser Glu Ile Val Ile Thr Ser Thr Arg
370 375 380
Gln Glu Ile Glu Glu Gln Trp His Leu Tyr Asp Gly Phe Asp Pro Val
385 390 395 400
Leu Glu Arg Lys Leu Arg Ala Arg Met Lys Arg Gly Val Ser Cys Tyr
405 410 415
Gly Arg Phe Met Pro Arg Met Val Val Ile Pro Pro Gly Met Glu Phe
420 425 430
Asn His Ile Val Pro His Glu Gly Asp Met Asp Gly Glu Thr Glu Glu
435 440 445
Thr Glu Glu His Pro Thr Ser Pro Asp Pro Pro Ile Trp Ala Glu Ile
450 455 460
Met Arg Phe Phe Ser Lys Pro Arg Lys Pro Met Ile Leu Ala Leu Ala
465 470 475 480
Arg Pro Asp Pro Lys Lys Asn Ile Thr Thr Leu Val Lys Ala Phe Gly
485 490 495
Glu Cys Arg Pro Leu Arg Glu Leu Ala Asn Leu Thr Leu Ile Met Gly
500 505 510
Asn Arg Asp Gly Ile Asp Glu Met Ser Ser Thr Ser Ser Ser Val Leu
515 520 525
Leu Ser Val Leu Lys Leu Ile Asp Gln Tyr Asp Leu Tyr Gly Gln Val
530 535 540
Ala Tyr Pro Lys His His Lys Gln Ala Asp Val Pro Glu Ile Tyr Arg
545 550 555 560
Leu Ala Ala Lys Thr Lys Gly Val Phe Ile Asn Pro Ala Phe Ile Glu
565 570 575
Pro Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala His Gly Leu Pro Met
580 585 590
Val Ala Thr Lys Asn Gly Gly Pro Val Asp Ile Gln Arg Val Leu Asp
595 600 605
Asn Gly Leu Leu Val Asp Pro His Glu Gln Gln Ser Ile Ala Thr Ala
610 615 620
Leu Leu Lys Leu Val Ala Asp Lys Gln Leu Trp Thr Lys Cys Gln Gln
625 630 635 640
Asn Gly Leu Lys Asn Ile His Leu Tyr Ser Trp Pro Glu His Ser Lys
645 650 655
Thr Tyr Leu Ser Arg Ile Ala Ser Ser Arg Gln Arg Gln Pro Gln Trp
660 665 670
Gln Arg Ser Ser Asp Glu Gly Leu Asp Asn Gln Glu Pro Glu Ser Pro
675 680 685
Ser Asp Ser Leu Arg Asp Ile Lys Asp Ile Ser Leu Asn Leu Glu Val
690 695 700
Leu Val Arg Pro Glu Lys Arg Val Lys Thr Leu Lys Ile Leu Gly Leu
705 710 715 720
Met Thr Lys Ala Asn Ser Arg Met Leu Leu Cys Ser Trp Ser Asn Gly
725 730 735
Val His Lys Met Leu Arg Lys Ala Arg Phe Ser Asp Lys Val Asp Gln
740 745 750
Ala Ser Ser Lys Tyr Pro Ala Phe Arg Arg Arg Lys Leu Ile Tyr Val
755 760 765
Ile Ala Val Asp Gly Asp Tyr Glu Asp Gly Leu Phe Asp Ile Val Arg
770 775 780
Arg Ile Phe Asp Ala Ala Gly Lys Glu Lys Ile Glu Gly Ser Ile Gly
785 790 795 800
Phe Ile Leu Ser Thr Ser Tyr Ser Met Pro Glu Ile Gln Asn Tyr Leu
805 810 815
Leu Ser Lys Gly Phe Asn Leu His Asp Phe Asp Ala Tyr Ile Cys Asn
820 825 830
Ser Gly Ser Glu Leu Tyr Tyr Ser Ser Leu Asn Ser Glu Glu Ser Asn
835 840 845
Ile Ile Ala Asp Ser Asp Tyr His Ser His Ile Glu Tyr Arg Trp Gly
850 855 860
Gly Glu Gly Leu Arg Arg Thr Leu Leu Arg Trp Ala Ala Ser Ile Thr
865 870 875 880
Glu Lys Asn Gly Glu Asn Glu Glu Gln Val Ile Thr Glu Asp Glu Glu
885 890 895
Val Ser Thr Gly Tyr Cys Phe Ala Phe Lys Ile Lys Asn Gln Asn Lys
900 905 910
Val Pro Pro Thr Lys Glu Leu Arg Lys Ser Met Arg Ile Gln Ala Leu
915 920 925
Arg Cys His Val Ile Tyr Cys Gln Asn Gly Ser Lys Met Asn Val Ile
930 935 940
Pro Val Leu Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Tyr Val Arg
945 950 955 960
Trp Gly Val Glu Leu Ser Lys Met Val Val Phe Val Gly Glu Cys Gly
965 970 975
Asp Thr Asp Tyr Glu Gly Leu Leu Gly Gly Val His Lys Thr Val Ile
980 985 990
Leu Lys Gly Val Ser Asn Thr Ala Leu Arg Ser Leu His Ala Asn Arg
995 1000 1005
Ser Tyr Pro Leu Ser His Val Val Ser Leu Asp Ser Pro Asn Ile Gly
1010 1015 1020
Glu Val Ser Lys Gly Cys Ser Ser Ser Glu Ile Gln Ser Ile Val Thr
1025 1030 1035 1040
Lys Leu Ser Lys Ala
1045
<210>8
<211>1068
<212>PRT
<213>玉米(Zea mays)
<400>8
Met Ala Gly Asn Glu Trp Ile Asn Gly Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Ser His Thr Ser Ser Arg Gly Ala Gly Gly Gly Gly Gly Gly Gly Asp
20 25 30
Pro Arg Ser Pro Thr Lys Ala Ala Ser Pro Arg Gly Ala His Met Asn
35 40 45
Phe Asn Pro Ser His Tyr Phe Val Glu Glu Val Val Lys Gly Val Asp
50 55 60
Glu Ser Asp Leu His Arg Thr Trp Ile Lys Val Val Ala Thr Arg Asn
65 70 75 80
Ala Arg Glu Arg Ser Thr Arg Leu Glu Asn Met Cys Trp Arg Ile Trp
85 90 95
His Leu Ala Arg Lys Lys Lys Gln Leu Glu Leu Glu Gly Ile Gln Arg
100 105 110
Ile Ser A1a Arg Arg Lys Glu Gln Glu Gln Val Arg Arg Glu Ala Thr
115 120 125
Glu Asp Leu Ala Glu Asp Leu Ser Glu Gly Glu Lys Gly Asp Thr Ile
130 135 140
Gly Glu Leu Ala Pro Val Glu Thr Thr Lys Lys Lys Phe Gln Arg Asn
145 150 155 160
Phe Ser Asp Leu Thr Val Trp Ser Asp Asp Asn Lys Glu Lys Lys Leu
165 170 175
Tyr Ile Val Leu Ile Ser Val His Gly Leu Val Arg Gly Glu Asn Met
180 185 190
Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr Val Val
195 200 205
Glu Leu Ala Arg Ala Met Ser Met Met Pro Gly Val Tyr Arg Val Asp
210 215 220
Leu Phe Thr Arg Gln Val Ser Ser Pro Asp Val Asp Trp Ser Tyr Gly
225 230 235 240
Glu Pro Thr Glu Met Leu Cys Ala Gly Ser Asn Asp Gly Glu Gly Met
245 250 255
Gly Glu Ser Gly Gly Ala Tyr Ile Val Arg Ile Pro Cys Gly Pro Arg
260 265 270
Asp Lys Tyr Leu Lys Lys Glu Ala Leu Trp Pro Tyr Leu Gln Glu Phe
275 280 285
Val Asp Gly Ala Leu Ala His Ile Leu Asn Met Ser Lys Ala Leu Gly
290 295 300
