CN116157009A - 大幅增加稻米产量的方式 - Google Patents
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- CN116157009A CN116157009A CN202180047035.9A CN202180047035A CN116157009A CN 116157009 A CN116157009 A CN 116157009A CN 202180047035 A CN202180047035 A CN 202180047035A CN 116157009 A CN116157009 A CN 116157009A
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Abstract
一种基因转殖稻米植物在其基因组中含有重组DNA构筑体,该构筑体包含第一核酸,该第一核酸具有与其天然启动子及5’非转译区(5’untranslated region,5’UTR)可操作地连接的第一Golden 2‑like转录因子(GLK)基因的序列,以及第二核酸,该第二核酸具有与其天然启动子及5’UTR可操作地连接的第二GLK基因的序列,该第二GLK基因不同于该第一GLK基因。该第一GLK基因以及该第二GLK基因均来自C4植物,且相较于未经转形的野生型稻米植物,该基因转殖稻米植物在枝条生物量上增加65‑106%且在谷物产量上增加50‑95%。还提供一种生产基因转殖稻米植物的方法以及一种可用于该方法的重组DNA构筑体。
Description
背景技术
不断扩大的人口需要增加作物产量。为了满足这一需求,近期致力于提高稻米等作物的光合作用效率上。自1960年代初至2000年,一系列目的在于提高全球农业产量的举措,亦即所谓的绿色革命,造成稻米产量大约倍增。然而,近年来产量的提升则较不引人注意。
由于超过90%的作物生物量来自光合作用,因此增加稻米等主要作物的光合作用可能会显著提高产量。
叶绿体在光合作用中发挥重要作用,也是调节植物生长、发育,以及逆境耐受性的植物激素生合成的场所。已知一对Golden 2-like(GLK)转录因子基因可调节高等植物如玉米(一种C4型植物)以及稻米(一种C3型植物)中叶绿体的发育。通过演化,相较于稻米的GLK基因,来自玉米的GLK基因,特别是GLK2,可能获得更新、更强的功能。
由于玉米的光合作用比稻米更有效,将两个玉米的GLK基因转形至稻米的基因组中可能会增强基因转殖稻米植物的光合作用,进而提高稻米的产量。
最近有报导称,转形由玉米泛素启动子所控制的两个玉米GLK基因中的任何一个,特别是玉米GLK2基因,稻米的产量提高了16-40%。然而,使用玉米泛素启动子,一种强组成型启动子,来驱动玉米GLK基因在基因转殖稻米中的表现导致稻米种子变小。
为了满足人口增长的需求,提高稻米产量仍是强烈的需求。
发明内容
为了满足不断增长的人口对粮食需求的日益增长,提供一种基因转殖稻米植物,其在其基因组中包含重组DNA构筑体,该构筑体包含第一核酸,该第一核酸具有与其天然启动子及5’非转译区(5’untranslated region,5’UTR)可操作地连接的第一Golden 2-like转录因子(GLK)基因的序列,以及第二核酸,该第二核酸具有与其天然启动子及5’UTR可操作地连接的第二GLK基因的序列,该第二GLK基因不同于该第一GLK基因。该第一GLK基因以及该第二GLK基因都是异源的,亦即并非来自稻米,且相较于一未经转形的野生型稻米植物,该基因转殖稻米植物在枝条生物量上(65-106%)及在谷物产量上(50-95%)具有显著,亦即至少50%的增加。
还提供一种生产上述基因转殖稻米植物的方法。
进一步公开一种可用于该方法的重组DNA构筑体。该重组DNA构筑体包含第一核酸序列,该第一核酸序列包含第一GLK基因,该第一GLK基因与其天然启动子及5’UTR可操作地连接,以及第二核酸序列,该第二核酸序列包含第二GLK基因,该第二GLK基因与其天然启动子及5’UTR可操作地连接,该第二GLK基因不同于该第一GLK基因。该第一GLK基因以及该第二GLK基因来自C4植物,例如玉米。
本发明的数个具体实施例的细节于以下描述及附图中阐述。本发明的所有特征、目的,及优点将从该描述及附图,以及从所附权利要求书中显而易见。
附图说明
以下描述参照附图,其中:
图1为用于农杆菌媒介的稻米转形的五种玉米GLK构筑体的示意图,以比较它们对基因表现及植物生长与产量的影响。pUbi:泛素启动子;p35S:35S启动子;Zm:Zea mays(玉米);pZmG1::ZmG1:玉米GLK1构筑体,具有其自身的启动子及5’UTR;pZmG2::ZmG2:玉米GLK2构筑体,具有其自身的启动子及5’UTR;HygR:潮霉素抗性基因;ZmG1:玉米GLK1基因;ZmG2:玉米GLK2基因;35ST:35S终止子;NOST:胭脂碱合成酶(nopaline synthase,nos)终止子;pro:启动子;LB:左边框;以及RB:右边框。
图2A所示为不同玉米组织中ZmG1(左)以及ZmG2(右)的相对基因表现量图,是通过分离自9天大幼苗的幼叶(YL)、2个月大植物的成熟叶(ML),以及幼茎(S)、雄花序(T),以及根(R)的总RNA进行定量即时聚合酶连锁反应(quantitative real-time polymerasechain reaction,“qRT-PCR”)来测定。数值为平均值±三个重复的标准差(S.D.),并标准化为17S rRNA的表现量。
图2B所示为ZmG1(左)以及ZmG2(右)的相对基因表现量的图,是通过来自野生型(WT)以及基因转殖稻米植物的总RNA进行qRT-PCR来确定。基因转殖稻米植物带有如图1所标记的转基因构筑体。从成熟的叶子(L)、茎(S)、小花(F),以及根(R)中分离出总RNA。异基因型组合的pZmG2::ZmG2植物以及其他四种基因转殖植物的同基因型组合的植物如图所示;由于高基因表现量,该同基因型组合的pZmG2::ZmG2植物在培养两个月后死亡。数值为平均值±三个重复的标准差(S.D.),并标准化为17S rRNA的表现量。
图2C包含以qRT-PCR测量的WT以及基因转殖稻米植物的不同组织中OsGLK1以及OsGLK2的表现量的图。组织来源及基因转殖稻米植物如图2B的图例所述。数值为平均值±三个重复的标准差(S.D.),并标准化为17S rRNA的表现量。
图2D所示为使用抗稻米及玉米GLK蛋白共有胜肽的抗体对玉米幼叶、成熟WT稻米旗叶,以及基因转殖稻米旗叶中的GLK蛋白进行的西方墨点分析。在每个泳道中加入相同量的总叶蛋白(20μg)。RbcL作为内部对照,并以玉米叶蛋白作为GLK的阳性对照。M:分子量标记。
图3A所示为4个月大的WT以及基因转殖稻米植物的照片。转基因的标记如上图1所示。
图3B为所示野生型以及基因转殖稻米植株旗叶中每叶面积的叶绿素含量的柱状图,以μg/cm2表示。图表上方所示为每叶面积的相对叶绿素含量,以相对于野生型植物的百分比表示,野生型植物设定为100%。相对于WT的统计显著性(*P<0.05;**P<0.01;***P<0.001)如图所示。数据=平均值±S.D.,n=4(从三种不同植物的叶部分获得的每个样本)。
图3C为所示野生型及基因转殖稻米植物小花中每10粒的叶绿素含量的柱状图,以μg/10粒表示。构筑体及统计显著性数值如图3B的图例所示。
图3D所示为通过酶活性分析的核酮糖双磷酸羧化酶(ribulose bisphosphatecarboxylase,rubisco)含量的柱状图(上图)以及旗叶中每叶面积的总可溶性蛋白质含量(下图)。统计检验基于成对t-检验。星号表示相较于WT的统计显著性程度(*P<0.05;**P<0.01;***P<0.001)。数据=平均值±S.D.,n=4(每个样品均来自三种不同植物的叶部分)。基因转殖植物如上所述。
图4所示为4个月大的WT以及GLK基因转殖稻米植株的旗叶中每叶面积的光饱和光合速率,以μmol/m2/s表示。该图所示为数值范围、25%至75%的四分位数(框)以及平均值(水平线)。测量条件为2000μmol/m2/s PPFD、30℃,以及400μL/LCO2。统计检验基于Wilcoxon-Mann-Whitney U-检验,用于两组的非参数比较。星号表示相较于WT具有统计学意义(**P<0.01,n=6)。构筑体如上所述。
图5A所示为WT以及5种GLK基因转殖稻米植株的总枝条生物量干重/植株。植物在5月至9月之间于温室中生长。统计检验如上针对图4所述。星号表示统计显著性程度(*P<0.05;**P<0.01;***P<0.001,n=9)。转基因构筑体如图1所示。
图5B为WT以及5种GLK基因转殖稻米植株每株植株总穗数的图。构筑及统计分析如图5A的图例所示。
图5C所示为WT以及上述5种GLK基因转殖稻米植株的每株植株总谷粒重。构筑及统计分析如图5A的图例所示。
图5D所示为WT以及5株GLK基因转殖稻米植物的每株植株1000颗谷粒的干重。构筑及统计分析如图5A的图例所示。
图6A所示为相对于WT,基因转殖植物pZmG1::ZmG1、pZmG2::ZmG2,以及pZmG1::ZmG1/pZmG2::ZmG2幼枝中差异表现基因的生物程序类别基因本体(Gene Ontology,GO)注释条目富集分析结果。涂色的格子表示上调(左图)或下调(右图)的生物程序。
图6B所示为相对于WT,基因转殖植物(如上标记)的幼枝中差异表现基因的GO注释条目富集分析。涂色的格子表示上调(左图)或下调(右图)的生物程序。
具体实施方式
如上所述,提供一种基因转殖稻米植物,其基因组中含有两个不同的异源GLK基因,每个基因都在其天然启动子及5’UTR的控制下。例如,两个GLK基因皆可来自C4植物。该GLK基因的示例性C4植物来源包含,但不限于,Zea mays L.(玉米)、Sorghum bicolor L.(高粱)、Saccharum officinarum L.(甘蔗),以及Setaria italica L.(粟)。
于一具体实施例中,该第一GLK基因为Zea mays GLK1,且该第二GLK基因为Zeamays GLK2,亦称为G2。更具体而言,由该GLK1基因表现的蛋白质具有如SEQ ID NO:3所示的胺基酸序列,由该GLK2基因表现的蛋白质具有如SEQ ID NO:6所示的胺基酸序列。
此外,该第一GLK基因的天然启动子及5’UTR可具有如SEQ ID NO:1所示的序列,而该第二GLK基因的天然启动子及5’UTR可具有如SEQ ID NO:4所示的序列。
如上所述,相较于未转形的野生型稻米植物,基因转殖稻米植物的枝条生长及谷物产量显著增加,亦即至少50%。
于一具体实施例中,相较于未转形的野生型稻米植物,例如稻米品种台农67(TNG67),pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物中枝条生长的增加,如通过生物量变化所测量的,可在65%至106%之间的范围内。
于同一实施例中,pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物的谷物产量可以比未转形的野生型稻米高出50%至95%。
发明内容部分还提到一种产生上述基因转殖植物的方法。该方法通过(i)将重组DNA构筑体引入宿主稻米植物中,该构筑体包含与其天然启动子及5’UTR可操作地连接的第一GLK基因以及与其天然启动子及5’UTR可操作地连接的第二GLK基因,以及(ii)鉴定基因转殖稻米植物,该基因转殖稻米植物相较于未转形的野生型稻米植物,例如TNG67,至少增加50%的枝条生物量及谷物产量。
于该方法中,该第一GLK基因以及该第二GLK基因彼此不同且与稻米异源。于一实施例中,该第一GLK基因以及该第二GLK基因来自C4植物,例如,玉米、高粱、甘蔗,以及粟。于一具体方法中,该第一GLK基因为Zea mays GLK1,且该第二GLK基因为Zea mays GLK2。
于一实施例中,该重组DNA构筑体可表现具有如SEQ ID NO:3所示的胺基酸序列的蛋白质,亦即Zea mays GLK1,以及具有如SEQ ID NO:6所示的胺基酸序列的蛋白质,亦即Zea mays GLK2。
GLK1以及GLK2的表现可分别在具有如SEQ ID NO:1所示的序列的天然启动子及5’UTR以及具有如SEQ ID NO:4所示的序列的天然启动子及5’UTR的控制下。
相较于未转形的野生型稻米植物,例如TNG67,已鉴定的基因转殖稻米植物的谷物产量显著更高。例如,相较于该未转形的野生型稻米植物,可以看到pZmG1::ZmG1/pZmG2::ZmG2基因转殖稻米植物的谷物产量增加了50%至95%。
基因转殖稻米植株的枝条生物量也显著高于未转形的野生型稻米植株。示例性的pZmG1::ZmG1/pZmG2::ZmG2基因转殖稻米植物的枝条质量比该未转形的野生型稻米植物的枝条质量高出65%至106%。
