CN101864429B - 利用玉米Lc基因培育抗棉铃虫植物的方法 - Google Patents
利用玉米Lc基因培育抗棉铃虫植物的方法 Download PDFInfo
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Abstract
本发明公开了玉米Lc基因在培育抗棉铃虫植物上的应用以及培育抗棉铃虫植物的方法，尤其适于在棉花植物上的应用。本发明转玉米Lc基因棉花植株对棉铃虫的抗性强，为抗棉铃虫育种提供了一种新的途径；其次，和使用杀虫剂相比，本发明方法成本低，且不会造成农药残留和环境污染。
Description
技术领域
本发明属于植物基因工程领域，具体地说涉及利用玉米Lc基因培育抗棉铃虫的转基因植物的方法及相关的转基因植物。
背景技术
棉花是我国第一大经济作物，在农业生产及整个国民经济中占有极其重要的战略地位和经济价值，随着棉花生产的迅猛发展，棉花枯黄萎病、棉铃虫、蚜虫、叶螨等病虫害越来越重，其中棉铃虫(Helicoverpa armigera Hubner)是危害中国棉花生产的最主要害虫，对我国棉花生产构成巨大威胁，常年损失达10～15％(王仁祥，作物研究2001，棉花专缉：6-9)。全国每年用于棉花害虫防治的杀虫剂用量约占杀虫剂使用总量的2/3，虽然对减轻虫害具有明显的作用，但也带来了严重的社会问题，如棉花的生产成本急剧增加，严重危害棉农的健康及生命安全，棉铃虫抗药性增强，环境污染，生态失衡等，棉田害虫的防治越来越困难。转基因抗虫棉的培育成为解决此类问题最经济最有效的途径之一(汪新业等.种子科技2003，3：156-157)。
目前使用最广泛也是最有效的抗虫基因是苏云金芽孢杆菌毒素蛋白基因(Bt基因)，应用于棉花的主要有CryIA(b)、CryIA(c)、CryIIA和CryIVA等几种，以培育抗棉铃虫的转基因棉花。经改良的CryIA(c)和CryIA(b)的表达量可达到棉株中可溶性蛋白的0.1％和0.05％，能有效地杀死棉铃虫、红铃虫等鳞翅目害虫。然而，我国所推广的针对棉铃虫的转基因抗虫棉，几乎都是涉及Bt及其相关的基因，存在很大的隐患。主要是：(1)Bt抗虫棉的杀虫活性主要在一代和二代，而在棉铃虫的第三、第四代时明显降低；(2)Bt抗虫棉抗性比较单一，只对棉铃虫、红铃虫等少数鳞翅目害虫有效果，而对棉田其它害虫没有抗性；(3)棉铃虫对Bt抗虫棉易产生抗性，存在Bt抗虫棉失效和Bt生物农药失效的巨大隐患，试验研究推测，大田连续种植8～10年后，转Bt基因棉可能丧失对棉铃虫的抗性，从而失去利用价值(汪若海等.生物技术通报2000，5：1-6；王仁祥，作物研究2001，棉花专缉：6-9)。因此，应积极筛选新的抗虫基因，培育高效广谱的转基因抗虫棉花。
花青素是植物果皮和种皮颜色的主要色素物质，是花瓣从红色、紫色到蓝色的主要呈色物质，是类黄酮的重要组成部分。花青素的生物合成途径有近20步化学反应，由两组基因调控，第一组大约涉及15个结构基因，编码催化花青素生化反应的多种酶类，第二组是调节基因，其编码的转录因子或属于MYB-型C1家族，或是碱性bHLH MYC-型R家族(HK.Dooner等.Annual Review of Genetics1991，25：173-199)，调控结构基因的时空表达。玉米C1基因在胚乳的糊粉层中调节花青素着色与否(KC.Cone等.Proc Natl Acad Sci U S A.1986，83：9631-9635)，而R基因调控花青素着色的时空分布(SR.Ludwig等.Proc NatlAcad Sci U S A.1989，86(18)：7092-7096)。Lc(lear color，简称Lc)基因属R基因家族，是玉米花青素生物合成途径中的一个调节基因，编码大约610个氨基酸的蛋白，典型的bHLH MYC型转录因子(SR.Ludwig等.Proc Natl Acad SciU S A.1989，86(18)：7092-7096)。在玉米中，Lc基因参与调节花青素生物合成途径中多个结构基因如C2基因的表达，调控花青素着色的数量、分布和时间。组成型表达单个Lc基因足以激活花青素生物合成途径，而且还与类黄酮化合物生物合成相关联，对植物育性和抵抗病虫害起到重要作用(HK.Dooner等.AnnualReview of Genetics 1991，25：173-199)，比如，转Lc基因苹果植株中，PAL、CHS、FHT、DFR、LAR、ANS和ANR的mRNA水平得以提高，叶片中花青素越橘花青苷(anthocyanin idaein)上升12倍，黄烷-3-表儿茶酸(flavan 3-olepicatechin)上升14倍，同分异构儿茶酸(isomeric catechin)上升41倍等(H.Li等.Planta 2007，226：1243-1254)；表达玉米C2基因的转基因水稻，稻瘟病抗性增强(M.Gandikota等.Molecular Breeding 2001，7：73-83)。
经检索，没有发现有关玉米Lc基因在抗棉铃虫植物上应用的报道.