Glu Gln Val Gly Asn Gly Arg Pro Val Leu Pro Tyr Val Ile His Gly
305 310 315 320
His Tyr Ala Asp Ala Gly Asp Val Ala Ala Leu Leu Ser Gly Ala Leu
325 330 335
Asn Val Pro Met Val Leu Thr Gly His Ser Leu Gly Arg Asn Lys Leu
340 345 350
Glu Gln Leu Leu Lys Gln Gly Arg Met Ser Lys Glu Glu Ile Asp Ser
355 360 365
Thr Tyr Lys Ile Met Arg Arg Ile Glu Gly Glu Glu Leu Ala Leu Asp
370 375 380
Ala Ser Glu Leu Val Ile Thr Ser Thr Arg Gln Glu Ile Asp Glu Gln
385 390 395 400
Trp Gly Leu Tyr Asp Gly Phe Asp Val Lys Leu Glu Lys Val Leu Arg
405 410 415
Ala Arg Ala Arg Arg Gly Val Ser Cys His Gly Arg Tyr Met Pro Arg
420 425 430
Met Val Val Ile Pro Pro Gly Met Asp Phe Ser Asn Val Val Val His
435 440 445
Glu Asp Ile Asp Gly Asp Gly Asp Val Lys Asp Asp Ile Val Gly Leu
450 455 460
Glu Gly Ala Ser Pro Lys Ser Met Pro Pro Ile Trp Ala Glu Val Met
465 470 475 480
Arg Phe Leu Thr Asn Pro His Lys Pro Met Ile Leu Ala Leu Ser Arg
485 490 495
Pro Asp Pro Lys Lys Asn Ile Thr Thr Leu Val Lys Ala Phe Gly Glu
500 505 510
Cys Arg Pro Leu Arg Glu Leu Ala Asn Leu Thr Leu Ile Met Gly Asn
515 520 525
Arg Asp Asp Ile Asp Asp Met Ser Ala Gly Asn Ala Ser Val Leu Thr
530 535 540
Thr Val Leu Lys Leu Ile Asp Lys Tyr Asp Leu Tyr Gly Ser Val Ala
545 550 555 560
Phe Pro Lys His His Asn Gln Ala Asp Val Pro Glu Ile Tyr Arg Leu
565 570 575
Ala Ala Lys Met Lys Gly Val Phe Ile Asn Pro Ala Leu Val Glu Pro
580 585 590
Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala His Gly Leu Pro Ile Val
595 600 605
Ala Thr Lys Asn Gly Gly Pro Val Asp Ile Thr Asn Ala Leu Asn Asn
610 615 620
Gly Leu Leu Val Asp Pro His Asp Gln Asn Ala Ile Ala Asp Ala Leu
625 630 635 640
Leu Lys Leu Val Ala Asp Lys Asn Leu Trp Gln Glu Cys Arg Arg Asn
645 650 655
Gly Leu Arg Asn Ile His Leu Tyr Ser Trp Pro Glu His Cys Arg Thr
660 665 670
Tyr Leu Thr Arg Val Ala Gly Cys Arg Leu Arg Asn Pro Arg Trp Leu
675 680 685
Lys Asp Thr Pro Ala Asp Ala Gly Ala Asp Glu Glu Glu Phe Leu Glu
690 695 700
Asp Ser Met Asp Ala Gln Asp Leu Ser Leu Arg Leu Ser Ile Asp Gly
705 710 715 720
Glu Lys Ser Ser Leu Asn Thr Asn Asp Pro Leu Trp Phe Asp Pro Gln
725 730 735
Asp Gln Val Gln Lys Ile Met Asn Asn Ile Lys Gln Ser Ser Ala Leu
740 745 750
Pro Pro Ser Met Ser Ser Val Ala Ala Glu Gly Thr Gly Ser Thr Met
755 760 765
Asn Lys Tyr Pro Leu Leu Arg Arg Arg Arg Arg Leu Phe Val Ile Ala
770 775 780
Val Asp Cys Tyr Gln Asp Asp Gly Arg Ala Ser Lys Lys Met Leu Gln
785 790 795 800
Val Ile Gln Glu Val Phe Arg Ala Val Arg Ser Asp Ser Gln Met Phe
805 810 815
Lys Ile Ser Gly Phe Thr Leu Ser Thr Ala Met Pro Leu Ser Glu Thr
820 825 830
Leu Gln Leu Leu Gln Leu Gly Lys Ile Pro Ala Thr Asp Phe Asp Ala
835 840 845
Leu Ile Cys Gly Ser Gly Ser Glu Val Tyr Tyr Pro Gly Thr Ala Asn
850 855 860
Cys Met Asp Ala Glu Gly Lys Leu Arg Pro Asp Gln Asp Tyr Leu Met
865 870 875 880
His Ile Ser His Arg Trp Ser His Asp Gly Ala Arg Gln Thr Ile Ala
885 890 895
Lys Leu Met Gly Ala Gln Asp Gly Ser Gly Asp Ala Val Glu Gln Asp
900 905 910
Val Ala Ser Ser Asn Ala His Cys Val Ala Phe Leu Ile Lys Asp Pro
915 920 925
Gln Lys Val Lys Thr Val Asp Glu Met Arg Glu Arg Leu Arg Met Arg
930 935 940
Gly Leu Arg Cys His Ile Met Tyr Cys Arg Asn Ser Thr Arg Leu Gln
945 950 955 960
Val Val Pro Leu Leu Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Ser
965 970 975
Val Arg Trp Gly Val Ser Val Gly Asn Met Tyr Leu Ile Thr Gly Glu
980 985 990
His Gly Asp Thr Asp Leu Glu Glu Met Leu Ser Gly Leu His Lys Thr
995 1000 1005
Val Ile Val Arg Gly Val Thr Glu Lys Gly Ser Glu Ala Leu Val Arg
1010 1015 1020
Ser Pro Gly Ser Tyr Lys Arg Asp Asp Val Val Pro Ser Glu Thr Pro
1025 1030 1035 1040
Leu Ala Ala Tyr Thr Thr Gly Glu Leu Lys Ala Asp Glu Ile Met Arg
1045 1050 1055
Ala Leu Lys Gln Val Ser Lys Thr Ser Ser Gly Met
1060 1065
<210>9
<211>1084
<212>PRT
<213>水稻(Oryza sativa)
<400>9
Met Ala Gly Asn Glu Trp Ile Asn Gly Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Ser Gly Gly Ala Ala Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly
20 25 30
Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly Gly Val Asp Pro
35 40 45
Ser Ser Pro Thr Thr Gly Thr Thr Ser Pro Arg Gly Pro His Met Asn
50 55 60
Phe Asn Pro Thr His Tyr Phe Val Glu Glu Val Val Lys Gly Val Asp
65 70 75 80
Glu Ser Asp Leu His Arg Thr Trp Ile Lys Val Val Ala Thr Arg Asn
85 90 95
Ala Arg Glu Arg Ser Thr Arg Leu Glu Asn Met Cys Trp Arg Ile Trp
100 105 110
His Leu Ala Arg Lys Lys Lys Gln Leu Glu Leu Glu Gly Ile Leu Arg
115 120 125
Ile Ser Ala Arg Arg Lys Glu Gln Glu Gln Val Arg Arg Glu Thr Ser
130 135 140
Glu Asp Leu Ala Glu Asp Leu Phe Glu Gly Glu Lys Ala Asp Thr Val
145 150 155 160
Gly Glu Leu Ala Gln Gln Asp Thr Pro Met Lys Lys Lys Phe Gln Arg
165 170 175
Asn Phe Ser Glu Leu Thr Val Ser Trp Ser Asp Glu Asn Lys Glu Lys
180 185 190
Lys Leu Tyr Ile Val Leu Ile Ser Leu His Gly Leu Val Arg Gly Asp
195 200 205
Asn Met Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr
210 215 220
Val Val Glu Leu Ala Arg Ala Leu Ala Met Met Pro Gly Val Tyr Arg
225 230 235 240
Val Asp Leu Phe Thr Arg Gln Val Ser Ser Pro Glu Val Asp Trp Ser
245 250 255
Tyr Gly Glu Pro Thr Glu Met Leu Thr Ser Gly Ser Thr Asp Gly Glu
260 265 270
Gly Ser Gly Glu Ser Ala Gly Ala Tyr Ile Val Arg Ile Pro Cys Gly
275 280 285
Pro Arg Asp Lys Tyr Leu Arg Lys Glu Ala Leu Trp Pro Tyr Leu Gln
290 295 300
Glu Phe Val Asp Gly Ala Leu Ala His Ile Leu Asn Met Ser Lys Ala
305 310 315 320
Leu Gly Glu Gln Val Ser Asn Gly Lys Leu Val Leu Pro Tyr Val Ile
325 330 335
His Gly His Tyr Ala Asp Ala Gly Asp Val Ala Ala Leu Leu Ser Gly
340 345 350
Ala Leu Asn Val Pro Met Val Leu Thr Gly His Ser Leu Gly Arg Asn
355 360 365
Lys Leu Glu Gln Ile Met Lys Gln Gly Arg Met Ser Lys Glu Glu Met
370 375 380
Asp Ser Thr Tyr Lys Ile Met Arg Arg Ile Glu Gly Glu Glu Leu Ala
385 390 395 400
Leu Asp Ala Ala Glu Leu Val Ile Thr Ser Thr Arg Gln Glu Ile Asp
405 410 415
Glu Gln Trp Gly Leu Tyr Asp Gly Phe Asp Val Lys Leu Glu Lys Val
420 425 430
Leu Arg Ala Arg Ala Arg Arg Gly Val Ser Cys His Gly Arg Phe Met
435 440 445
Pro Arg Met Val Val Ile Pro Pro Gly Met Asp Phe Ser Ser Val Val
450 455 460
Val Pro Glu Asp Thr Ser Asp Gly Asp Asp Gly Lys Asp Phe Glu Ile
465 470 475 480
Ala Ser Pro Arg Ser Leu Pro Pro Ile Trp Ala Glu Val Ser Arg Phe
485 490 495
Trp Thr Asn Pro His Lys Pro Met Ile Leu Ala Leu Ser Arg Pro Asp
500 505 510
Pro Lys Lys Asn Ile Thr Thr Leu Val Lys Ala Phe Gly Glu Cys Arg
515 520 525
Pro Leu Arg Glu Leu Ala Asn Leu Ile Leu Ser Met Gly Thr Arg Asp
530 535 540
Asp Ile Asp Gly Met Ser Ala Gly Asn Ala Ser Val Leu Thr Thr Val
545 550 555 560
Leu Lys Leu Ile Asp Lys Tyr Asp Leu Tyr Gly Ser Val Ala Phe Pro
565 570 575
Lys Tyr His Lys Gln Ser Asp Val Pro Glu Ile Tyr Arg Leu Thr Gly
580 585 590
Lys Met Lys Gly Val Phe Ile Asn Pro Ala Leu Val Glu Pro Phe Gly
595 600 605
Leu Thr Leu Ile Glu Ala Ala Ala His Gly Leu Pro Ile Val Gly Thr
610 615 620
Lys Asn Gly Gly Pro Val Asp Ile Lys Asn Ala Leu Asn Asn Gly Leu
625 630 635 640
Leu Val Asp Pro His Asp Gln His Ala Ile Ala Asp Ala Leu Leu Lys
645 650 655
Leu Val Ala Asp Lys Asn Leu Trp Gln Glu Cys Arg Lys Asn Gly Leu
660 665 670
Arg Asn Ile Gln Leu Tyr Ser Trp Pro Glu His Cys Arg Thr Tyr Leu
675 680 685
Thr Arg Ile Ala Gly Cys Arg Ile Arg Asn Pro Arg Trp Leu Met Asp
690 695 700
Thr Pro Ala Asp Ala Ala Ala Glu Glu Glu Glu Ala Leu Glu Asp Ser
705 710 715 720
Leu Met Asp Val Gln Asp Leu Ser Leu Arg Leu Ser Ile Asp Gly Glu
725 730 735
Arg Gly Ser Ser Met Asn Asp Ala Pro Ser Ser Asp Pro Gln Asp Ser
740 745 750
Val Gln Arg Ile Met Asn Lys Ile Lys Arg Ser Ser Pro Ala Glu Thr
755 760 765
Asp Gly Ala Lys Ile Pro Ala Glu Ala Ala Ala Thr Ala Thr Ser Gly
770 775 780
Ala Met Asn Lys Tyr Pro Leu Leu Arg Arg Arg Arg Arg Leu Phe Val
785 790 795 800
Ile Ala Val Asp Cys Tyr Gly Asp Asp Gly Ser Ala Ser Lys Arg Met
805 810 815
Leu Gln Val Ile Gln Glu Val Phe Arg Ala Val Arg Ser Asp Ser Gln
820 825 830
Met Ser Arg Ile Ser Gly Phe Ala Leu Ser Thr Xaa Met Pro Leu Pro
835 840 845
Glu Thr Leu Lys Leu Leu Gln Leu Gly Lys Ile Pro Pro Thr Asp Phe
850 855 860
Asp Ala Leu Ile Cys Gly Ser Gly Ser Glu Val Tyr Tyr Pro Ser Thr
865 870 875 880
Ala Gln Cys Val Asp Ala Gly Gly Arg Leu Arg Pro Asp Gln Asp Tyr
885 890 895
Leu Leu His Ile Asn His Arg Trp Ser His Asp Gly Ala Lys Gln Thr
900 905 910
Ile Ala Lys Leu Ala His Asp Gly Ser Gly Thr Asn Val Glu Pro Asp
915 920 925
Val Glu Ser Cys Asn Pro His Cys Val Ser Phe Phe Ile Lys Asp Pro
930 935 940
Asn Lys Val Arg Thr Met Asp Glu Met Arg Glu Arg Val Arg Met Arg
945 950 955 960
Gly Leu Arg Cys His Leu Met Tyr Cys Arg Asn Ala Thr Arg Leu Gln
965 970 975
Val Val Pro Leu Leu Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Phe
980 985 990
Val Arg Trp Gly Leu Ser Val Gly Asn Met Tyr Leu Ile Val Gly Glu
995 1000 1005
His Gly Asp Thr Asp His Glu Glu Met Leu Ser Gly Leu His Lys Thr
1010 1015 1020
Val Ile Ile Arg Gly Val Thr Glu Lys Gly Ser Glu Gln Leu Val Arg
1025 1030 1035 1040
Ser Ser Gly Ser Tyr Gln Arg Glu Asp Val Val Pro Ser Glu Ser Pro
1045 1050 1055
Leu Ile Ala Phe Thr Lys Gly Asp Leu Lys Ala Asp Glu Ile Met Arg
1060 1065 1070
Ala Leu Lys Glu Val Thr Lys Ala Ala Ser Gly Met
1075 1080
<210>10
<211>1049
<212>PRT
<213>水稻(Oryza sativa)
<400>10
Met Ala G1y Asn Glu Trp Ile Asn Gly Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Ser Gly Gly Ala Ala Gly Gly Gly Gly Gly Gly Gly Gly Val Asp Pro
20 25 30
Arg Ser Pro Ala Ala Gly Ala Ala Ser Pro Arg Gly Pro His Met Asn
35 40 45
Phe Asn Pro Thr His Tyr Phe Val Glu Glu Val Val Lys Gly Val Asp
50 55 60
Glu Ser Asp Leu His Arg Thr Trp Ile Lys Val Val Ala Thr Arg Asn
65 70 75 80
Ala Arg Glu Arg Ser Thr Arg Leu Glu Asn Met Cys Trp Arg Ile Trp
85 90 95
His Leu Ala Arg Lys Lys Lys Gln Leu Glu Leu Glu Gly Ile Leu Arg
100 105 110