接下来讨论的是上述发明内容部分中提到的重组DNA构筑体。该重组DNA构筑体包含一与其天然启动子以及5’UTR可操作地连接的第一GLK基因,以及一与其天然启动子及5’UTR可操作地连接的第二GLK基因。该第二GLK基因与该第一GLK基因不同,该两个GLK基因来自一C4植物。因此,它们与稻米异源。
于一示例性重组DNA构筑体中,该第一GLK基因为Zea mays GLK1,且该第二GLK基因为Zea mays GLK2。
一特定的重组DNA构筑体包含(i)一编码如SEQ ID NO:3所示的胺基酸序列的第一GLK基因,且其受其具有如SEQ ID NO:1所示的序列的天然启动子/5’UTR的控制,以及(ii)一编码如SEQ ID NO:6所示的胺基酸序列的第二GLK基因,且其受其具有如SEQ ID NO:4所示的序列的天然启动子/5’UTR的控制。
在特定的重组DNA构筑体中,该第一GLK基因由具有如SEQ ID NO:2所示的序列的核酸编码,而该第二GLK基因由具有如SEQ ID NO:5所示的序列的核酸编码。
无需进一步阐述,相信本领域技术人员可基于本文的公开内容最大限度地利用本发明内容。因此,以下具体实施例被解释为仅为描述性的,并且无论如何不以任何方式限制本发明的其余部分。本文引用的所有出版物及专利文件均通过引用整体并入
实施例1:用于农杆菌媒介的稻米转形作用的玉米GLK构筑体
五种玉米GLK构筑体通过以下实施例7中提到的标准技术制备,包含图1中所示的基因及调控序列。
稻米(Oryza sativa cv.TNG67)愈伤组织诱导、与农杆菌共培养、转形愈伤组织的潮霉素筛选,以及植株再生依照Yeh等人2015年(Rice 8:36-48)的方法进行。将所有阳性转基因幼苗移植到土壤中并在一温室中培养以进行分子、生理,以及解剖学分析。通过热不对称交错(thermal asymmetric interlaced,TAIL)-PCR确定转基因插入位置。TAIL-PCR的条件先前在Liu等人2005年(Methods Mol.Biol.286:341-348)以及等人2009年(PlantCell 21:2163-2178)中描述过。使用根据Tail-PCR确定的转基因插入位点侧翼的3’以及5’端区域设计的基因分型引子,以基因分型PCR筛选同基因型组合的基因转殖植物。
实施例2:玉米GLK构筑体在基因转殖稻米植物中的表现
通过qRT-PCR以及西方墨点分析确定玉米GLK基因的表现量,如下文实施例7中所述。结果如图2A-2D中所示。
泛素以及35S启动子过度驱动了所有基因转殖稻米组织中玉米GLK基因的表现。在pZmUbi::ZmG1基因转殖植物的叶片中检测到大量GLK蛋白,反映ZmG1在叶片中的高表现量。参见图2B。
异基因型组合的pZmG2::ZmG2植株生长发育延迟1个月,而同基因型组合的的pZmG2::ZmG2幼苗在培养两个月后死亡。这表示pZmG2::ZmG2植物中玉米GLK2的高量表现抑制了植物的生长及发育。
在pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物的所有研究组织中,ZmG1的表现得到促进而ZmG2的表现受到抑制,进而提供这两个基因在pZmG1::ZmG1/pZmG2::ZmG2基因转殖稻米中协调表现的证据。
稻米内源性GLK1以及GLK2基因在玉米GLK基因转殖稻米植株中的表现量普遍降低。
实施例3:基因转殖稻米植物中生长及叶绿体的特征描述。
相较于所有其他测试的稻米植物,pZmG2::ZmG2(异基因型组合)稻米植物的开花及结实延迟了一个月。参见图3A。
穿透式电子显微镜(Transmission electron microscopy,TEM)分析显示,稻米中玉米GLK基因的表现诱导了所有五个基因转殖稻米品系的叶肉(M)以及束鞘(BS)细胞中大叶绿体的发育。与WT相似,所有GLK基因转殖稻米植株的M及BS叶绿体结构都具有完整的颗粒结构及类囊体膜。
相对于WT,所有5种GLK基因转殖植物的旗叶及小花内的叶绿素含量分别增加18-39%以及10-127%,旗叶中核酮糖双磷酸羧化酶(Rubisco)活性以及蛋白质含量分别增加了27-28%以及21-28%。见图3B、3C,以及3D。这些观察结果与增加的叶绿体发育以及相关基因的表现一致。
显然,这五种基因转殖稻米品系的所有光合功能都显著增强,特别是在pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物中。
实施例4:光饱和光合速率
分析旗叶以测量光合作用,因为它们比其他叶子对谷粒生长的贡献更大。相较于WT,所有5种GLK基因转殖植物的旗叶光合速率增加了5-11%。参见图4。在pZmG1::ZmG1/pZmG2::ZmG2基因转殖稻米植物中观察到最高的光合速率。
实施例5:生长性状及谷物产量
相较于WT(100%),所有5种GLK基因转殖植物的每株植物的总枝条生物量(109-180%)以及总穗数(137-224%)均增加,这主要是由于产生了额外的分蘗。参见图5A及5B。
基因转殖pZmG1::ZmG1以及pZmG2::ZmG2(异基因型组合)稻米植株的每株植物的总谷粒重分别增加了51%以及47%,同时伴随着1000粒谷粒重量的适度下降,亦即1-10%。参见图5C及5D。
令人惊讶的是,pZmG1::ZmG1/pZmG2::ZmG2稻米植株每株植物的总谷粒重增加最高达95%,平均增加70%。参见图5C。
pZmUbi::ZmG1植物由于谷粒重量减少15%而未显示总谷物产量显著增加。因此,虽然组成型泛素启动子增加了生物量(参见图5A),但其导致生殖生长减少。
不受理论的束缚,据信在强玉米泛素启动子的驱动下,玉米GLK1的高量表现可能会抑制生殖生长及发育。
实施例6:基因转殖稻米植株的转录组分析
如前所述,自WT、pZmG1::ZmG1、pZmG2::ZmG2,以及pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物的4天大幼苗的枝条及根中分离总RNA。参见刘等人2013年(PNAS 110:3979-3984)。转录组分析亦按照Liu等人(2013年)的描述进行。结果示于图6A及6B。
上调差异表现基因
pZmG2::ZmG2枝条在与生物/非生物刺激以及几丁质分解代谢过程相关的基因本体(GO)注释条目中显著富集(参见图6A),而pZmG1::ZmG1/pZmG2::ZmG2枝条主要在脂质运输的GO注释条目中富集(图6B)。
pZmG1::ZmG1根显著在转录调节相关的GO注释条目中富集,而pZmG2::ZmG2根在生物逆境反应的GO注释条目中富集,包含脂氧素(oxylipin)生合成过程(茉莉酸甲酯生合成途径)、几丁质分解代谢过程,以及植物防御素代谢过程。显然,在稻米中,两个玉米GLK基因都刺激叶绿体发育,但ZmG1的作用较像是转录调节因子,而ZmG2的作用较像是防御调节因子。
pZmG1::ZmG1/pZmG2::ZmG2根在转录调节相关的GO注释条目中富集(也见于pZmG1::ZmG1根)、几丁质分解代谢/植物防御素生合成过程(也见于pZmG2::ZmG2根)以及光合作用活化相关途径、吉贝素(GA)代谢过程、类异戊二烯生合成过程、木聚醣分解代谢过程,以及蛋白质泛素化。
总之,这些结果表示这两个玉米GLK基因以互补的方式促进稻米光合作用、代谢,以及逆境反应。
下调差异表现基因
pZmG2::ZmG2根表现出对参与离子转运、胺生合成过程、硫化合物生合成过程,以及天门冬胺酸家族胺基酸生合成过程的基因下调中富集,而pZmG1::ZmG1/pZmG2::ZmG2根主要在胺生合成以及离子转运的下调中富集。见图6A及6B。这表示当ZmG1与ZmG2在稻米中共表现时,ZmG1可能会部分抵消ZmG2的抑制功能。这与pZmG2::ZmG2植物的幼苗在萌发后在枝条及根中均表现出发育迟缓的观察结果一致,以及与具有ZmG2基因高量表现的同基因型组合的pZmG2::ZmG2植物在生长两个月后死亡的观察一致。
实施例7:材料与方法
启动子及基因选殖
使用十六烷基三甲基溴化铵方法从玉米叶(White Crystal,一种糯质玉米品种)中萃取基因组DNA。基于Ensembl植物资料库中的ZmG1(GRMZM2G026833)以及ZmG2(GRMZM2G087804)的序列(可在plants.ensembl.org/index.html网站上找到),以聚合酶连锁反应(Polymerase Chain Reaction,“PCR”)选殖玉米GLK1启动子(ZmG1;下表1中的引子SEQ ID NOs:9及10)以及G2启动子(ZmG2;引子SEQ ID NOs:11及12),两者均含有5’-UTR区域。如Liu等人,PNAS 110:3979-3984(2013年)所述,从玉米胚叶中萃取总RNA并用于合成cDNA。以PCR使用下表1中所示的特异性引子(ZmG1:SEQ ID NOs:13及14;ZmG2:SEQ ID NOs:15及16)选殖ZmG1以及ZmG2 cDNA。ZmG1以及ZmG2的启动子及全长cDNA的序列通过定序得到证实。
表1.引子序列
转形载体的构筑
在玉米或组成型启动子的控制下,制备五种构筑体以在稻米中表现ZmG1以及ZmG2基因。以PCR选殖这两个基因的玉米启动子及全长cDNA,并次选殖至转形载体pCAMBIA或Geteway(Invitrogen公司)中。使用特异性引子(表1中的SEQ ID NOs:17-24)产生具有相容限制性位点的PCR片段。ZmG1启动子及cDNA片段的5’及3’端分别以XbaI/BamHI以及BglII处理,ZmG2启动子及cDNA片段的5’及3’端分别以HindIII/BamHI以及BglII处理。融合至其cDNA(1428bp;SEQ ID NO:2)的ZmG1启动子(2134bp;SEQ ID NO:1)以及融合至其cDNA(1386bp;SEQ ID NO:5)的ZmG2启动子(1942bp;SEQ ID NO:4)分别连接到含有胭脂碱合成酶终止子的中间载体中,以获得ZmG1p::ZmG1以及ZmG2p::ZmG2片段。中间载体中的所有片段均通过限制性内切酶消化分析确认。随后,将来自中间载体的以EcoRI消化的ZmG1p::ZmG1片段以及以HindIIII/EcoRI消化的ZmG2p::ZmG2片段连接到pCAMBIA1300的EcoRI以及HindIIII/EcoRI位点,以产生pZmG1::ZmG1以及pZmG2::ZmG2构筑体。此外,将以EcoRI切割的ZmG1p::ZmG1片段选殖至ZmG2p::ZmG2载体的相同限制酶切位以生成pZmG1::ZmG1/pZmG2::ZmG2构筑体。
为了在组成型启动子控制下在稻米中表现玉米G1及G2,将含有ZmG1或ZmG2cDNA的Gateway进入选殖株分别转移到在玉米泛素启动子控制下的Gateway供体载体pCAMBIA1302或在35S启动子控制下的pH2GW7以产生pZmUbi::ZmG1以及p35S::ZmG2构筑体。
包含用于筛选的潮霉素磷酸转移酶II基因的pZmUbi::ZmG1、p35S::ZmG2、pZmG1::ZmG1、pZmG2::ZmG2,以及pZmG1::ZmG1/pZmG2::ZmG2构筑体分别通过电穿孔而转染至农杆菌(Agrobacterium tumefaciens)EHA105菌株中。所有的构筑体都在图1中以图表形式显示。
稻米转形
根据Yeh等人,2015年,Rice 8:36-48进行稻米(Oryza sativa栽培品种TNG67)愈伤组织诱导、与农杆菌共培养、转形愈伤组织的潮霉素筛选,以及植株再生。将所有基因转殖阳性幼苗移植到土壤中并于温室中进行分子、生理以及解剖学分析。
通过热不对称交错(TAIL)-PCR确定转基因插入位置
TAIL-PCR的条件如Liu等人(2013年)以及等人(2009年)所述。以T0基因转殖植物的基因组DNA作为模板用于三个连续运行的PCR,该PCR使用短的任意简并引子(AD;SEQID NOs:28-30)以及三个特异性巢式引子(SP0;SEQ IDNO:25,SP1;SEQ ID NO:26以及SP2;SEQ ID NO:27)以扩增T-DNA侧翼基因组DNA区域。Tail-PCR产物被纯化并定序。以BLAST将PCR产物序列与在Ensemble Plants中的温带粳稻(Oryza sativa Japonica)群DNA序列(可在plants.ensembl.org/Multi/Tools/Blast?db=core网站上获得)进行比较,以确定转基因在稻米染色体上的插入位点。然后将鉴定的序列用于评估基因转殖植物的接合状态。
通过基因分型PCR筛选同源基因转殖植物