发明内容
本发明的目的在于提供玉米Lc基因在培育抗棉铃虫植物上的用途。
本发明另一目的在于提供含有玉米Lc基因的转基因植物、组织或细胞。
本发明第三目的在于提供含有玉米Lc基因的转基因棉花、棉花组织或棉花细胞。
本发明第四目的在于提供利用玉米Lc基因培育抗棉铃虫植物的方法。
为实现上述目的，本发明的技术方案如下：
玉米Lc基因在培育抗棉铃虫植物上的应用，其中所述的玉米Lc基因编码由SEQ ID No：2所示的氨基酸序列组成的蛋白质。
上述应用中所述的玉米Lc基因由SEQ ID No：1所示的核苷酸序列组成；或者由与SEQ ID No：1所示的核苷酸序列具有95％以上同源性并编码相同功能蛋白的核苷酸序列组成。
所述的玉米Lc基因为玉米叶色基因(leaf color gene)，在Genbank中的编号为ID M26227。
上述应用中所述的植物是指棉花。
含有所述的外源玉米Lc基因的植物、组织或细胞。
含有所述的外源玉米Lc基因的棉花、棉花组织或棉花细胞。
利用玉米Lc基因培育抗棉铃虫植物的方法，将玉米Lc基因克隆到质粒载体上并导入农杆菌菌株，再经农杆菌介导转入植物，可得抗棉铃虫的转基因植物。
上述方法中所述的质粒载体可以是pBI121。
上述方法中所述的农杆菌菌株可以是LBA4404。
利用玉米Lc基因培育抗棉铃虫植物的方法，包括如下步骤：
(1)将外源的玉米Lc基因转入植物细胞或组织，获得含有外源玉米Lc基因的植物细胞或组织；
(2)将含有外源玉米Lc基因的植物细胞或组织，再生成植物植株，即得抗棉铃虫的植物。
上述方法中所述的植物优选棉花。
在一优选例中，在步骤(1)中，将外源的玉米Lc基因转入棉花细胞或组织。
在另一优选例中，在步骤(1)中，用携带玉米Lc基因表达载体的农杆菌感染棉花下胚轴。
利用上述所得的转玉米Lc基因棉花培育棉花品种的方法，以上述转Lc基因棉花为亲本之一，利用回交或者杂交的方法，可培育出抗棉铃虫的棉花品种。
本发明的优点：(1)本发明转玉米Lc基因棉花植株对棉铃虫的抗性强，为抗棉铃虫育种提供了一种新的途径；(2)和使用杀虫剂相比，本发明方法成本低，且不会造成农药残留和环境污染。
附图说明
图1.转玉米Lc基因棉花T0代植株Southern杂交印迹图。
图2.转玉米Lc基因棉花T1代植株Southern杂交印迹图。
图3.转玉米Lc基因棉花表型照片，其中A为转玉米Lc基因愈伤组织，B为转玉米Lc基因体细胞胚，C为转玉米Lc基因植株叶片，D为转玉米Lc基因植株茎杆，E为转玉米Lc基因植株花蕾，F为转玉米Lc基因植株花药，G、H为转玉米Lc基因植株叶片栅栏细胞，I为黑暗条件下培养一周转玉米Lc基因胚珠，J、K为光照条件下培养一周转玉米Lc基因胚珠，L为转基因T1代幼苗中Lc基因遗传分离，阳性苗表现出红色根。
图4.转玉米Lc基因棉花植株叶片粗提物光谱分析曲线图。
图5.转玉米Lc基因棉花植株叶片花青素含量柱形图；其中1为转玉米Lc基因植株干叶，2为野生型植株干叶，3为转玉米Lc基因植株鲜叶，4为野生型植株鲜叶。
图6.转玉米Lc基因棉花植株叶片花青素成分含量柱形图；其中1为转玉米Lc基因植株干叶，2为野生型植株干叶，3为转玉米Lc基因植株鲜叶，4为野生型植株鲜叶。
图7.转玉米Lc基因叶片和非转基因叶片同时饲喂棉铃虫幼虫1天后叶片照片，其中a1、c1：转玉米Lc基因叶片，b1：非转基因叶片。
图8.转玉米Lc基因叶片和非转基因叶片同时饲喂棉铃虫幼虫3天后照片，其中A2、c2：转Lc基因叶片，b2：非转基因叶片。
图9.转玉米Lc基因叶片和非转基因叶片分别饲喂棉铃虫幼虫5天后照片，其中d：转玉米Lc基因红色叶片，e：转玉米Lc基因绿色叶片，f、g：非转基因叶片。
图10.转玉米Lc基因叶片和非转基因叶片分别饲喂棉铃虫幼虫5天后幼虫体重柱形图；其中1为转玉米Lc基因植株叶片喂养，2为野生型植株叶片喂养。
具体实施方式
以下通过实施例对本发明做进一步的说明，但是不构成对本发明的任何限制。如无特别说明，实施例中所涉及的方法等皆为本领域技术人员所熟知的方法，可参见《分子克隆实验指南》等。