Ile Ser Ala Arg Arg Lys Glu Gln Glu Gln Val Arg Arg Glu Thr Ser
115 120 125
Glu Asp Leu Ala Glu Asp Leu Phe Glu Gly Glu Lys Ala Asp Thr Val
130 135 140
Gly Glu Leu Ala Gln Gln Asp Thr Pro Met Lys Lys Lys Phe Gln Arg
145 150 155 160
Asn Phe Ser Glu Leu Thr Val Ser Trp Ser Asp Glu Asn Lys Glu Lys
165 170 175
Lys Leu Tyr Ile Val Leu Ile Ser Leu His Gly Leu Val Ser Gly Asp
180 185 190
Asn Met Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr
195 200 205
Val Val Glu Leu Ala Arg Ala Leu Ala Met Met Pro Gly Val Tyr Arg
210 215 220
Val Asp Leu Phe Thr Arg Gln Val Ser Ser Pro Glu Val Asp Trp Ser
225 230 235 240
Tyr Gly Glu Pro Thr Glu Met Leu Thr Pro Val Pro Leu Thr Glu Arg
245 250 255
Glu Ala Val Arg Val Leu Val Arg Thr Leu Cys Ala Phe Arg Ala Val
260 265 270
Gln Gly Thr Ser Thr Ser Val Lys Ser Pro Val Ala Leu Pro Pro Arg
275 280 285
Val Cys Arg Arg Ser Ser Arg Ala Tyr Leu Asn Met Ser Lys Ala Leu
290 295 300
Gly Glu Gln Val Ser Asn Gly Lys Leu Val Leu Pro Tyr Val Ile His
305 310 315 320
Gly His Tyr Ala Asp Ala Gly Asp Val Ala Ala Leu Leu Ser Gly Ala
325 330 335
Leu Asn Val Pro Met Val Leu Thr Gly His Ser Leu Gly Arg Asn Lys
340 345 350
Leu Glu Gln Ile Met Lys Gln Gly Arg Met Ser Lys Glu Glu Ile Asp
355 360 365
Ser Thr Tyr Lys Ile Met Arg Arg Ile Glu Gly Glu Glu Leu Ala Leu
370 375 380
Asp Ala Thr Glu Pro Val Ile Thr Ser Thr Arg Gln Glu Asn Asp Glu
385 390 395 400
Gln Trp Gly Leu Tyr Asp Gly Phe Asp Val Lys Leu Glu Lys Val Leu
405 410 415
Arg Ala Arg Ala Arg Arg Gly Val Ser Cys His Gly Arg Phe Met Pro
420 425 430
Arg Met Val Val Ile Pro Pro Gly Met Asp Phe Ser Ser Val Val Val
435 440 445
Pro Glu Asp Thr Ser Asp Gly Asp Asp Gly Lys Asp Phe Glu Ile Ala
450 455 460
Ser Pro Arg Ser Leu Pro Pro Ile Trp Ala Glu Val Met Arg Phe Leu
465 470 475 480
Thr Asn Pro His Lys Pro Met Ile Leu Ala Leu Ser Arg Pro Asp Pro
485 490 495
Lys Lys Asn Ile Thr Thr Leu Val Lys Ala Phe Gly Glu Cys Arg Pro
500 505 510
Leu Arg Glu Leu Ala Asn Leu Ile Leu Ile Met Gly Asn Arg Asp Asp
515 520 525
Ile Asp Glu Met Ser Ala Gly Asn Ala Ser Val Leu Thr Thr Val Leu
530 535 540
Lys Leu Ile Asp Lys Tyr Asp Leu Tyr Gly Ser Val Ala Phe Pro Lys
545 550 555 560
His His Lys Gln Ser Asp Val Pro Glu Ile Tyr Arg Leu Thr Gly Lys
565 570 575
Met Lys Gly Val Phe Ile Asn Pro Ala Leu Val Glu Pro Phe Gly Leu
580 585 590
Thr Leu Ile Glu Ala Ala Ala His Gly Leu Pro Ile Val Ala Thr Lys
595 600 605
Asn Gly Gly Pro Val Asp Ile Lys Asn Ala Leu Asn Asn Gly Leu Leu
610 615 620
Val Asp Pro His Asp Gln His Ala Ile Ala Asp Ala Leu Leu Lys Leu
625 630 635 640
Val Ala Asp Lys Asn Leu Trp Gln Glu Cys Arg Lys Asn Gly Leu Arg
645 650 655
Asn Ile Gln Leu Tyr Ser Trp Pro Glu His Cys Arg Thr Tyr Leu Thr
660 665 670
Arg Ile Ala Gly Cys Arg Ile Arg Asn Pro Arg Trp Leu Met Asp Thr
675 680 685
Pro Ala Asp Ala Ala Ala Glu Glu Glu Glu Ala Leu Glu Asp Ser Leu
690 695 700
Met Asp Val Gln Asp Leu Ser Leu His Leu Ser Ile Asp Gly Glu Arg
705 710 715 720
Gly Ser Ser Met Asn Asp Ala Pro Ser Ser Asp Pro Gln Asp Ser Val
725 730 735
Gln Arg Ile Met Asn Lys Ile Lys Arg Ser Ser Pro Ala Asp Thr Asp
740 745 750
Gly Ala Lys Ile Arg Gln Ala Ala Ala Thr Ala Thr Ser Gly Ala Met
755 760 765
Asn Lys Tyr Pro Leu Leu Arg Arg Arg Arg Arg Leu Phe Val Ile Ala
770 775 780
Val Asp Cys Tyr Gly Asp Asp Gly Ser Ala Ser Lys Arg Met Leu Gln
785 790 795 800
Val Ile Gln Glu Val Phe Arg Ala Val Arg Ser Asp Ser Gln Met Ser
805 810 815
Arg Ile Ser Gly Phe Ala Leu Ser Thr Ala Met Pro Leu Pro Glu Thr
820 825 830
Leu Lys Leu Leu Gln Leu Gly Lys Ile Pro Pro Thr Asp Phe Asp Ala
835 840 845
Leu Ile Cys Gly Ser Gly Ser Glu Val Tyr Tyr Pro Gly Thr Ala Gln
850 855 860
Cys Val Asp Ala Gly Gly Leu Arg Pro Asp Gln Asp Tyr Leu Leu His
865 870 875 880
Ile Asn His Arg Trp Ser His Asp Gly Ala Lys Gln Thr Ile Ala Asn
885 890 895
Val Ala His Asp Gly Ser Gly Thr Asn Val Glu Pro Asp Val Glu Ser
900 905 910
Cys Asn Pro His Cys Val Ser Phe Phe Ile Lys Asp Pro Asn Lys Val
915 920 925
Arg Thr Ala Asp Glu Met Arg Glu Arg Met Arg Met Arg Gly Leu Arg
930 935 940
Cys His Leu Met Tyr Cys Arg Asn Ala Thr Arg Leu Gln Val Val Pro
945 950 955 960
Leu Leu Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Phe Val Arg Trp
965 970 975
Gly Leu Ser Val Gly Asn Met Tyr Leu Ile Val Gly Glu His Gly Asp
980 985 990
Thr Asp His Glu Glu Met Leu Ser Gly Leu His Lys Thr Val Ile Ile
995 1000 1005
Arg Gly Val Thr Glu Lys Gly Ser Glu Gln Leu Val Arg Ser Ser Gly
1010 1015 1020
Ser Tyr Gln Arg Glu Asp Val Phe Pro Ser Glu Ser Pro Leu Ile Ala
1025 1030 1035 1040
Phe Thr Lys Gly Asp Leu Lys Ala Asp
1045
<210>11
<211>1083
<212>PRT
<213>拟南芥(Arabidopsis thaliana)
<400>11
Met Ala Arg Asn Asp Trp Ile Asn Ser Tyr Leu Glu Ala Ile Leu Asp
1 5 10 15
Val Gly Thr Ser Lys Lys Lys Arg Phe Glu Ser Asn Ser Lys Ile Val
20 25 30
Gln Lys Leu Gly Asp Ile Asn Ser Lys Asp His Gln Glu Lys Val Phe
35 40 45