基因分型引子系从以Tail-PCR确定的转基因插入位点侧翼的3’以及5’端区域设计的。基因组特异性引子对(GT-F以及GT-R;SEQ ID NOs:31-36)是基于跨假定插入位点的染色体DNA序列设计的。此外,转基因特异性SP2以及基因组特异性引子GT-R用于筛选具有转基因插入的基因转殖植物。使用Phire Plant Direct PCR Master Mix(ThermoScientific公司)按照操作手册从T1基因转殖植物的叶组织中萃取DNA。在PCR反应中,以两个引子组(基因组特异性GT-F与GT-R引子,以及转基因特异性SP2与基因组GT-R引子;表1)进行PCR扩增。由于不含转基因,WT仅产生一种来自基因组特异性引子对的PCR产物。异基因型组合的基因转殖植物通过(i)由基因组特异性引子对扩增基因组序列的未被破坏的DNA链以及(ii)由SP2与GT-R引子扩增存在转基因的一股DNA链而产生两种PCR产物。相较之下,由于两股DNA链都被特定位点的转基因插入破坏,因此在同基因型组合的基因转殖植物中只能看到来自SP2以及GT-R引子的一种PCR产物。筛选后,从阳性同基因型组合的植物的幼叶中分离基因组DNA,以通过PCR进一步确认玉米GLK基因的存在。
DNA萃取以及PCR
为了检测基因转殖稻米植物中的玉米GLK基因,在PCR分析中使用特异性引子(参见表1,SEQ ID NOs:37-43)。通过上述CTAB方法从WT以及代表性基因转殖稻米植株的幼叶中分离基因组DNA。玉米GLK基因的PCR扩增在以下条件下于热循环仪中进行:94℃/5分钟,94℃/30秒、58℃/30秒,以及72℃/30秒的30个循环,以及最后在72℃/5分钟进行延伸。
南方氏墨点分析
基因组DNA的制备以及南方氏凝胶墨点分析按照Yeh等人(2015年)先前的描述进行。从以基因组PCR证实存在玉米GLK基因的基因转殖植物中分离基因组DNA,以HindⅢ于37℃下消化16-18小时,在1%琼脂凝胶上进行电泳,并转移至一尼龙膜上进行探针杂交。按照DIG High Prime DNA Labeling and Detection Starter Kit II(Roche公司)中的指示,使用长叶毛地黄配质-dUTP探针进行随机引子DNA标记。
RNA萃取以及即时定量PCR(qRT-PCR)
使用TRIZOL试剂(Invitrogen公司)自WT以及基因转殖植物的叶片中分离总RNA,并通过酸性苯酚-氯仿萃取进行纯化。cDNA合成以及qRT-PCR按照Yeh等人的描述进行。用于扩增目标基因(SEQ ID NOs:44-57)以及17S基因(登录号X00755;SEQ ID NOs:58及59)作为标准化表现值的内部对照的基因特异性引子列于表1中。
转录组分析
使用TRIZOL试剂(Invitrogen公司)从WT、pZmG1::ZmG1、pZmG2::ZmG2,以及pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物的4天大的幼苗的枝条及根中分离总RNA。样品制备以及转录组分析使用Liu等人的方法完成。使用Tophat(2.0.10版)处理定序读数并将其定位至稻米基因组(IRGSP-1.0)。每个读数都对齐,最多允许10次命中。使用Cufflinks(版本2.1.1)评估每个基因的表现量,亦即每百万个定位的读数中每千个碱基外显子的读数(reads per kilobase exon per million reads mapped,“RPKM”)。至少一个样本中RPKM≥1的基因被认为是“表现的”并被选择用于进一步分析。为了比较WT、pZmG1::ZmG1、pZmG2::ZmG2,以及pZmG1::ZmG1/pZmG2::ZmG2基因转殖植物的枝条及根中所选基因的表现量,采用Bullard等人,2010年,BMC Bioinform.11:94中所述的上四分位数标准化程序。Tarazona等人的非参数方法(2011年,Genome Research 21:2213-2223)被用来鉴定两个样本之间的差异表现基因(differentially expressed genes,“DEGs”),该方法中的q值(差异表现概率)设置为0.740。以本研究中所有表现基因的背景集进行功能富集分析。将错误发现率(false discovery rate,“FDR”)<0.05的Fisher精确检验与MapMan的功能注释一起应用(请参阅mapman.gabipd.org网站)。针对DEGs的功能分类,GO分析采用AgriGO V2软体进行Fisher精确检验,FDR≤0.05,得到基于生物程序的GO注释。
DEGs的GO分析
为了对DEGs的生物学功能进行分类,使用基于生物学过程的AgriGO V2软体进行GO分析。有向无环图(direct acyclic graph,“DAG”)绘图是一种可视化工具,用于说明重要的GO注释条目。DAG基于GO结构的性质,指示注释条目之间的相互关系。为了研究GLK基因对每个基因转殖稻米的影响,使用≤0.05的错误发现率(FDR)分析来自根及枝条的上调与下调的DEGs。
蛋白质纯化以及西方墨点分析
如前所述,从新成熟的叶子中萃取总可溶性蛋白质用于SDS-PAGE以及西方免疫检测分析(参见Dai等人,1994年,Planta 192:287-294以及Ku等人,1999年,Nat.Biotechnol.17:76-80)。如Ku等人所述,制备抗玉米Rubisco大次单位的抗体。此外,将与玉米GLK基因(SEQ ID NO:7)共有的OsGLK1部分681bp序列选殖至pET21b载体中以产生用于产生抗GLK抗体的重组蛋白。通过亲和色层分析(Yao-Hong Biotechnology Inc.,中国台湾)纯化针对稻米/玉米GLK部分胜肽产生的抗体。
叶绿素测量以及Rubisco测定
收集旗叶或小花的叶段并以96%乙醇萃取。叶绿素含量由波长665nm以及649nm处的吸光度计算,并以叶面积或鲜重表示。如前所述测定Rubisco的活性。参见Pyankov等人,2000年,Photosynth.Res.63:69-84。酶活性基于叶绿素含量或叶面积。
光合作用测量
在夏季开花期间在温室内生长的四个月大稻米植物的主要分蘗上完整旗叶的中段用于使用CIRAS2便携式光合作用系统(PP系统,Amesbury,麻州)测量CO2以及H2O的交换。测量在晴天的上午7:00至12:30进行,条件为2000μmol m-2s-1光子通量密度、30℃叶温、70%相对湿度,以及415ppm CO2。记录稳态光合速率(Pn)、气孔导度(Gs)、细胞间二氧化碳浓度(Ci),以及蒸散速率(E)。
穿透式电子显微镜
将旗叶样品(1mm x 2mm)固定在含有1%戊二醛的0.1M磷酸盐-柠檬酸盐缓冲液中,pH 7.2,4℃过夜。在缓冲液中冲洗3次,每次20分钟之后,将样品在含有1% OsO4的相同缓冲液中固定2小时,然后在室温下冲洗3次,每次20分钟,并更换缓冲液。样品在丙酮系列中脱水,嵌入Spurr树脂中,并以Lecia Reichert Ultracut S或Lecia EM UC7超薄切片机切片。超薄切片(70-90nm)以5%醋酸双氧铀/50%甲醇以及0.4%柠檬酸铅/0.1N NaOH染色。使用80KV的FEI G2 Tecnai Spirit Twin穿透式电子显微镜进行观察,并使用GatanOrius CCD相机记录影像。
农艺性状研究
WT以及GLK基因转殖稻米植物在3月至8月之间,于自然阳光条件下在温室中的5升盆(一株/盆)中栽培。植物每天浇水,每周施肥。成熟后,种子于43℃而枝条于60℃下干燥两天后,分析总地上生物量、总穗数、1000粒谷粒重量,以及每株植物的总谷粒重。
其他具体实施例
本说明书中所公开的所有特征可以任何组合进行组合。本说明书中所公开的每个特征可被用于相同、等效或类似目的的替代特征替换。因此,除非另有明确说明,所公开的每个特征仅为等效或相似特征的通用系列的示例。
由以上描述,本领域技术人员可以很容易地确定本发明的本质特征,并且在不脱离本发明的精神及范围的情况下,可对本发明进行各种变化及修改以适应各种用途及条件。因此,其他具体实施例也在所附权利要求书的范围内。
序列表
<110> 刘扶东
<120> 增加稻米产量的方式
<130> 218048-0285PCT
<150> US 63/019,672
<151> 2020-05-04
<160> 63
<170> PatentIn version 3.5
<210> 1
<211> 2134
<212> DNA
<213> 玉米
<220>
<221> 启动子
<222> (1)..(1972)
<223> ZMG1
<220>
<221> 5'UTR
<222> (1973)..(2134)
<223> ZMG1
<400> 1
ccaaccgctg catatcttgc acgcggaccg acgacatccg cgacacgcca aaatcaacaa 60
aatcgccggc gcgcaactcc tccgccatca aactactcag cgtctcctca gacgaggcgt 120
cggccgacgg tgaggcggcg gccggcggcg tagcggcagc ggaagaagag gaggtcggca 180
tcgctatgta ccgtcggcgg cggcgggcgg tctgcttcac tcgagccaga tctccggcga 240
acaagagcac ggcgggcaga cgagcagcaa ggcggtgaaa aaggcggcta gggttttcac 300
ggagcgcgga aagtgaagtt caaaaaccgc cgacccccgc ccccttttat agacagggcg 360
cggctcttcg ggaaacccgc aatccaacag ggcgcggcgc ttcgggaaac ccgcaatcca 420
atagtacgct gtcaatcaac ggtcatgtca aaaacccccg cggaaccact tcgagcgagg 480
gccgcgtctc cgccatctag cgacgtcttc ggcaccaggt gactttgtcg aactggtccc 540
tcggagggca aatgttgggg cgaaggcaaa gacgccaccc ttcgctcgag gccttcgctg 600
cagtcgctgg tctgacagag acaaaacggg cagggacacc cttcgctcga ctcggcacca 660
gacgaaggcc tgcgacggcg tcatcccaca gtgcggcctc gtccagctca gaggcccacg 720
tgtgatccgg cccagtgtaa cgggccctgc gtggccgccg cgtgttacgg gcctaatttg 780
taaaggcatc cctgtaatta cagtctgtaa ccctgcttta tgggaatatt ctggggataa 840
cctaggtagc tcagggcaca tgcgtcctta gcacaaggcg ctgggcgctc aggtacctat 900
aaataccctc gcacagtgac cgtgggaggc cagattaaca gagctattgc catctagcgc 960
gtaaccctat taacgtcact gttcaccatt gttggccccc cttgcgagtg agagcaagtc 1020
ccaacattat acatcactct tgtagtttca taaactacaa gagagacata taagttttat 1080
aaataaattt acttttattt ttcctataaa gaaataacca aaacaaaaat atacatctcg 1140
atgagttata caactttata gttgaaaact tttacagctc aattattttt tcttcaaaat 1200
ttgatttgca tagtgtaaaa aaaacattca gcgaatgatt tctctagctg tttgccgagt 1260
gttaaaaaac actcgccaaa atgtttcttt gctgagtgtt ttttagtcga gcgttttttg 1320
tttgacaatt agtaaagagt tttttttttg ttttgttgac tgtggaaaga aaaacaataa 1380
gcaaattatt taacgctcgg caaagggcca gaattgtgaa ctatatatac tgctccgtct 1440
cgaactaggc catcggagct tctgcgagac actgagaatc cacgaatcca gcagcctcca 1500
tgcatgacct ccaagctcaa cgcagacgtg aagttcaggg cagcactcca cgatgtcccg 1560
ttggggaaga cctaatagga ccgccagtag aaaacgtcac tgtactagta cagtctccag 1620
ccgatataca tgcagctgag ccgctgtcat atcttgttct cttcctgttg ttcgccaccg 1680
tctcacgaga tcgagtccac ctctctgccc agagttgcat caacgaaaag aaaaaggaaa 1740