实施例1转玉米Lc基因棉花的培育
通过美国国家生物技术信息中心(National Center for BiotechnologyInformation http://www.ncbi.nlm.nih.gov)，检索获取玉米Lc基因序列数据，设计如下引物：
LcB-F：5’-ccggatccatggcgctttcagcttcccg-3’(SEQ ID No：3)，
LcH-R：5’-ggaagctttcaccgcttccctatagctttg-3’(SEQ ID No：4)，
选取玉米授粉后10天的雌穗，用热酚法(参见XB Li等.Plant Cell 2005，17：859-875)提取总RNA，根据QIANGEN一步RT-PCR方案做反转录，即RNA模板加入到反应液中，50℃下反转录30分钟，之后灭活反转录酶(QIANGEN公司产品使用指南)，cDNA模板95℃变性15分钟，以LcB-F和LcH-R为引物PCR扩增玉米Lc基因编码区cDNA片段，反应程序为94℃45秒、60℃45秒、72℃2分钟，计30循环。所获cDNA片段经Bam HI/Hind III处理后，-20℃保存备用。质粒pBI121用BamHI/SacI切开，连接上一个：
Hind III-SacI接头：5’-AGCTTGAGCT-3’(SEQ ID No：5)，
与前述玉米Lc基因cDNA连接，经测序(北京三博志远基因技术有限公司)确认玉米Lc基因(序列见SEQ ID No：1)正确克隆到35S启动子下游，得到植物表达载体pBI-Lc。采用电击法将表达载体pBI-Lc导入农杆菌菌株LBA4404。
将棉花种子(珂字312，Coker312)用70％乙醇处理1分钟、10％双氧水处理1-2小时进行表面消毒，放置在1/2MS固体培养基(组成成份及其比例为：1/2MS无机盐、1/2B5维生素、1/2铁盐、0.5％葡萄糖、0.5g/L MgCl2·6H2O、0.25％phytagel、pH6.0)上，在25～28℃、14小时光照/10小时黑暗的条件下培养发芽。挑取含有质粒pBI-Lc的农杆菌LBA4404单菌落，放入10ml LB液体培养基中，28℃摇床中振荡培养3天，取2ml菌液5000rpm离心3分钟，去除上清液，加入1ml MS共培养基重悬菌体，吸取适量重悬后的菌液，加入到50ml液体MS共培养基中，使OD值约0.1。选取发芽6～9天的棉花幼苗，将其下胚轴切成5mm的小段，浸入上述处理好的农杆菌LBA4404菌液中15min，之后将外植体转至固体MS共培养基(组成成份及其比例为：MS无机盐、维生素B5、铁盐、3.0％葡萄糖、0.1mg/L KT、0.05mg/L 2，4-D、100umol/L乙酰丁香酮、0.5g/L MgCl2·6H2O、0.25％phytagel、pH6.0)上，在25～28℃、14小时光照/10小时黑暗的条件下共培养3天。将外植体转到筛选培养基(组成成份及其比例为：MS无机盐、B5维生素、铁盐、3.0％葡萄糖、0.1mg/L KT、0.05mg/L 2，4-D、0.5g/L MgCl2·6H2O、0.3％phytagel、pH6.0，附加头孢霉素500mg/L、潮霉素5mg/L)上诱导愈伤组织，3～4周继代培养一次。将抗性愈伤组织转至体细胞胚诱导培养基(组成成份及其比例：MS无机盐、B5维生素、铁盐、3.0％葡萄糖、0.5g/L MgCl2·6H2O、1.9g/L KNO3、1.0g/L谷氨酰胺、0.5g/L天门冬酰胺、0.3％phytagel、pH6.0，附加头孢霉素500mg/L、潮霉素5-10mg/L)，在25～28℃、14小时光照/10小时黑暗的条件下继代培养直到长出胚状体。将成熟体细胞胚转至壮苗培养基(组成成份及其比例：无NH4NO3的MS无机盐、B5维生素、铁盐、3.0％葡萄糖、0.5g/L MgCl2·6H20、1.9g/L KNO3、0.3％phytagel、pH6.0，附加头孢霉素500mg/L、潮霉素5-10mg/L)培养直到长出幼芽和根。