Gly Asp Met Asn Gly Lys Asp His Gln Glu Lys Val Phe Ser Pro Ile
50 55 60
Lys Tyr Phe Val Glu Glu Val Val Asn Ser Phe Asp Glu Ser Asp Leu
65 70 75 80
Tyr Lys Thr Trp Ile Lys Val Ile Ala Thr Arg Asn Thr Arg Glu Arg
85 90 95
Ser Asn Arg Leu Glu Asn Ile Cys Trp Arg Ile Trp His Leu Ala Arg
100 105 110
Lys Lys Lys Gln Ile Val Trp Asp Asp Gly Val Arg Leu Ser Lys Arg
115 120 125
Arg Ile Glu Arg Glu Gln Gly Arg Asn Asp Ala Glu Glu Asp Leu Leu
130 135 140
Ser Glu Leu Ser Glu Gly Glu Lys Asp Lys Asn Asp Gly Glu Lys Glu
145 150 155 160
Lys Ser Glu Val Val Thr Thr Leu Glu Pro Pro Arg Asp His Met Pro
165 170 175
Arg Ile Arg Ser Glu Met Gln Ile Trp Ser Glu Asp Asp Lys Ser Ser
180 185 190
Arg Asn Leu Tyr Ile Val Leu Ile Arg Gln Val Glu Ile Gly Phe Ser
195 200 205
Asp Leu Phe Val Val Phe Asn Met Leu Val Gly Leu Thr Trp Cys Leu
210 215 220
Tyr Leu Val Pro Cys Phe Thr Asn Cys Ser Met His Gly Leu Val Arg
225 230 235 240
Gly Glu Asn Met Glu Leu Gly Arg Asp Ser Asp Thr Gly Gly Gln Val
245 250 255
Lys Tyr Val Val Glu Leu Ala Arg Ala Leu Ala Asn Thr Glu Gly Val
260 265 270
His Arg Val Asp Leu Leu Thr Arg Gln Ile Ser Ser Pro Glu Val Asp
275 280 285
Tyr Ser Tyr Gly Glu Pro Val Glu Met Leu Ser Cys Pro Pro Glu Gly
290 295 300
Ser Asp Ser Cys Gly Ser Tyr Ile Ile Arg Ile Pro Cys Gly Ser Arg
305 310 315 320
Asp Lys Tyr Ile Pro Lys Glu Ser Leu Trp Pro His Ile Pro Glu Phe
325 330 335
Val Asp Gly Ala Leu Asn His Ile Val Ser Ile Ala Arg Ser Leu Gly
340 345 350
Glu Gln Val Asn Gly Gly Lys Pro Ile Trp Pro Tyr Val Ile His Gly
355 360 365
His Tyr Ala Asp Ala Gly Glu Val Ala Ala His Leu Ala Gly Ala Leu
370 375 380
Asn Val Pro Met Val Leu Thr Gly His Ser Leu Gly Arg Asn Lys Phe
385 390 395 400
Glu Gln Leu Leu Gln Gln Gly Arg Ile Thr Arg Glu Asp Ile Asp Arg
405 410 415
Thr Tyr Lys Ile Met Arg Arg Ile Glu Ala Glu Glu Gln Ser Leu Asp
420 425 430
Ala Ala Glu Met Val Val Thr Ser Thr Arg Gln Glu Ile Asp Ala Gln
435 440 445
Trp Gly Leu Tyr Asp Gly Phe Asp Ile Lys Leu Glu Arg Lys Leu Arg
450 455 460
Val Arg Arg Arg Arg Gly Val Ser Cys Leu Gly Arg Tyr Met Pro Arg
465 470 475 480
Met Val Val Ile Pro Pro Gly Met Asp Phe Ser Tyr Val Leu Thr Gln
485 490 495
Asp Ser Gln Glu Pro Asp Gly Asp Leu Lys Ser Leu Ile Gly Pro Asp
500 505 510
Arg Asn Gln Ile Lys Lys Pro Val Pro Pro Ile Trp Ser Glu Ile Met
515 520 525
Arg Phe Phe Ser Asn Pro His Lys Pro Thr Ile Leu Ala Leu Ser Arg
530 535 540
Pro Asp His Lys Lys Asn Val Thr Thr Leu Val Lys Ala Phe Gly Glu
545 550 555 560
Cys Gln Pro Leu Arg Glu Leu Ala Asn Leu Val Leu Ile Leu Gly Asn
565 570 575
Arg Asp Asp Ile Glu Glu Met Pro Asn Ser Ser Ser Val Val Leu Met
580 585 590
Asn Val Leu Lys Leu Ile Asp Gln Tyr Asp Leu Tyr Gly Gln Val Ala
595 600 605
Tyr Pro Lys His His Lys Gln Ser Glu Val Pro Asp Ile Tyr Arg Leu
610 615 620
Ala Ala Lys Thr Lys Gly Val Phe Ile Asn Pro Ala Leu Val Glu Pro
625 630 635 640
Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala Tyr Gly Leu Pro Ile Val
645 650 655
Ala Thr Arg Asn Gly Gly Pro Val Asp Ile Val Lys Ala Leu Asn Asn
660 665 670
Gly Leu Leu Val Asp Pro His Asp Gln Gln Ala Ile Ser Asp Ala Leu
675 680 685
Leu Lys Leu Val Ala Asn Lys His Leu Trp Ala Glu Cys Arg Lys Asn
690 695 700
Gly Leu Lys Asn Ile His Arg Phe Ser Trp Pro Glu His Cys Arg Asn
705 710 715 720
Tyr Leu Ser His Val Glu His Cys Arg Asn Arg His Pro Thr Ser Ser
725 730 735
Leu Asp Ile Met Lys Val Pro Glu Glu Leu Thr Ser Asp Ser Leu Arg
740 745 750
Asp Val Asp Asp Ile Ser Leu Arg Phe Ser Thr Glu Gly Asp Phe Thr
755 760 765
Leu Asn Gly Glu Leu Asp Ala Gly Thr Arg Gln Lys Lys Leu Val Asp
770 775 780
Ala Ile Ser Gln Met Asn Ser Met Lys Gly Cys Ser Ala Ala Ile Tyr
785 790 795 800
Ser Pro Gly Arg Arg Gln Met Leu Phe Val Val Ala Val Asp Ser Tyr
805 810 815
Asp Asp Asn Gly Asn Ile Lys Ala Asn Leu Asn Glu Ile Ile Lys Asn
820 825 830
Met Ile Lys Ala Ala Asp Leu Thr Ser Gly Lys Gly Lys Ile Gly Phe
835 840 845
Val Leu Ala Ser Gly Ser Ser Leu Gln Glu Val Val Asp Ile Thr Gln
850 855 860
Lys Asn Leu Ile Asn Leu Glu Asp Phe Asp Ala Ile Val Cys Asn Ser
865 870 875 880
Gly Ser Glu Ile Tyr Tyr Pro Trp Arg Asp Met Met Val Asp Ala Asp
885 890 895
Tyr Glu Thr His Val Glu Tyr Lys Trp Pro Gly Glu Ser Ile Arg Ser
900 905 910
Val Ile Leu Arg Leu Ile Cys Thr Glu Pro Ala Ala Glu Asp Asp Ile
915 920 925
Thr Glu Tyr Ala Ser Ser Cys Ser Thr Arg Cys Tyr Ala Ile Ser Val
930 935 940
Lys Gln Gly Val Lys Thr Arg Arg Val Asp Asp Leu Arg Gln Arg Leu
945 950 955 960
Arg Met Arg Gly Leu Arg Cys Asn Ile Val Tyr Thr His Ala Ala Thr
965 970 975
Arg Leu Asn Val Ile Pro Leu Cys Ala Ser Arg Ile Gln Ala Leu Arg
980 985 990
Tyr Leu Ser Ile Arg Trp Gly Ile Asp Met Ser Lys Thr Val Phe Phe
995 1000 1005
Leu Gly Glu Lys Gly Asp Thr Asp Tyr Glu Asp Leu Leu Gly Gly Leu
1010 1015 1020
His Lys Thr Ile Ile Leu Lys Gly Val Val Gly Ser Asp Ser Glu Lys
1025 1030 1035 1040
Leu Leu Arg Ser Glu Glu Asn Phe Lys Arg Glu Asp Ala Val Pro Gln