aaaaaccggc ggttccaaaa aagactcggc cacacacagg cacacgctcg ctacgaggcc 1800
acaataattc gcagaagcga ccaagctgca cttgcgctcg cggacaaggc cacccatcta 1860
aaggattcag agagatcttt gcgaggcttg tggtcgagat gccttcgccg ttcgccacaa 1920
atactccccg gagagcgagt acttactggc tcgccatatg gcaccaacag caaaacagcg 1980
aaacagtgca gagacaagct acaacaacct accctaacag gccattcctt ccactgcact 2040
tcattcgatc ctgagctata gctggctgcc tgagctcgct ccaagaagtg tatctatagt 2100
atagacacta ggcttcgtgt ggcgtgctcg agat 2134
<210> 2
<211> 1428
<212> DNA
<213> 玉米
<220>
<221> CDS
<222> (1)..(1428)
<223> ZMG1
<400> 2
atg ctt gca gtg tcg ccg tcg ccg gtg cgg tgt gcc gat gcg gag gag 48
Met Leu Ala Val Ser Pro Ser Pro Val Arg Cys Ala Asp Ala Glu Glu
1 5 10 15
tgc ggc gga gga ggc gcc agc aag gaa atg gag gag acc gcc gtc ggg 96
Cys Gly Gly Gly Gly Ala Ser Lys Glu Met Glu Glu Thr Ala Val Gly
20 25 30
cct gtg tcc gac tcg gac ctg gat ttc gac ttc acg gtc gac gac ata 144
Pro Val Ser Asp Ser Asp Leu Asp Phe Asp Phe Thr Val Asp Asp Ile
35 40 45
gac ttc ggg gac ttc ttc ctc agg cta gac gac ggg gat gac gcg ctg 192
Asp Phe Gly Asp Phe Phe Leu Arg Leu Asp Asp Gly Asp Asp Ala Leu
50 55 60
ccg ggc ctc gag gtc gac cct gcc gag atc gtc ttc gct gac ttc gag 240
Pro Gly Leu Glu Val Asp Pro Ala Glu Ile Val Phe Ala Asp Phe Glu
65 70 75 80
gca atc gcc acc gcc ggc ggc gat ggc ggc gtc acg gac cag gag gtg 288
Ala Ile Ala Thr Ala Gly Gly Asp Gly Gly Val Thr Asp Gln Glu Val
85 90 95
ccc agt gtc ctg ccc ttt gcg gac gcg gcg cac ata ggc gcc gtg gat 336
Pro Ser Val Leu Pro Phe Ala Asp Ala Ala His Ile Gly Ala Val Asp
100 105 110
ccg tgt tgt ggt gtc ctt ggc gag gac aac gac gca gcg tgc gca gac 384
Pro Cys Cys Gly Val Leu Gly Glu Asp Asn Asp Ala Ala Cys Ala Asp
115 120 125
gtg gaa gaa ggg aaa ggg gag tgc gac cat gcc gac gag gta gca gcc 432
Val Glu Glu Gly Lys Gly Glu Cys Asp His Ala Asp Glu Val Ala Ala
130 135 140
gcc ggt aat aat aat agc gac tcc ggt gag gcc ggc tgt gga gga gcc 480
Ala Gly Asn Asn Asn Ser Asp Ser Gly Glu Ala Gly Cys Gly Gly Ala
145 150 155 160
ttt gcc ggc gaa aaa tca ccg tcg tcg acg gca tcg tcg tcg cag gag 528
Phe Ala Gly Glu Lys Ser Pro Ser Ser Thr Ala Ser Ser Ser Gln Glu
165 170 175
gct gag agc cgg cgc aag gtg tcc aag aag cac tcc caa ggg aag aag 576
Ala Glu Ser Arg Arg Lys Val Ser Lys Lys His Ser Gln Gly Lys Lys
180 185 190
aaa gca aag gtg gat tgg acg ccg gag ctt cac cgg aga ttc gtt cag 624
Lys Ala Lys Val Asp Trp Thr Pro Glu Leu His Arg Arg Phe Val Gln
195 200 205
gcg gtg gag gag ctg ggc atc gac aag gcg gtg ccg tcc agg atc ctc 672
Ala Val Glu Glu Leu Gly Ile Asp Lys Ala Val Pro Ser Arg Ile Leu
210 215 220
gag atc atg ggg atc gac tcc ctc acg cgg cat aac ata gcc agc cat 720
Glu Ile Met Gly Ile Asp Ser Leu Thr Arg His Asn Ile Ala Ser His
225 230 235 240
ctg cag aag tac cgt tcc cac agg aag cac atg ctt gcg agg gag gtg 768
Leu Gln Lys Tyr Arg Ser His Arg Lys His Met Leu Ala Arg Glu Val
245 250 255
gag gca gcg acg tgg acg acg cac cgg cgg ccg atg tac gct gcc ccc 816
Glu Ala Ala Thr Trp Thr Thr His Arg Arg Pro Met Tyr Ala Ala Pro
260 265 270
agc ggc gcc gtg aag agg ccc gac tct aac gcg tgg acc gtg ccg acc 864
Ser Gly Ala Val Lys Arg Pro Asp Ser Asn Ala Trp Thr Val Pro Thr
275 280 285
atc ggt ttc cct ccg ccg gcg ggg acc cct cct cgt ccg gtg cag cac 912
Ile Gly Phe Pro Pro Pro Ala Gly Thr Pro Pro Arg Pro Val Gln His
290 295 300
ttc ggg agg cca ctg cac gtc tgg ggc cat ccg agt ccg acg cca gcg 960
Phe Gly Arg Pro Leu His Val Trp Gly His Pro Ser Pro Thr Pro Ala
305 310 315 320
gtg gag tca ccc cgg gtg cca atg tgg cct cgg cat ctc gcc ccc cgc 1008
Val Glu Ser Pro Arg Val Pro Met Trp Pro Arg His Leu Ala Pro Arg
325 330 335
gcc ccg ccg ccg ccg ccg tgg gct ccg cca ccg cca gct gac ccg gcg 1056
Ala Pro Pro Pro Pro Pro Trp Ala Pro Pro Pro Pro Ala Asp Pro Ala
340 345 350
tcg ttc tgg cac cat gct tac atg agg ggg cct gct gcc cat atg cca 1104
Ser Phe Trp His His Ala Tyr Met Arg Gly Pro Ala Ala His Met Pro
355 360 365
gac cag gtg gcg gtg act cca tgc gtg gca gtg cca atg gca gca gcg 1152
Asp Gln Val Ala Val Thr Pro Cys Val Ala Val Pro Met Ala Ala Ala
370 375 380
cgt ttc cct gct cca cac gtg agg ggt tct ttg cca tgg cca cct ccg 1200
Arg Phe Pro Ala Pro His Val Arg Gly Ser Leu Pro Trp Pro Pro Pro
385 390 395 400
atg tac aga cct ctc gtt cct cca gca ctc gca ggc aag agc cag caa 1248
Met Tyr Arg Pro Leu Val Pro Pro Ala Leu Ala Gly Lys Ser Gln Gln
405 410 415
gac gcg ctg ttt cag cta cag ata cag cca tca agc gag agc ata gat 1296
Asp Ala Leu Phe Gln Leu Gln Ile Gln Pro Ser Ser Glu Ser Ile Asp
420 425 430
gca gca ata ggt gat gtc tta acg aag ccg tgg ctg ccg ctg ccc ctc 1344
Ala Ala Ile Gly Asp Val Leu Thr Lys Pro Trp Leu Pro Leu Pro Leu
435 440 445
gga ctg aag ccc cct tcg gta gac agt gtc atg ggc gag ctg cag agg 1392
Gly Leu Lys Pro Pro Ser Val Asp Ser Val Met Gly Glu Leu Gln Arg
450 455 460
caa ggc gta gcg aat gtg ccg caa gct tgt gga tga 1428
Gln Gly Val Ala Asn Val Pro Gln Ala Cys Gly
465 470 475
<210> 3
<211> 475
<212> PRT
<213> 玉米
<400> 3
Met Leu Ala Val Ser Pro Ser Pro Val Arg Cys Ala Asp Ala Glu Glu
1 5 10 15
Cys Gly Gly Gly Gly Ala Ser Lys Glu Met Glu Glu Thr Ala Val Gly
20 25 30
Pro Val Ser Asp Ser Asp Leu Asp Phe Asp Phe Thr Val Asp Asp Ile
35 40 45
Asp Phe Gly Asp Phe Phe Leu Arg Leu Asp Asp Gly Asp Asp Ala Leu
50 55 60
Pro Gly Leu Glu Val Asp Pro Ala Glu Ile Val Phe Ala Asp Phe Glu
65 70 75 80
Ala Ile Ala Thr Ala Gly Gly Asp Gly Gly Val Thr Asp Gln Glu Val
85 90 95
Pro Ser Val Leu Pro Phe Ala Asp Ala Ala His Ile Gly Ala Val Asp
100 105 110
Pro Cys Cys Gly Val Leu Gly Glu Asp Asn Asp Ala Ala Cys Ala Asp
115 120 125
Val Glu Glu Gly Lys Gly Glu Cys Asp His Ala Asp Glu Val Ala Ala
130 135 140