将2-4cm并长出真叶的幼苗转到生根培养基(组成成份及其比例：1/2MS无机盐、1/2B5维生素、1/2铁盐、0.5％葡萄糖、0.1mg/L IAA、0.1mg/L KT、0.25％phytagel、pH6.0)上，25～28℃、14小时光照/10小时黑暗的条件下培养4周，得到转化再生植株并移栽到温室中。共获得陆地棉(Gossypium hirsutum，Coker 312)6个独立的转玉米Lc基因棉花株系(株系编号分别为：L28、L82、L73、L74、L143、L144)。
实施例2转玉米Lc基因棉花中玉米Lc基因的分子检测
CTAB方法提取实施例1中所得的6个转玉米Lc基因的T0代植株基因组DNA(L.Wang.Plant Cell 1998，10：1733-46)，以下述序列为引物：
Lc-F：5’-atggcgctttcagcttcccgagt-3’(SEQ ID No：6)，
Lc-R：5’-tcaccgcttccctatagctttgc-3’(SEQ ID No：7)，
PCR扩增Lc基因片段，制备Southern杂交探针，50ug基因组DNA用限制性内切酶BamHI 37℃酶切过夜，琼脂糖凝胶电泳后转移至Hybond N+尼龙膜上(具体方法参见Amersham公司产品使用指南)，地高辛-11-dUTP标记Lc基因探针，42℃下与尼龙膜杂交过夜(Roche公司产品使用指南)，结果表明，转基因T0代植株中，L82含3个Lc基因拷贝，L73、L74、L143、L144含单拷贝Lc基因，L28含6个Lc基因拷贝(见图1)。
将6个转基因T0代植株盆栽种植于温室，单株自交后收获T1代种子，并种植于相同温室中，每株系种植20株。CTAB方法提取L143T1代植株基因组DNA，50ug基因组DNA用限制性内切酶BamHI 37℃酶切过夜，琼脂糖凝胶电泳后转移至Hybond N+尼龙膜上(Amersham公司产品使用指南)，地高辛-11-dUTP标记Lc基因探针，42℃下与尼龙膜杂交过夜(Roche公司产品使用指南)，结果显示，外源玉米Lc基因能稳定传递到T1代，并表现为单基因位点遗传(见图2)。
实施例3转基因棉花植株中Lc基因表达鉴定
热酚法(参见XB Li等.Plant Cell 2005，17：859-875)提取实施例1所得的转基因植株嫩叶、茎、花器官包括花瓣、子房、花药和萼片的总RNA，根据QIANGEN一步RT-PCR方案做反转录，即RNA模板加入到反应液中，50℃下反转录30分钟，之后灭活反转录酶(QIANGEN公司产品使用指南)，cDNA模板95℃变性15分钟，前述Lc基因引物Lc-F(SEQ ID No：6)和Lc-R(SEQ ID No：7)做PCR扩增，反应程序为94℃45秒、56℃45秒、72℃2分钟，计30循环，同时PCR扩增肌动蛋白基因(Actin基因)作为内标，所用引物为：
Actin-F：5’-tttgctggtgatgatgctcc-3’(SEQ ID No：8)
Actin-R：5’-ctccaatccagacactgtact-3’(SEQ ID No：9)。
RT-PCR分析显示，Lc基因在转基因棉花T0代植株叶片和花器官中均有表达，而野生型棉花植株中检测不到Lc基因表达；不同组织器官中Lc基因表达水平稍有差异，花瓣中未检测到Lc基因表达，叶片中Lc基因持续高水平表达，4～16天子房中Lc基因表达水平最高，7～13天的棉纤维中Lc基因中度表达，而23天及以后的成熟棉纤维细胞中，几乎检测不到Lc基因表达。
实施例4转化棉花细胞和转基因棉花植株表型观测