1045 1050 1055
Glu Ser Pro Asn Ile Ser Tyr Val Lys Glu Asn Gly Gly Ser Gln Glu
1060 1065 1070
Ile Met Ser Thr Leu Glu Ala Tyr Gly Ile Lys
1075 1080
<210>12
<211>963
<212>PRT
<213>拟南芥(Arabidopsis thaliana)
<400>12
Met Ala Gly Asn Asp Asn Trp Ile Asn Ser Tyr Leu Asp Gly Ile Leu
1 5 10 15
Asp Ala Gly Lys Ala Ala Ile Gly Gly Asn Arg Pro Ser Leu Leu Leu
20 25 30
Arg Glu Arg Gly His Phe Ser Pro Ala Arg Tyr Phe Val Glu Glu Val
35 40 45
Ile Thr Gly Tyr Asp Glu Thr Asp Leu Tyr Lys Thr Trp Leu Arg Ala
50 55 60
Asu Ala Met Arg Ser Arg Arg Glu Glu His Ala Leu Glu Asn Met Thr
65 70 75 80
Trp Arg Ile Trp Asn Leu Ala Arg Lys Lys Lys Glu Phe Glu Lys Glu
85 90 95
Glu Ala Cys Arg Leu Ser Lys Arg Gln Pro Glu Thr Glu Lys Thr Arg
100 105 110
Ala Asp Ala Thr Ala Asp Met Ser Glu Asp Leu Phe Glu Gly Glu Lys
115 120 125
Gly Glu Asp Ala Gly Asp Pro Ser Val Ala Tyr Gly Asp Ser Thr Thr
130 135 140
Gly Ser Ser Pro Lys Thr Ser Ser Ile Asp Lys Leu Tyr Ile Val Leu
145 150 155 160
Ile Ser Leu His Gly Leu Val Arg Gly Glu Asn Met Glu Leu Gly Arg
165 170 175
Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr Val Val Glu Leu Ala Lys
180 185 190
Ala Leu Ser Ser Ser Pro Gly Val Tyr Arg Val Asp Leu Leu Thr Arg
195 200 205
Gln Ile Leu Ala Pro Asn Phe Asp Arg Ser Tyr Gly Glu Pro Ala Glu
210 215 220
Leu Leu Val Ser Thr Ser Gly Lys Asn Ser Lys Gln Glu Lys Gly Glu
225 230 235 240
Asn Ser Gly Ala Tyr Ile Ile Arg Ile Pro Phe Gly Pro Lys Asp Lys
245 250 255
Tyr Leu Ala Lys Glu His Leu Trp Pro Phe Ile Gln Glu Phe Val Asp
260 265 270
Gly Ala Leu Ser His Ile Val Arg Met Ser Lys Ala Ile Gly Glu Glu
275 280 285
Thr Gly Arg Gly His Pro Val Trp Pro Ser Val Ile His Gly His Tyr
290 295 300
Ala Ser Ala Gly Ile Ala Ala Ala Leu Leu Leu Gly Ala Leu Asn Leu
305 310 315 320
Pro Met Ala Phe Thr Gly His Phe Leu Gly Lys Asp Lys Leu Glu Gly
325 330 335
Leu Leu Lys Gln Gly Arg Gln Thr Arg Glu Gln Ile Asn Met Thr Tyr
340 345 350
Lys Ile Met Cys Arg Ile Glu Ala Glu Glu Leu Ser Leu Asp Ala Ser
355 360 365
Glu Ile Val Ile Ala Ser Thr Arg Gln Glu Ile Glu Glu Gln Trp Asn
370 375 380
Leu Tyr Asp Gly Phe Glu Val Ile Leu Ala Arg Lys Leu Arg Ala Arg
385 390 395 400
Val Lys Arg Gly Ala Asn Cys Tyr Gly Arg Phe Met Pro Arg Met Val
405 410 415
Ile Ile Pro Pro Gly Val Glu Phe Gly His Ile Ile His Asp Phe Asp
420 425 430
Met Asp Gly Glu Glu Glu Asn Pro Ser Pro Ala Ser Glu Asp Pro Pro
435 440 445
Ile Trp Ser Gln Ile Met Arg Phe Phe Thr Asn Pro Arg Lys Pro Met
450 455 460
Ile Leu Ala Val Ala Arg Pro Tyr Pro Glu Lys Asn Ile Thr Thr Leu
465 470 475 480
Val Lys Ala Phe Gly Glu Cys Arg Pro Leu Arg Glu Leu Ala Asn Leu
485 490 495
Thr Leu Ile Met Gly Asn Arg Glu Ala Ile Ser Lys Met His Asn Met
500 505 510
Ser Ala Ala Val Leu Thr Ser Val Leu Thr Leu Ile Asp Glu Tyr Asp
515 520 525
Leu Tyr Gly Gln Val Ala Tyr Pro Lys His His Lys His Ser Glu Val
530 535 540
Pro Asp Ile Tyr Arg Leu Ala Ala Arg Thr Lys Gly Ala Phe Val Asn
545 550 555 560
Val Ala Tyr Phe Glu Gln Phe Gly Val Thr Leu Ile Glu Ala Ala Met
565 570 575
Asn Gly Leu Pro Ile Ile Ala Thr Lys Asn Gly Ala Pro Val Glu Ile
580 585 590
Asn Gln Val Leu Asn Asn Gly Leu Leu Val Asp Pro His Asp Gln Asn
595 600 605
Ala Ile Ala Asp Ala Leu Tyr Lys Leu Leu Ser Asp Lys Gln Leu Trp
610 615 620
Ser Arg Cys Arg Glu Asn Gly Leu Thr Asn Ile His Gln Phe Ser Trp
625 630 635 640
Pro Glu His Cys Lys Asn Tyr Leu Ser Arg Ile Leu Thr Leu Gly Pro
645 650 655
Arg Ser Pro Ala Ile Gly Asn Arg Glu Glu Arg Ser Asn Thr Pro Ile
660 665 670
Ser Gly Arg Arg Gln Ile Ile Val Ile Ser Val Asp Ser Val Asn Lys
675 680 685
Glu Asp Leu Val Arg Ile Ile Arg Asn Ala Ile Glu Val Ile His Thr
690 695 700
Gln Asn Met Ser Gly Ser Ala Gly Phe Val Leu Ser Thr Ser Leu Thr
705 710 715 720
Ile Ser Glu Ile His Ser Leu Leu Leu Ser Gly Gly Met Leu Pro Thr
725 730 735
Asp Phe Asp Ala Phe Ile Cys Asn Ser Gly Ser Asn Ile Tyr Tyr Pro
740 745 750
Ser Tyr Ser Gly Glu Thr Pro Asn Asn Ser Lys Ile Thr Phe Ala Leu
755 760 765
Asp Gln Asn His Gln Ser His Ile Glu Tyr Arg Trp Gly Gly Glu Gly
770 775 780
Leu Arg Lys Tyr Leu Val Lys Trp Ala Thr Ser Val Val Glu Arg Lys
785 790 795 800
Gly Arg Thr Glu Arg Gln Ile Ile Phe Glu Asp Pro Glu His Ser Ser
805 810 815
Ala Tyr Cys Leu Ala Phe Arg Val Val Asn Pro Asn His Leu Pro Pro
820 825 830
Leu Lys Glu Leu Arg Lys Leu Met Arg Ile Gln Ser Leu Arg Cys Asn
835 840 845
Ala Leu Tyr Asn His Ser Ala Thr Arg Leu Ser Val Val Pro Ile His
850 855 860
Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Cys Ile Arg Trp Gly Ile
865 870 875 880
Glu Val Pro Asn Val Ala Val Leu Val Gly Glu Ser Gly Asp Ser Asp
885 890 895
Tyr Glu Glu Leu Leu Gly Gly Leu His Arg Thr Val Ile Leu Lys Gly
900 905 910
Glu Phe Asn Thr Pro Ala Asn Arg Ile His Thr Val Arg Arg Tyr Pro
915 920 925
Leu Gln Asp Val Val Pro Leu Asp Ser Ser Asn Ile Thr Gly Val Glu
930 935 940