Ala Gly Asn Asn Asn Ser Asp Ser Gly Glu Ala Gly Cys Gly Gly Ala
145 150 155 160
Phe Ala Gly Glu Lys Ser Pro Ser Ser Thr Ala Ser Ser Ser Gln Glu
165 170 175
Ala Glu Ser Arg Arg Lys Val Ser Lys Lys His Ser Gln Gly Lys Lys
180 185 190
Lys Ala Lys Val Asp Trp Thr Pro Glu Leu His Arg Arg Phe Val Gln
195 200 205
Ala Val Glu Glu Leu Gly Ile Asp Lys Ala Val Pro Ser Arg Ile Leu
210 215 220
Glu Ile Met Gly Ile Asp Ser Leu Thr Arg His Asn Ile Ala Ser His
225 230 235 240
Leu Gln Lys Tyr Arg Ser His Arg Lys His Met Leu Ala Arg Glu Val
245 250 255
Glu Ala Ala Thr Trp Thr Thr His Arg Arg Pro Met Tyr Ala Ala Pro
260 265 270
Ser Gly Ala Val Lys Arg Pro Asp Ser Asn Ala Trp Thr Val Pro Thr
275 280 285
Ile Gly Phe Pro Pro Pro Ala Gly Thr Pro Pro Arg Pro Val Gln His
290 295 300
Phe Gly Arg Pro Leu His Val Trp Gly His Pro Ser Pro Thr Pro Ala
305 310 315 320
Val Glu Ser Pro Arg Val Pro Met Trp Pro Arg His Leu Ala Pro Arg
325 330 335
Ala Pro Pro Pro Pro Pro Trp Ala Pro Pro Pro Pro Ala Asp Pro Ala
340 345 350
Ser Phe Trp His His Ala Tyr Met Arg Gly Pro Ala Ala His Met Pro
355 360 365
Asp Gln Val Ala Val Thr Pro Cys Val Ala Val Pro Met Ala Ala Ala
370 375 380
Arg Phe Pro Ala Pro His Val Arg Gly Ser Leu Pro Trp Pro Pro Pro
385 390 395 400
Met Tyr Arg Pro Leu Val Pro Pro Ala Leu Ala Gly Lys Ser Gln Gln
405 410 415
Asp Ala Leu Phe Gln Leu Gln Ile Gln Pro Ser Ser Glu Ser Ile Asp
420 425 430
Ala Ala Ile Gly Asp Val Leu Thr Lys Pro Trp Leu Pro Leu Pro Leu
435 440 445
Gly Leu Lys Pro Pro Ser Val Asp Ser Val Met Gly Glu Leu Gln Arg
450 455 460
Gln Gly Val Ala Asn Val Pro Gln Ala Cys Gly
465 470 475
<210> 4
<211> 1942
<212> DNA
<213> 玉米
<220>
<221> 启动子
<222> (1)..(1288)
<223> ZMG2
<220>
<221> 5'UTR
<222> (1289)..(1942)
<223> ZMG2
<400> 4
aagcttcagg ttattggata aggtgaggaa aattccatat ttctgaaaaa ccgatgatca 60
aagttagtcc aagtgtctag gtcgcggatc gttcgcgata gcatcggacc gtgcaagcaa 120
agacaccgga ctgttcgacg gcatcagaaa tgtcgcctta gttgattcgg ctaacaatgc 180
ggccgatgca gttgattgga gaatgtcttt gattgtatac ctgcataatc caggtgttag 240
gacagacatg atcatttgac agatagattt caagtatgtt ttaatcgatg atgaacttta 300
tcgctgaact tctagtgatg gcttgcttaa ttgcttgggc cctaacgatg ctttattagc 360
tatggacaaa gtaaatgaag gaatttgtgg tactcatcaa tcggctctaa agatgaagtg 420
gctcttaagg tggtctatgg tttattagcc tgatatgata tcgattgttt taaatactag 480
aaagggtgtg aaatatgtca ggagttcggc gatctgcaat tagttctgag ccccgcgtgc 540
ggaccgtccg gtgtgacacg gagaagaccc gctcatgcag cgagatcgcg gaccgtccgc 600
gctcccgcag agagcaccgc caggagatac atccctaatg tttggctcca aatcggagcc 660
aacactaccc tacatgttct aaattacaat gtgttttagt atttttttta ttgtatagtt 720
tttattatat aacttattat aagtagatga atagtaaaat ctgttggaat atgactctaa 780
tttattttat atatttggaa tatgtcttga aatatgcttt catgtacgcc tataaatata 840
gatctttcat gtaacttgta ttctgcggtt gatagttgat ctgtttcctc gtggtcgttt 900
gctcctccat attggaggga attttcgcat gtaaatcttt gtgtctattt tatttcgtaa 960
caaaatctat agattctaaa aagctaaaac ttattatagt ttagaacgga ggggcagtgg 1020
atataggccg agtggtatgg caccgtaact gtaaataaca tgcatgcatg gagacgacga 1080
cgcatgtaga ttatatatcg tagtggggtt ccggaacaat atacaaaaaa catatatgcg 1140
tgtatttaat tattatttgt ggcggctggt gcatgtgaac tactggaata ataatggtga 1200
aattgaggag ggtgcactga ctgagatacg gtagacatgc atagataagc actgcggagc 1260
catgctgcct gcctgcaata atgcttgtgt gtatacagcg acacgacacg agacgagagg 1320
gagggtgaac tgaaagagag acccggctag agctataaag caagtggggg aagagagaga 1380
gagaggagaa ggagaaggag aggtttttat gtatgtgtgg tggacatgcg tcctcctgtc 1440
gtctcgagag agcggaggcc ctaatactcc aatcaccacg caccaaacta gctagctagc 1500
agtcgtcttg cattgtagct agtccattgc tctcatcttc tcttcttctt cccccagtcc 1560
tccccccctt tcccctcttc ggcctctctc gctcagctca ctcttcatta agcgagctca 1620
cgtcgttcct cccttcttct tcttcttatc cgttgttcaa ttcgttcagc tagccggcca 1680
gagatcgagc atcatctcca tcatcgattc atccatctca tctttctctt cttctattct 1740
atgtcgtcgt cgtcccagat tagatcgaag ctgctagcag tctatccagt ctctagctag 1800
ctagctagat caagcccgca gactatacaa tataatacaa gctagctacg tgcttattat 1860
tgctttatta gtctagaata atcttgatat atacgatcga acatatatct cgatctccac 1920
cgatacacac ccggccctat cc 1942
<210> 5
<211> 1386
<212> DNA
<213> 玉米
<220>
<221> CDS
<222> (1)..(1386)
<223> ZMG2
<400> 5
atg ctt gag gtg tcg acg ctg cgc ggc cct act agc agc ggc agc aag 48
Met Leu Glu Val Ser Thr Leu Arg Gly Pro Thr Ser Ser Gly Ser Lys
1 5 10 15
gcg gag cag cac tgc ggc ggc ggc ggc ggc ttc gtc ggc gac cac cat 96
Ala Glu Gln His Cys Gly Gly Gly Gly Gly Phe Val Gly Asp His His
20 25 30
gtg gtg ttc ccg acg tcc ggc gac tgc ttc gcc atg gtg gac gac aac 144
Val Val Phe Pro Thr Ser Gly Asp Cys Phe Ala Met Val Asp Asp Asn
35 40 45
ctc ctg gac tac atc gac ttc agc tgc gac gtg ccc ttc ttc gac gct 192
Leu Leu Asp Tyr Ile Asp Phe Ser Cys Asp Val Pro Phe Phe Asp Ala
50 55 60
gac ggg gac atc ctc ccc gac ctg gag gta gac acc acg gag ctc ctc 240
Asp Gly Asp Ile Leu Pro Asp Leu Glu Val Asp Thr Thr Glu Leu Leu
65 70 75 80
gcc gag ttc tcg tcc acc cct cct gcg gac gac ctg ctg gca gtg gca 288
Ala Glu Phe Ser Ser Thr Pro Pro Ala Asp Asp Leu Leu Ala Val Ala
85 90 95
gta ttc ggc gcc gac gac cag ccg gcg gcg gca gta gca caa gag aag 336
Val Phe Gly Ala Asp Asp Gln Pro Ala Ala Ala Val Ala Gln Glu Lys
100 105 110
ccg tcg tcg tcg ttg gag caa aca tgt ggt gac gac aaa ggt gta gca 384
Pro Ser Ser Ser Leu Glu Gln Thr Cys Gly Asp Asp Lys Gly Val Ala
115 120 125
gta gcc gcc gcc aga aga aag ctg cag acg acg acg acg acg acg acg 432
Val Ala Ala Ala Arg Arg Lys Leu Gln Thr Thr Thr Thr Thr Thr Thr
130 135 140
acg gag gag gag gat tct tct cct gcc ggg tcc ggg gcc aac aag tcg 480
Thr Glu Glu Glu Asp Ser Ser Pro Ala Gly Ser Gly Ala Asn Lys Ser
145 150 155 160
tcg gcg tcg gca gag ggc cac agc agc aag aag aag tcg gcg ggc aag 528