玉米Lc基因转化棉花过程中，观察到卡那抗性愈伤组织和体细胞胚显现红色或紫色，体细胞成熟胚中，胚根、胚轴和子叶有花青素积累(见图3.A、B)。转基因幼苗的绿色真叶同样积累花青素，根保持红色。田间生长的转基因T0代植株，幼嫩叶片通常表现为红色或紫色斑块，间有绿色区域，茎、萼片及花药明显有着色，而花瓣无着色(见图3.C、D、E、F)。转基因T0代植株叶片经徒手切片观察到，花青素积累在叶片的栅栏细胞及维管中，表皮细胞中未发现有花青素(见图3.G、H)。非转基因植株衰老叶片保持黄颜色，转基因植株老叶片则变为红色。上述着色方式可以遗传到T1和T2代，并与外源Lc基因共分离(见图3.L)。转基因植株及其后代(T1和T2)未发现形态变化。
实施例5转玉米Lc基因棉花胚珠离体培养观察
实施例1中转Lc基因棉花T1代植株开花后2天的棉铃，70％乙醇灭菌5-10分钟后，取幼嫩胚珠放入液体培养基(CA Beasley等.Am.J.Bot.61：188-194)中离体培养一周，光照条件下棉纤维细胞转为深红色，且在中央液泡中积累红色或紫色油状液态物(见图3.J、K)，而在黑暗条件下棉纤维细胞保持白色(见图3.I)，表明玉米Lc基因参与花青素合成受光照调控。
实施例6转Lc基因棉花植株中花青素含量测定
选取实施例1中转Lc基因棉花植株(L143)的新鲜或风干叶片材料提取花青素(MG Choung等J Agric.Food.Chem.2001，49：5848-5851)，pH差减法处理提取样品(J Lee等J Ass.Off.Ana.Chem.Inter.2005，88：1269-1278)，采用UV-VIS分光光度计进行定量分析(ZZ Gong等Plant Mol.Biol.1997，35：915-927)，花青素含量计为氰化-3-葡糖苷，依照Lambert-Beer’s定律计算(A＝εCL，A-吸收峰值，C-浓度，L-光路长度以厘米计，ε-摩尔消光系数，为26900L/mol；摩尔质量取449g/mol)。对比非转基因棉花，转Lc基因棉花植株叶片提取物呈现深红色，光谱分析显示，转Lc基因叶片粗提物分别在210-230nm、280nm、510nm波长处吸收峰有明显增加(见图4)。pH差减法(JLee等.J Ass.Off.Ana.Chem.Inter.2005，88：1269-1278)定量分析花青素含量，结果转玉米Lc基因植株叶片中氰化-3-葡糖苷达53.6mg/100g(干重)，为野生型棉花的3.5倍(见图5)。转Lc基因植株干叶中，单体花青素占总花青素的27.2％、多聚体花青素占43.8％、其它占29.0％，非转基因植株干叶中分别为17.3％、67.2、15.5％(见图6)。说明转Lc基因棉花植株中Lc基因的表达显著促进了棉花叶片中总花青素的产生量。
实施例7转玉米Lc基因棉花植株棉铃虫抗性鉴定
摘取实施例1中转玉米Lc基因棉花的T1代植株(L143-5、L143-6、L143-7)与相应的非转基因棉株的第四片叶，清水冲洗晾干后喂食三龄棉铃幼虫。
试验1：将转玉米Lc基因棉株叶与非转基因棉株叶放入同一个培养皿中，放入3条棉铃幼虫饲喂1-3天，重复10皿；
试验2：将转玉米Lc基因棉株叶与非转基因棉株叶分别放在不同的皿中，每皿放入3条相同大小、相同龄期的棉铃幼虫饲喂5天，重复13对皿。
结果，同时饲喂转玉米Lc基因和非转基因叶片时，棉铃虫幼虫偏向取食非转基因叶片，饲喂1天后，转玉米Lc基因叶片保持完整，非转基因叶片则被取食(见图7)，3天后这一结果更加明显(见图8)，转玉米Lc基因棉株叶片被取食通常是叶片的绿色部分而非红色区块。胁迫饲喂试验中，棉铃虫幼虫在转玉米Lc基因叶片上几乎不取食(见图9)，幼虫体重明显低于非转基因叶片饲喂的幼虫(见图10)。以上结果说明表达Lc基因的转基因棉花具有较好的棉铃虫抗性。
序列表
<110>中国科学院遗传与发育生物学研究所
<120>利用玉米Lc基因培育抗棉铃虫植物的方法
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accgagtggt actacgtggt ctccatgacc tacgccttcc ggccaggcca agggttgccc 420
ggcaggagtt tcgcgagcga cgagcatgtc tggctgtgca acgcgcacct cgccggcagc 480