Gly Tyr Thr Thr Asp Asp Leu Lys Ser Ala Leu Gln Gln Met Gly Ile
945 950 955 960
Leu Thr Gln
<210>13
<211>963
<212>PRT
<213>甘蔗(Saccharum officinarum)
<400>13
Met Ala Gly Asn Asp Asn Trp Ile Asn Ser Tyr Leu Asp Gly Ile Leu
1 5 10 15
Asp Ala Gly Lys Ala Ala Ile Gly Gly Asn Arg Pro Ser Leu Leu Leu
20 25 30
Arg Glu Arg Gly His Phe Ser Pro Ala Arg Tyr Phe Val Glu Glu Val
35 40 45
Ile Thr Gly Tyr Asp Glu Thr Asp Leu Tyr Lys Thr Trp Leu Arg Ala
50 55 60
Asn Ala Met Arg Ser Arg Arg Glu Glu His Ala Leu Glu Asn Met Thr
65 70 75 80
Trp Arg Ile Trp Asn Leu Ala Arg Lys Lys Lys Glu Phe Glu Lys Glu
85 90 95
Glu Ala Cys Arg Leu Ser Lys Arg Gln Pro Glu Thr Glu Lys Thr Arg
100 105 110
Ala Asp Ala Thr Ala Asp Met Ser Glu Asp Leu Phe Glu Gly Glu Lys
115 120 125
Gly Glu Asp Ala Gly Asp Pro Ser Val Ala Tyr Gly Asp Ser Thr Thr
130 135 140
Gly Ser Ser Pro Lys Thr Ser Ser Ile Asp Lys Leu Tyr Ile Val Leu
145 150 155 160
Ile Ser Leu His Gly Leu Val Arg Gly Glu Asn Met Glu Leu Gly Arg
165 170 175
Asp Ser Asp Thr Gly Gly Gln Val Lys Tyr Val Val Glu Leu Ala Lys
180 185 190
Ala Leu Ser Ser Ser Pro Gly Val Tyr Arg Val Asp Leu Leu Thr Arg
195 200 205
Gln Ile Leu Ala Pro Asn Phe Asp Arg Ser Tyr Gly Glu Pro Ala Glu
210 215 220
Leu Leu Val Ser Thr Ser Gly Lys Asn Ser Lys Gln Glu Lys Gly Glu
225 230 235 240
Asn Ser Gly Ala Tyr Ile Ile Arg Ile Pro Phe Gly Pro Lys Asp Lys
245 250 255
Tyr Leu Ala Lys Glu His Leu Trp Pro Phe Ile Gln Glu Phe Val Asp
260 265 270
Gly Ala Leu Ser His Ile Val Arg Met Ser Lys Ala Ile Gly Glu Glu
275 280 285
Thr Gly Arg Gly His Pro Val Trp Pro Ser Val Ile His Gly His Tyr
290 295 300
Ala Ser Ala Gly Ile Ala Ala Ala Leu Leu Leu Gly Ala Leu Asn Leu
305 310 315 320
Pro Met Ala Phe Thr Gly His Phe Leu Gly Lys Asp Lys Leu Glu Gly
325 330 335
Leu Leu Lys Gln Gly Arg Gln Thr Arg Glu Gln Ile Asn Met Thr Tyr
340 345 350
Lys Ile Met Cys Arg Ile Glu Ala Glu Glu Leu Ser Leu Asp Ala Ser
355 360 365
Glu Ile Val Ile Ala Ser Thr Arg Gln Glu Ile Glu Glu Gln Trp Asn
370 375 380
Leu Tyr Asp Gly Phe Glu Val Ile Leu Ala Arg Lys Leu Arg Ala Arg
385 390 395 400
Val Lys Arg Gly Ala Asn Cys Tyr Gly Arg Phe Met Pro Arg Met Val
405 410 415
Ile Ile Pro Pro Gly Val Glu Phe Gly His Ile Ile His Asp Phe Asp
420 425 430
Met Asp Gly Glu Glu Glu Asn Pro Ser Pro Ala Ser Glu Asp Pro Pro
435 440 445
Ile Trp Ser Gln Ile Met Arg Phe Phe Thr Asn Pro Arg Lys Pro Met
450 455 460
Ile Leu Ala Val Ala Arg Pro Tyr Pro Glu Lys Asn Ile Thr Thr Leu
465 470 475 480
Val Lys Ala Phe Gly Glu Cys Arg Pro Leu Arg Glu Leu Ala Asn Leu
485 490 495
Thr Leu Ile Met Gly Asn Arg Glu Ala Ile Ser Lys Met His Asn Met
500 505 510
Ser Ala Ala Val Leu Thr Ser Val Leu Thr Leu Ile Asp Glu Tyr Asp
515 520 525
Leu Tyr Gly Gln Val Ala Tyr Pro Lys His His Lys His Ser Glu Val
530 535 540
Pro Asp Ile Tyr Arg Leu Ala Ala Arg Thr Lys Gly Ala Phe Val Asn
545 550 555 560
Val Ala Tyr Phe Glu Gln Phe Gly Val Thr Leu Ile Glu Ala Ala Met
565 570 575
Asn Gly Leu Pro Ile Ile Ala Thr Lys Asn Gly Ala Pro Val Glu Ile
580 585 590
Asn Gln Val Leu Asn Asn Gly Leu Leu Val Asp Pro His Asp Gln Asn
595 600 605
Ala Ile Ala Asp Ala Leu Tyr Lys Leu Leu Ser Asp Lys Gln Leu Trp
610 615 620
Ser Arg Cys Arg Glu Asn Gly Leu Thr Asn Ile His Gln Phe Ser Trp
625 630 635 640
Pro Glu His Cys Lys Asn Tyr Leu Ser Arg Ile Leu Thr Leu Gly Pro
645 650 655
Arg Ser Pro Ala Ile Gly Asn Arg Glu Glu Arg Ser Asn Thr Pro Ile
660 665 670
Ser Gly Arg Arg Gln Ile Ile Val Ile Ser Val Asp Ser Val Asn Lys
675 680 685
Glu Asp Leu Val Arg Ile Ile Arg Asn Ala Ile Glu Val Ile His Thr
690 695 700
Gln Asn Met Ser Gly Ser Ala Gly Phe Val Leu Ser Thr Ser Leu Thr
705 710 715 720
Ile Ser Glu Ile His Ser Leu Leu Leu Ser Gly Gly Met Leu Pro Thr
725 730 735
Asp Phe Asp Ala Phe Ile Cys Asn Ser Gly Ser Asn Ile Tyr Tyr Pro
740 745 750
Ser Tyr Ser Gly Glu Thr Pro Asn Asn Ser Lys Ile Thr Phe Ala Leu
755 760 765
Asp Gln Asn His Gln Ser His Ile Glu Tyr Arg Trp Gly Gly Glu Gly
770 775 780
Leu Arg Lys Tyr Leu Val Lys Trp Ala Thr Ser Val Val Glu Arg Lys
785 790 795 800
Gly Arg Thr Glu Arg Gln Ile Ile Phe Glu Asp Pro Glu His Ser Ser
805 810 815
Ala Tyr Cys Leu Ala Phe Arg Val Val Asn Pro Asn His Leu Pro Pro
820 825 830
Leu Lys Glu Leu Arg Lys Leu Met Arg Ile Gln Ser Leu Arg Cys Asn
835 840 845
Ala Leu Tyr Asn His Ser Ala Thr Arg Leu Ser Val Val Pro Ile His
850 855 860
Ala Ser Arg Ser Gln Ala Leu Arg Tyr Leu Cys Ile Arg Trp Gly Ile
865 870 875 880
Glu Val Pro Asn Val Ala Val Leu Val Gly Glu Ser Gly Asp Ser Asp
885 890 895
Tyr Glu Glu Leu Leu Gly Gly Leu His Arg Thr Val Ile Leu Lys Gly
900 905 910
Glu Phe Asn Thr Pro Ala Asn Arg Ile His Thr Val Arg Arg Tyr Pro
915 920 925
Leu Gln Asp Val Val Pro Leu Asp Ser Ser Asn Ile Thr Gly Val Glu
930 935 940
Gly Tyr Thr Thr Asp Asp Leu Lys Ser Ala Leu Gln Gln Met Gly Ile
945 950 955 960
Leu Thr Gln
<210>14
<211>720
<212>PRT
<213>集胞藻(Synechocystis sp.)