Ser Ala Ser Ala Glu Gly His Ser Ser Lys Lys Lys Ser Ala Gly Lys
165 170 175
aac tcc aac ggc ggc aag cgc aag gtg aag gtg gac tgg acg ccg gag 576
Asn Ser Asn Gly Gly Lys Arg Lys Val Lys Val Asp Trp Thr Pro Glu
180 185 190
ctg cac cgg cgg ttc gtg cag gcg gtg gag cag ctg ggc atc gac aag 624
Leu His Arg Arg Phe Val Gln Ala Val Glu Gln Leu Gly Ile Asp Lys
195 200 205
gcc gtg ccg tcc agg atc ctg gag atc atg ggc acg gac tgc ctc aca 672
Ala Val Pro Ser Arg Ile Leu Glu Ile Met Gly Thr Asp Cys Leu Thr
210 215 220
agg cac aac att gcc agc cac ctc cag aag tac cgg tcg cac aga aag 720
Arg His Asn Ile Ala Ser His Leu Gln Lys Tyr Arg Ser His Arg Lys
225 230 235 240
cac ctg atg gcg cgg gag gcg gag gcc gcc acc tgg gcg cag aag cgc 768
His Leu Met Ala Arg Glu Ala Glu Ala Ala Thr Trp Ala Gln Lys Arg
245 250 255
cac atg tac gcg ccg cca gct cca agg acg acg acg acg acg gac gcc 816
His Met Tyr Ala Pro Pro Ala Pro Arg Thr Thr Thr Thr Thr Asp Ala
260 265 270
gcc agg ccg ccg tgg gtg gtg ccg acg acc atc ggg ttc ccg ccg ccg 864
Ala Arg Pro Pro Trp Val Val Pro Thr Thr Ile Gly Phe Pro Pro Pro
275 280 285
cgc ttc tgc cgc ccg ctg cac gtg tgg ggc cac ccg ccg ccg cac gcc 912
Arg Phe Cys Arg Pro Leu His Val Trp Gly His Pro Pro Pro His Ala
290 295 300
gcc gcg gct gaa gca gca gcg gcg act ccc atg ctg ccc gtg tgg ccg 960
Ala Ala Ala Glu Ala Ala Ala Ala Thr Pro Met Leu Pro Val Trp Pro
305 310 315 320
cgt cac ctg gcg ccg ccc cgg cac ctg gcg ccg tgg gcg cac ccg acg 1008
Arg His Leu Ala Pro Pro Arg His Leu Ala Pro Trp Ala His Pro Thr
325 330 335
ccg gtg gac ccg gcg ttc tgg cac cag cag tac agc gct gcc agg aaa 1056
Pro Val Asp Pro Ala Phe Trp His Gln Gln Tyr Ser Ala Ala Arg Lys
340 345 350
tgg ggc cca cag gca gcc gcc gtg acg caa ggg acg cca tgc gtg ccg 1104
Trp Gly Pro Gln Ala Ala Ala Val Thr Gln Gly Thr Pro Cys Val Pro
355 360 365
ctg ccg agg ttt ccg gtg cct cac ccc atc tac agc aga ccg gcg atg 1152
Leu Pro Arg Phe Pro Val Pro His Pro Ile Tyr Ser Arg Pro Ala Met
370 375 380
gta cct ccg ccg cca agc acc acc aag cta gct caa ctg cat ctg gag 1200
Val Pro Pro Pro Pro Ser Thr Thr Lys Leu Ala Gln Leu His Leu Glu
385 390 395 400
ctc caa gcg cac ccg tcc aag gag agc atc gac gca gcc atc gga gat 1248
Leu Gln Ala His Pro Ser Lys Glu Ser Ile Asp Ala Ala Ile Gly Asp
405 410 415
gtt tta gtg aag cca tgg ctg ccg ctt cca ctg ggg ctc aag ccg ccg 1296
Val Leu Val Lys Pro Trp Leu Pro Leu Pro Leu Gly Leu Lys Pro Pro
420 425 430
tcg ctc gac agc gtc atg tcg gag ctg cac aag caa ggc gta cca aaa 1344
Ser Leu Asp Ser Val Met Ser Glu Leu His Lys Gln Gly Val Pro Lys
435 440 445
atc cca ccg gcg gct gcc acc acc acc ggc gcc acc gga tga 1386
Ile Pro Pro Ala Ala Ala Thr Thr Thr Gly Ala Thr Gly
450 455 460
<210> 6
<211> 461
<212> PRT
<213> 玉米
<400> 6
Met Leu Glu Val Ser Thr Leu Arg Gly Pro Thr Ser Ser Gly Ser Lys
1 5 10 15
Ala Glu Gln His Cys Gly Gly Gly Gly Gly Phe Val Gly Asp His His
20 25 30
Val Val Phe Pro Thr Ser Gly Asp Cys Phe Ala Met Val Asp Asp Asn
35 40 45
Leu Leu Asp Tyr Ile Asp Phe Ser Cys Asp Val Pro Phe Phe Asp Ala
50 55 60
Asp Gly Asp Ile Leu Pro Asp Leu Glu Val Asp Thr Thr Glu Leu Leu
65 70 75 80
Ala Glu Phe Ser Ser Thr Pro Pro Ala Asp Asp Leu Leu Ala Val Ala
85 90 95
Val Phe Gly Ala Asp Asp Gln Pro Ala Ala Ala Val Ala Gln Glu Lys
100 105 110
Pro Ser Ser Ser Leu Glu Gln Thr Cys Gly Asp Asp Lys Gly Val Ala
115 120 125
Val Ala Ala Ala Arg Arg Lys Leu Gln Thr Thr Thr Thr Thr Thr Thr
130 135 140
Thr Glu Glu Glu Asp Ser Ser Pro Ala Gly Ser Gly Ala Asn Lys Ser
145 150 155 160
Ser Ala Ser Ala Glu Gly His Ser Ser Lys Lys Lys Ser Ala Gly Lys
165 170 175
Asn Ser Asn Gly Gly Lys Arg Lys Val Lys Val Asp Trp Thr Pro Glu
180 185 190
Leu His Arg Arg Phe Val Gln Ala Val Glu Gln Leu Gly Ile Asp Lys
195 200 205
Ala Val Pro Ser Arg Ile Leu Glu Ile Met Gly Thr Asp Cys Leu Thr
210 215 220
Arg His Asn Ile Ala Ser His Leu Gln Lys Tyr Arg Ser His Arg Lys
225 230 235 240
His Leu Met Ala Arg Glu Ala Glu Ala Ala Thr Trp Ala Gln Lys Arg
245 250 255
His Met Tyr Ala Pro Pro Ala Pro Arg Thr Thr Thr Thr Thr Asp Ala
260 265 270
Ala Arg Pro Pro Trp Val Val Pro Thr Thr Ile Gly Phe Pro Pro Pro
275 280 285
Arg Phe Cys Arg Pro Leu His Val Trp Gly His Pro Pro Pro His Ala
290 295 300
Ala Ala Ala Glu Ala Ala Ala Ala Thr Pro Met Leu Pro Val Trp Pro
305 310 315 320
Arg His Leu Ala Pro Pro Arg His Leu Ala Pro Trp Ala His Pro Thr
325 330 335
Pro Val Asp Pro Ala Phe Trp His Gln Gln Tyr Ser Ala Ala Arg Lys
340 345 350
Trp Gly Pro Gln Ala Ala Ala Val Thr Gln Gly Thr Pro Cys Val Pro
355 360 365
Leu Pro Arg Phe Pro Val Pro His Pro Ile Tyr Ser Arg Pro Ala Met
370 375 380
Val Pro Pro Pro Pro Ser Thr Thr Lys Leu Ala Gln Leu His Leu Glu
385 390 395 400
Leu Gln Ala His Pro Ser Lys Glu Ser Ile Asp Ala Ala Ile Gly Asp
405 410 415
Val Leu Val Lys Pro Trp Leu Pro Leu Pro Leu Gly Leu Lys Pro Pro
420 425 430
Ser Leu Asp Ser Val Met Ser Glu Leu His Lys Gln Gly Val Pro Lys
435 440 445
Ile Pro Pro Ala Ala Ala Thr Thr Thr Gly Ala Thr Gly
450 455 460
<210> 7
<211> 681
<212> DNA
<213> 稻米
<220>
<221> misc_feature
<223> OsGLK1基因部分序列
<400> 7
gcgaaggtgg actggacgcc tgagcttcac cggaggttcg tgcaggcggt ggagcagctc 60
ggcatcgaca aggccgtgcc gtcgaggata cttgagatca tggggatcga ctctctcacc 120
cggcacaaca tagccagcca tcttcagaag taccggtcac acagaaaaca catgattgcg 180
agagaggcgg aggcagcgag ttggacccaa cggcggcaga tttacgccgc cggtggaggt 240
gctgttgcga agaggccgga gtccaacgcg tggaccgtgc caaccattgg cttccctcct 300
cctccgccac caccaccatc accggctccg attcaacatt ttgctcgccc gttgcatgtt 360
tggggccacc cgacgatgga cccgtcccga gttccagtgt ggccaccgcg gcacctcgtt 420
ccccgtggcc cggcgccacc atgggttcca ccgccgccgc cgtcggaccc tgctttctgg 480
caccaccctt acatgagggg gccagcacat gtgccaactc aagggacacc ttgcatggcg 540