aaagccttcc cccgcgcgct cctggccaag agcgcgtcca ttcagtcaat cctctgcatc 540
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ggacgagcaa acgagaccgg cgaggccgca gcagacgacg gcacgtttgc gttcgaggaa 720
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cacatcacca aggagatcga ggagttctac agcctctgcg acgaaatgga cctgcaggcg 900
ctaccactac cgctagagga cggctggacc gtggacgcgt ccaatttcga ggtcccctgc 960
tcttccccgc agccagcgcc gcctccggtg gacagggcta ccgctaacgt cgccgccgac 1020
gcctcaaggg cacccgtcta cggctctcgc gcgacgagtt tcatggcttg gacgaggtcc 1080
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ggcgcgacgg gagcagcaca ggaaatgagt ggcactggca ccaagaacca cgtcatgtcg 1260
gagcgaaagc gacgagagaa gctcaacgag atgttcctcg tcctcaagtc actgcttccg 1320
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acgacgacaa ggctaataac aaggccctcc cgtggcaata atgagagtgt gaggaaggag 1500
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Claims (4)
1.玉米Lc基因在培育抗棉铃虫棉花上的应用，其中所述的玉米Lc基因编码由SEQ ID No：2所示的氨基酸序列组成的蛋白质。
2.按照权利要求1所述的玉米Lc基因在培育抗棉铃虫棉花上的应用，其特征在于所述的玉米Lc基因由SEQ ID No：1所示的核苷酸序列组成。
3.利用玉米Lc基因培育抗棉铃虫棉花的方法，其特征在于包括如下步骤：
(1)将外源的玉米Lc基因转入棉花细胞或组织，获得含有外源玉米Lc基因的棉花细胞或组织；所述的玉米Lc基因由SEQ ID No：1所示的核苷酸序列组成；所述的棉花组织是指棉花下胚轴的细胞或组织；
(2)将上述含有外源玉米Lc基因的棉花细胞或组织，再生成棉花植株，即得抗棉铃虫的棉花。
4.利用转玉米Lc基因棉花培育棉花品种的方法，其特征在于以权利要求3利用玉米Lc基因培育抗棉铃虫棉花的方法中所获得的抗棉铃虫棉花为亲本之一，利用回交或者杂交的方法，培育出抗棉铃虫的棉花品种；所述的玉米Lc基因由SEQ ID No：1所示的核苷酸序列组成。
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| CN103387995A (zh) * | 2013-07-27 | 2013-11-13 | 山西省农业科学院棉花研究所 | 玉米Lc基因作为筛选标记在棉花转基因育种中的应用 |
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| CN103387995A (zh) * | 2013-07-27 | 2013-11-13 | 山西省农业科学院棉花研究所 | 玉米Lc基因作为筛选标记在棉花转基因育种中的应用 |
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