<400>14
Met Ser Tyr Ser Ser Lys Tyr Ile Leu Leu Ile Ser Val His Gly Leu
1 5 10 15
Ile Arg Gly Glu Asn Leu Glu Leu Gly Arg Asp Ala Asp Thr Gly Gly
20 25 30
Gln Thr Lys Tyr Val Leu Glu Leu Ala Arg Ala Leu Val Lys Asn Pro
35 40 45
Gln Val Ala Arg Val Asp Leu Leu Thr Arg Leu Ile Lys Asp Pro Lys
50 55 60
Val Asp Ala Asp Tyr Ala Gln Pro Arg Glu Leu Ile Gly Asp Arg Ala
65 70 75 80
Gln Ile Val Arg Ile Glu Cys Gly Pro Glu Glu Tyr Ile Ala Lys Glu
85 90 95
Met Leu Trp Asp Tyr Leu Asp Asn Phe Ala Asp His Ala Leu Asp Tyr
100 105 110
Leu Lys Glu Gln Pro Glu Leu Pro Asp Val Ile His Ser His Tyr Ala
115 120 125
Asp Ala Gly Tyr Val Gly Thr Arg Leu Ser His Gln Leu Gly Ile Pro
130 135 140
Leu Val His Thr Gly His Ser Leu Gly Arg Ser Lys Arg Thr Arg Leu
145 150 155 160
Leu Leu Ser Gly Ile Lys Ala Asp Glu Ile Glu Ser Arg Tyr Asn Met
165 170 175
Ala Arg Arg Ile Asn Ala Glu Glu Glu Thr Leu Gly Ser Ala Ala Arg
180 185 190
Val Ile Thr Ser Thr His Gln Glu Ile Ala Glu Gln Tyr Ala Gln Tyr
195 200 205
Asp Tyr Tyr Gln Pro Asp Gln Met Leu Val Ile Pro Pro Gly Thr Asp
210 215 220
Leu Glu Lys Phe Tyr Pro Pro Lys Gly Asn Glu Trp Glu Thr Pro Ile
225 230 235 240
Val Gln Glu Leu Gln Arg Phe Leu Arg His Pro Arg Lys Pro Ile Ile
245 250 255
Leu Ala Leu Ser Arg Pro Asp Pro Arg Lys Asn Ile His Lys Leu Ile
260 265 270
Ala Ala Tyr Gly Gln Ser Pro Gln Leu Gln Ala Gln Ala Asn Leu Val
275 280 285
Ile Val Ala Gly Asn Arg Asp Asp Ile Thr Asp Leu Asp Gln Gly Pro
290 295 300
Arg Glu Val Leu Thr Asp Leu Leu Leu Thr Ile Asp Arg Tyr Asp Leu
305 310 315 320
Tyr Gly Lys Val Ala Tyr Pro Lys Gln Asn Gln Ala Glu Asp Val Tyr
325 330 335
Ala Leu Phe Arg Leu Thr Ala Leu Ser Gln Gly Val Phe Ile Asn Pro
340 345 350
Ala Leu Thr Glu Pro Phe Gly Leu Thr Leu Ile Glu Ala Ala Ala Cys
355 360 365
Gly Val Pro Ile Val Ala Thr Glu Asp Gly Gly Pro Val Asp Ile Ile
370 375 380
Lys Asn Cys Gln Asn Gly Tyr Leu Ile Asn Pro Leu Asp Glu Val Asp
385 390 395 400
Ile Ala Asp Lys Leu Leu Lys Val Leu Asn Asp Lys Gln Gln Trp Gln
405 410 415
Phe Leu Ser Glu Ser Gly Leu Glu Gly Val Lys Arg His Tyr Ser Trp
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Pro Ser His Val Glu Ser Tyr Leu Glu Ala Ile Asn Ala Leu Thr Gln
435 440 445
Gln Thr Ser Val Leu Lys Arg Ser Asp Leu Lys Arg Arg Arg Thr Leu
450 455 460
Tyr Tyr Asn Gly Ala Leu Val Thr Ser Leu Asp Gln Asn Leu Leu Gly
465 470 475 480
Ala Leu Gln Gly Gly Leu Pro Gly Asp Arg Gln Thr Leu Asp Glu Leu
485 490 495
Leu Glu Val Leu Tyr Gln His Arg Lys Asn Val Gly Phe Cys Ile Ala
500 505 510
Thr Gly Arg Arg Leu Asp Ser Val Leu Lys Ile Leu Arg Glu Tyr Arg
515 520 525
Ile Pro Gln Pro Asp Met Leu Ile Thr Ser Met Gly Thr Glu Ile Tyr
530 535 540
Ser Ser Pro Asp Leu Ile Pro Asp Gln Ser Trp Arg Asn His Ile Asp
545 550 555 560
Tyr Leu Trp Asn Arg Asn Ala Ile Val Arg Ile Leu Gly Glu Leu Pro
565 570 575
Gly Leu Ala Leu Gln Pro Lys Glu Glu Leu Ser Ala Tyr Lys Ile Ser
580 585 590
Tyr Phe Tyr Asp Ala Ala Ile Ala Pro Asn Leu Glu Glu Ile Arg Gln
595 600 605
Leu Leu His Lys Gly Glu Gln Thr Val Asn Thr Ile Ile Ser Phe Gly
610 615 620
Gln Phe Leu Asp Ile Leu Pro Ile Arg Ala Ser Lys Gly Tyr Ala Val
625 630 635 640
Arg Trp Leu Ser Gln Gln Trp Asn Ile Pro Leu Glu His Val Phe Thr
645 650 655
Ala Gly Gly Ser Gly Ala Asp Glu Asp Met Met Arg Gly Asn Thr Leu
660 665 670
Ser Val Val Val Ala Asn Arg His His Glu Glu Leu Ser Asn Leu Gly
675 680 685
Glu Ile Glu Pro Ile Tyr Phe Ser Glu Lys Arg Tyr Ala Ala Gly Ile
690 695 700
Leu Asp Gly Leu Ala His Tyr Arg Phe Phe Glu Leu Leu Asp Pro Val
705 710 715 720
Claims (7)
1.一种提高由棉花植物所产生的棉纤维的品质的方法,所述方法包括:
将一种嵌合DNA结构导入棉花植物,该DNA结构包含植物专一性转录起始区和编码蔗糖磷酸酯合成酶的DNA,所述蔗糖磷酸酯合成酶包含选自下列的氨基酸序列:SEQ ID NO:1、SEQ ID NO:2、SEQ ID NO:3、SEQ ID NO:4、SEQ ID NO:5、SEQ ID NO:6、SEQ ID NO:7、SEQ ID NO:8、SEQ ID NO:9、SEQ ID NO:10、SEQ ID NO:11、SEQ ID NO:12、SEQID NO:13、SEQ ID NO:14;
其中被提高的品质选自:与缺少嵌合DNA结构的棉花植物相比,提高了强度,提高了长度、提高了纤维成色、提高了纤维成熟率、降低了未成熟纤维含量、提高了纤维均匀度、提高了纤维细度。
2.如权利要求1的方法,还包括:
种植所述棉花植物。
3.如权利要求1的方法,其中,所述嵌合DNA结构能表达包含SEQID NO:1的氨基酸序列的蔗糖磷酸酯合成酶。
4.如权利要求1的方法,其中,所述转录起始区是棉纤维专一性的。
5.如权利要求1的方法,其中,所述转录起始区是35S启动子、复式S35启动子、FMV启动子、选自E6、H6、FbL2A、rac、CelA、CAP、ACP和LTP的棉纤维表达基因的棉纤维强化启动子。
6.如权利要求1的方法,其中,所述嵌合DNA结构被稳定整合到所述棉花植物的基因组中。
7.如权利要求1的方法,其中,所述将嵌合DNA结构导入所述植物是通过选自下列一组的方法完成的:电穿孔、农杆菌属介导的转化、粒子轰击基因转化、化学介导的转化和显微注射。
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US09/394,272 US6472588B1 (en) | 1999-09-10 | 1999-09-10 | Transgenic cotton plants with altered fiber characteristics transformed with a sucrose phosphate synthase nucleic acid |
US09/394,272 | 1999-09-10 |
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CN2006100037355A Expired - Fee Related CN101092632B (zh) | 1999-09-10 | 2000-09-07 | 增强了蔗糖磷酸酯合成酶表达的转基因产纤维植物 |
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EP1220602A1 (en) | 2002-07-10 |
ZA200201855B (en) | 2002-12-24 |
US6472588B1 (en) | 2002-10-29 |
TR200201477T2 (tr) | 2002-11-21 |
ATE337701T1 (de) | 2006-09-15 |
CN101092632B (zh) | 2011-01-26 |
AU7119000A (en) | 2001-04-10 |
US20030070191A1 (en) | 2003-04-10 |
US7091400B2 (en) | 2006-08-15 |
AU784405B2 (en) | 2006-03-30 |
EP1220602B1 (en) | 2006-08-30 |
MXPA02002497A (es) | 2002-07-30 |
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