CN111171127A - 紫云英lhy基因及其应用 - Google Patents
紫云英lhy基因及其应用 Download PDFInfo
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- CN111171127A CN111171127A CN202010120702.9A CN202010120702A CN111171127A CN 111171127 A CN111171127 A CN 111171127A CN 202010120702 A CN202010120702 A CN 202010120702A CN 111171127 A CN111171127 A CN 111171127A
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- lhy
- plant
- lhy gene
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Abstract
本发明提供紫云英LHY基因及其应用。紫云英LHY基因及其编码蛋白的序列见SEQ ID NO:1和2。本发明首次从紫云英中克隆得到LHY基因，并通过农杆菌介导的遗传转化方法，在拟南芥中验证了LHY基因具有调控植物开花的功能，具体为延迟植株开花。转LHY基因的拟南芥株系出现抽薹提前，开花提前，抽薹天数比野生型平均晚18.36天，开花天数比野生型平均晚20.72天。这表明紫云英LHY基因与成花关系密切，其具有调节开花时间的功能。将该基因应用于植物性状改良，具有良好的应用前景。本发明提供的LHY基因及其编码蛋白为培育植物新品种提供了宝贵资源。
Description
技术领域
本发明涉及基因工程和植物遗传育种领域，具体地说，涉及紫云英LHY基因及其应用。
背景技术
紫云英(Astragalus sinicus L.)是豆科，黄耆属二年生草本植物，主要分布于中国长江流域，是重要的蜜源植物。紫云英对土壤资源的可持续利用，具有重要意义。
成花是植物从营养生长向生殖生长的关键转换，同时也是是实现世代交替的中心环节，在很大程度上决定了植物繁殖的成功与否。紫云英按开花早晚可分特早花型、早花型、中花型和晚花型，开花时间直接决定了有效生长季的长短。作为绿肥，紫云英的种植和收获时间取决于主作物，保证紫云英产量的同时还要与主作物的生育期相协调。在合适的时间促使或避免开花，根据当地的耕作制度选育花期适当的品种是重要的育种目标。因此，成花基因研究对紫云英生长发育及遗传改良具有重要意义。但对于紫云英成花机制的研究，以及开花相关基因在此过程中的作用尚不清楚。
发明内容
本发明的目的是提供紫云英LHY基因及其应用。
为了实现本发明目的，第一方面，本发明提供紫云英LHY基因，LHY基因为编码如下蛋白质(a)或(b)的基因：
(a)由SEQ ID NO:2所示的氨基酸序列组成的蛋白质；
(b)SEQ ID NO:2所示序列经取代、缺失或添加一个或几个氨基酸且具有同等功能的由(a)衍生的蛋白质。
本发明利用RACE技术获得LHY基因的全长cDNA序列大小为2865bp(SEQ ID NO:1)，含有一个2259bp的开放阅读框，5’和3’非编码区的长度分别为407bp和199bp。PolyA尾巴信号肽区域位于2854-2859bp处。
第二方面，本发明提供含有所述LHY基因的生物材料，所述生物材料包括但不限于重组DNA、表达盒、转座子、质粒载体、噬菌体载体、病毒载体或工程菌或非可再生的植物部分。
第三方面，本发明提供所述LHY基因或含有该基因的生物材料在调控植物开花时间中的应用。
所述调控是指延迟开花时间。
前述应用包括：
1)使植物包含LHY基因；或者，
2)使植物过表达LHY基因。
所述应用包括但不限于转基因、杂交、回交、自交或无性繁殖。
可选地，过表达LHY基因的方法选自以下1)～4)，或任选的组合：
1)通过向植物中导入具有LHY基因的质粒；
2)通过增加植物染色体上LHY基因的拷贝数；
3)通过将强启动子与LHY基因可操作地连接；
4)通过导入增强子。
本发明中，所述植物为拟南芥、紫云英。