atgcccatgc cagctgcgag atttcctgct ccaccggtgc caggagttgt cccgtgtcca 600
atgtataggc cattgactcc accagcactg gcgagcaaga atcagcagga cgcacagctt 660
caactccagg ttcaaccatc a 681
<210> 8
<211> 227
<212> PRT
<213> 稻米
<220>
<221> misc_feature
<223> OsGLK1基因部分蛋白质序列
<400> 8
Ala Lys Val Asp Trp Thr Pro Glu Leu His Arg Arg Phe Val Gln Ala
1 5 10 15
Val Glu Gln Leu Gly Ile Asp Lys Ala Val Pro Ser Arg Ile Leu Glu
20 25 30
Ile Met Gly Ile Asp Ser Leu Thr Arg His Asn Ile Ala Ser His Leu
35 40 45
Gln Lys Tyr Arg Ser His Arg Lys His Met Ile Ala Arg Glu Ala Glu
50 55 60
Ala Ala Ser Trp Thr Gln Arg Arg Gln Ile Tyr Ala Ala Gly Gly Gly
65 70 75 80
Ala Val Ala Lys Arg Pro Glu Ser Asn Ala Trp Thr Val Pro Thr Ile
85 90 95
Gly Phe Pro Pro Pro Pro Pro Pro Pro Pro Ser Pro Ala Pro Ile Gln
100 105 110
His Phe Ala Arg Pro Leu His Val Trp Gly His Pro Thr Met Asp Pro
115 120 125
Ser Arg Val Pro Val Trp Pro Pro Arg His Leu Val Pro Arg Gly Pro
130 135 140
Ala Pro Pro Trp Val Pro Pro Pro Pro Pro Ser Asp Pro Ala Phe Trp
145 150 155 160
His His Pro Tyr Met Arg Gly Pro Ala His Val Pro Thr Gln Gly Thr
165 170 175
Pro Cys Met Ala Met Pro Met Pro Ala Ala Arg Phe Pro Ala Pro Pro
180 185 190
Val Pro Gly Val Val Pro Cys Pro Met Tyr Arg Pro Leu Thr Pro Pro
195 200 205
Ala Leu Ala Ser Lys Asn Gln Gln Asp Ala Gln Leu Gln Leu Gln Val
210 215 220
Gln Pro Ser
225
<210> 9
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> G1 启动子_F
<400> 9
ccaaccctgc atatcttgc 19
<210> 10
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G1 启动子_R
<400> 10
ggcgacactg caagcatatc 20
<210> 11
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G2 启动子_F
<400> 11
gctatctcca accacggctc 20
<210> 12
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G2 启动子_R
<400> 12
cttgctgccg ctgctagtag 20
<210> 13
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G1 cDNA_F
<400> 13
atgcttgcag tgtcgccgtc 20
<210> 14
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> G1 cDNA_R
<400> 14
tcatccacaa gcttgcggca c 21
<210> 15
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> G2 cDNA_F
<400> 15
atgcttgagg tgtcgacgct gcgc 24
<210> 16
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G2 cDNA_R
<400> 16
tcatccggtg gcgccggtgg 20
<210> 17
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> G1p_F (XbaI)
<400> 17
cgtctagacc aaccgctgca tatcttg 27
<210> 18
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> G1p_R (BhI)
<400> 18
caggatccgg cgacactgca agcata 26
<210> 19
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G2p_mF (HdIII)
<400> 19
tggcttggca catgaagctt 20
<210> 20
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> G2p_R (BhI)
<400> 20
caggatccac ttgctgccgc tgctag 26
<210> 21
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> G1_cDNA_F (BglII)
<400> 21
ggagatctat gcttgcagtg tcg 23
<210> 22
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> G1_cDNA_R (BglII)
<400> 22
ggagatcttc atccacaagc ttgcg 25
<210> 23
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> G2_cDNA_F (BglII)
<400> 23
ggagatctat gcttgaggtg tcg 23
<210> 24
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> G2_cDNA_F (BglII)
<400> 24
atagatcttc atccggtggc gcc 23
<210> 25
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> SP0
<400> 25
tcgagtttct ccataataat gtgtga 26
<210> 26
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> SP1
<400> 26
taatgtgtga gtagttccca gata 24
<210> 27
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> SP2
<400> 27
aatccagtac taaaatccag atcc 24
<210> 28
<211> 16
<212> DNA
<213> 人工序列
<220>
<223> AD1
<220>
<221> 其他特征
<222> (3)..(3)
<223> n is a, c, g, or t
<220>
<221> 其他特征
<222> (11)..(11)
<223> n is a, c, g, or t
<400> 28
gtncgaswca nawgtt 16
<210> 29
<211> 15
<212> DNA
<213> 人工序列
<220>
<223> AD2
<220>
<221> 其他特征
<222> (4)..(4)
<223> n is a, c, g, or t
<220>
<221> 其他特征
<222> (9)..(9)
<223> n is a, c, g, or t
<400> 29
gwgnagwanc asaga 15
<210> 30
<211> 15
<212> DNA
<213> 人工序列
<220>
<223> AD3
<220>
<221> 其他特征
<222> (1)..(1)
<223> n is a, c, g, or t
<400> 30
ntcgastwts gwgtt 15
<210> 31
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> GT_F_G1
<400> 31
cgatcgtgtt ggcacgtgat g 21
<210> 32
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> GT_R_G1
<400> 32
tcgaagagtc aaccttcgag gtc 23
<210> 33
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> GT_F_G2
<400> 33
gccttggagg aagtagaaca gc 22
<210> 34
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> GT_R_G2
<400> 34
gtaggtttgg gaaatctttc agc 23
<210> 35
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> GT_F_G1G2
<400> 35
gtgaccggaa tccctctcaa g 21
<210> 36
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> GT_R_G1G2
<400> 36
gttgcactgt gtgtactagt c 21
<210> 37
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> G1p_F
<400> 37
ccaaccgctg catatcttg 19
<210> 38
<211> 18
<212> DNA
<213> 人工序列
<220>
<223> G1p_R
<400> 38
ggcgacactg caagcata 18
<210> 39
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> G2p_F
<400> 39
tggcttggca catgaagctt 20
<210> 40
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> G2p_R
<400> 40
cacttgctgc cgctgctag 19
<210> 41
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> G1_mF
<400> 41
gtcgttctgg caccatgctt ac 22
<210> 42
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> G2_mF
<400> 42
cgcacagaaa gcacctgatg g 21
<210> 43
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> NOS_R
<400> 43
tcatcgcaag accggcaaca 20
<210> 44
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> G1-F1 qPCR
<400> 44
gcacataggc gccgtggatc c 21
<210> 45
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> G1-R1 qPCR
<400> 45
ttaccggcgg ctgctacctc g 21
<210> 46
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> G2-F qPCR
<400> 46
gacaaaggtg tagcagtagc cgccgc 26
<210> 47
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> G2-R qPCR
<400> 47
cgttggagtt cttgcccgcc gacttc 26
<210> 48
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> OsGLK1-F qPCR
<400> 48
gatgctggca tggaggtcgt c 21
<210> 49
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> OsGLK1-R qPCR
<400> 49
tgcacacctt cgcctccact c 21
<210> 50
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> OsGLK2-F qPCR
<400> 50
gtgacgacgg aggattcctc g 21
<210> 51
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> OsGLK2-R qPCR
<400> 51
ccgatgacga cgacgacgac g 21
<210> 52
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_F_exon 1
<400> 52
cgcgctgtcc aaggcctatc tct 23
<210> 53
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_R_exon 1
<400> 53
gcaacagcat ggggagcatt gg 22
<210> 54
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_F_exon 6
<400> 54
ggcttggcat aactggagct gc 22
<210> 55
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_R_exon 6-7
<400> 55
aactcaaagc cccctgggag ag 22
<210> 56
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_F_exon 7
<400> 56
ggtggcatgc tgttaggaag aaccc 25
<210> 57
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> OsFRD3_R_exon 7
<400> 57
gccatagctg ttgggccttg tcg 23
<210> 58
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 17S rRNA_F
<400> 58
gataactcga cggatcgcac gg 22
<210> 59
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> 17S rRNA_R
<400> 59
gcctgctgcc ttccttggat gtg 23
<210> 60
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> OsActin_F
<400> 60
atccttgtat gctagcggtc ga 22
<210> 61
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> OsActin_R
<400> 61
atccaaccgg aggatagcat g 21
<210> 62
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> HptII_F
<400> 62
gacctgatgc agctctcgga g 21
<210> 63
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> HptII_R
<400> 63
tgctccatac aagccaacca cg 22
Claims (20)
1.一种基因转殖稻米植物,在其基因组中包含重组DNA构筑体,该构筑体包含第一核酸,该第一核酸具有与其天然启动子及5’非转译区(5’untranslated region,5’UTR)可操作地连接的第一Golden 2-like转录因子(GLK)基因的序列,以及第二核酸,该第二核酸具有与其天然启动子及5’UTR可操作地连接的第二GLK基因的序列,该第二GLK基因不同于该第一GLK基因,
其中该第一GLK基因以及该第二GLK基因皆为异源的,且相较于未转形的野生型稻米植物,该基因转殖稻米植物表现至少50%增加的枝条生物量及谷物产量。
2.如权利要求1所述的基因转殖稻米植物,其中该第一GLK基因以及该第二GLK基因来自C4植物。
3.如权利要求2所述的基因转殖稻米植物,其中该第一GLK基因为玉米(Zea mays)GLK1,且该第二GLK基因为Zea mays GLK2。
4.如权利要求1所述的基因转殖稻米植物,其中该第一核酸序列包含编码具有如SEQID NO:3所示的胺基酸序列的蛋白质的第一编码序列,以及该第二核酸序列包含编码具有如SEQ ID NO:6所示的胺基酸序列的蛋白质的第二编码序列。
5.如权利要求4所述的基因转殖稻米植物,其中该第一GLK基因的天然启动子及5’UTR具有如SEQ ID NO:1所示的序列,以及该第二GLK基因的天然启动子及5’UTR具有如SEQ IDNO:4所示的序列。
6.如权利要求4所述的基因转殖稻米植物,其中该第一编码序列为SEQ ID NO:2,以及该第二编码序列为SEQ ID NO:5。
7.如权利要求1所述的基因转殖稻米植物,其中相较于该未转形的野生型稻米植物,该基因转殖稻米植物的谷物产量提高50%至95%。
8.一种生产基因转殖稻米植物的方法,该方法包含:
将一重组DNA构筑体引入宿主稻米植物,该构筑体包含第一核酸序列,该第一核酸序列包含与其天然启动子及5’UTR可操作地连接的第一Golden2-like转录因子(GLK)基因,以及第二核酸序列,该第二核酸序列包含与其天然启动子及5’UTR可操作地连接的第二GLK基因,该第二GLK基因不同于该第一GLK基因,该第一GLK基因以及该第二GLK基因为异源的;以及
鉴定基因转殖稻米植物,该基因转殖稻米植物相较于未转形的野生型稻米植物表现至少增加50%的枝条生物量及谷物产量。
9.如权利要求8所述的方法,其中该第一GLK基因以及该第二GLK基因来自C4植物。
10.如权利要求9所述的方法,其中该第一GLK基因为玉米(Zea mays)GLK1,该第二GLK基因为玉米(Zea mays)GLK2。
11.如权利要求8所述的方法,其中该第一核酸序列包含编码具有如SEQ ID NO:3所示的胺基酸序列的蛋白质的第一编码序列,以及该第二核酸序列包含编码具有如SEQ ID NO:6所示的胺基酸序列的蛋白质的第二编码序列。
12.如权利要求11所述的方法,其中该第一GLK基因的天然启动子及5’UTR具有如SEQID NO:1所示的序列,以及该第二GLK基因的天然启动子及5’UTR具有如SEQ ID NO:4所示的序列。
13.如权利要求8所述的方法,其中该第一编码序列为SEQ ID NO:2,以及该第二编码序列为SEQ ID NO:5。
14.如权利要求8所述的方法,其中相较于该未转形的野生型稻米植物,该基因转殖稻米植物的谷物产量提高50%至95%。
15.如权利要求8所述的方法,其中相较于该未转形的野生型稻米植物,该基因转殖稻米植物的枝条生物量提高65%至106%。
16.一种重组DNA构筑体,包含
第一核酸序列,该第一核酸序列包含与其天然启动子及5’UTR可操作地连接的第一Golden 2-like转录因子(GLK)基因,以及第二核酸序列,该第二核酸序列包含与其天然启动子及5’UTR可操作地连接的第二GLK基因,该第二GLK基因不同于该第一GLK基因,其中该第一GLK基因以及该第二GLK基因来自C4植物。
17.如权利要求16所述的重组DNA构筑体,其中该第一GLK基因为玉米(Zea mays)GLK1,且该第二GLK基因为玉米(Zea mays)GLK2。
18.如权利要求16所述的重组DNA构筑体,其中该第一核酸序列包含编码具有如SEQ IDNO:3所示的胺基酸序列的蛋白质的第一编码序列,以及该第二核酸序列包含编码具有如SEQ ID NO:6所示的胺基酸序列的蛋白质的第二编码序列。
19.如权利要求18所述的重组DNA构筑体,其中该第一GLK基因的天然启动子及5’UTR具有如SEQ ID NO:1所示的序列,以及该第二GLK基因的天然启动子及5’UTR具有如SEQ IDNO:4所示的序列。
20.如权利要求18所述的重组DNA构筑体,其中该第一编码序列为SEQ ID NO:2,以及该第二编码序列为SEQ ID NO:5。
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US202063019672P | 2020-05-04 | 2020-05-04 | |
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Citations (3)
Publication number | Priority date | Publication date | Assignee | Title |
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CN1884548A (zh) * | 2006-06-07 | 2006-12-27 | 谢旭亮 | Dna构建体以及其用途 |
US20090241218A1 (en) * | 2006-05-30 | 2009-09-24 | Cropdesign N.V. | Plants having enhanced yield-related traits and a method for making the same |
CN101688215A (zh) * | 2007-06-01 | 2010-03-31 | 克罗普迪塞恩股份有限公司 | 在植物中通过调节GARP转录因子ZmRR10_p的产量增强 |
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US20080148432A1 (en) * | 2005-12-21 | 2008-06-19 | Mark Scott Abad | Transgenic plants with enhanced agronomic traits |
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- 2021-04-29 WO PCT/US2021/029863 patent/WO2021225862A1/en active Application Filing
- 2021-04-29 US US17/923,455 patent/US20230203521A1/en active Pending
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Publication number | Priority date | Publication date | Assignee | Title |
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US20090241218A1 (en) * | 2006-05-30 | 2009-09-24 | Cropdesign N.V. | Plants having enhanced yield-related traits and a method for making the same |
CN1884548A (zh) * | 2006-06-07 | 2006-12-27 | 谢旭亮 | Dna构建体以及其用途 |
CN101688215A (zh) * | 2007-06-01 | 2010-03-31 | 克罗普迪塞恩股份有限公司 | 在植物中通过调节GARP转录因子ZmRR10_p的产量增强 |
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