进一步地，所述应用包括利用Ti质粒、植物病毒载体、直接DNA转化、显微注射、基因枪、电导或农杆菌介导等方法将包含LHY基因的重组表达载体导入植物，获得转基因植株。
第四方面，本发明提供所述LHY基因或含有该基因的生物材料在植物育种中的应用。育种目的为延迟开花时间。
借由上述技术方案，本发明至少具有下列优点及有益效果：
本发明首次从紫云英中克隆得到LHY基因，并通过农杆菌介导的遗传转化方法，在拟南芥中验证了LHY基因具有调控植物开花的功能，具体为延迟植株开花。转LHY基因的拟南芥株系出现抽薹延迟，开花延迟，抽薹天数比野生型平均晚18.36天，开花天数比野生型平均晚20.72。这表明紫云英LHY基因与成花关系密切，其具有调节开花时间的功能。将该基因应用于植物性状改良，具有良好的应用前景。本发明提供的LHY基因及其编码蛋白为培育植物新品种提供了宝贵资源。
附图说明
图1为本发明实施例1中紫云英LHY基因编码蛋白的氨基酸序列与其它九个物种的序列同源性比对结果。
图2为本发明实施例1中LHY蛋白质二级结构预测及功能位点注释。
图3为本发明实施例1中利用MEGA 5.2构建的LHY基因氨基酸序列的NJ系统进化树(数字为置信度)。
图4为本发明实施例1中预测的LHY蛋白的三维结构图及其与模板碳主链的重叠图。其中，A：紫云英LHY的三维结构图。B：紫云英LHY结构和质型多角体病毒RNA聚合酶蛋白的碳主链的重叠图。彩色结构的是紫云英LHY；紫色线是质型多角体病毒RNA聚合酶碳主链。
图5为本发明实施例1中紫云英LHY基因5-RACE PCR电泳结果分析。其中，M:DL5000Marker：100，250，500，750，1000，1500，2000，3000，5000；1：LHY 5-RACE PCR电泳结果(约750bp)。
图6为本发明实施例1中紫云英LHY基因3-RACE PCR电泳结果分析。其中，M:DL5000Marker：100，250，500，750，1000，1500，2000，3000，5000；1：LHY 3-RACE PCR电泳结果(约580bp)。
图7为本发明实施例1中紫云英LHY基因3端第一步移基因PCR扩增结果。其中，M:DL5K DNA marker：100，250，500，750，1000，15000，2000，3000，5000bp；1：LHY3端第一步移基因PCR扩增产物(约2000bp)。
图8为本发明实施例2中转基因(紫云英LHY基因)拟南芥的表型分析结果。其中，左侧为野生型，右侧为转基因植株。
具体实施方式
以下实施例用于说明本发明，但不用来限制本发明的范围。若未特别指明，实施例均按照常规实验条件，如Sambrook等分子克隆实验手册(Sambrook J&Russell DW,Molecular Cloning:a Laboratory Manual,2001)，或按照制造厂商说明书建议的条件。实施例1紫云英LHY基因的克隆及序列分析
本发明利用RACE技术从紫云英中获得LHY基因的全长cDNA序列大小为2865bp(SEQID NO:1)，含有一个2259bp的开放阅读框，5’和3’非编码区的长度分别为407bp和199bp。PolyA尾巴信号肽区域位于2854-2859bp处。其编码蛋白的氨基酸序列如SEQ ID NO:2所示。具体方法如下：
1、试验材料：新鲜紫云英组织样本。
2、RNA提取：紫云英总RNA提取采用RNeasy Plant Mini Kit(Qiagen货号：74904)进行，具体提取过程参照试剂盒说明书。
3、5-RACE-PCR实验
5-RACE模板采用GeneRacerTM Kit(Invitrogen，货号：L1500-01)进行合成。
(1)引物设计及序列
采用Primer Premier 6.0软件进行5-RACE引物的设计，由生工生物工程(上海)股份有限公司进行合成。引物序列见表1。
表1 5-RACE引物序列
(2)5-RACE PCR反应体系及条件见表2～表5。
表2 5-RACE PCR第一轮反应体系
表3 5-RACE PCR第一轮反应条件
表4 5-RACE PCR第二轮反应体系
表5 5-RACE PCR第二轮反应条件
(3)5-RACE PCR电泳结果
PCR结束后，采用1.5％琼脂糖凝胶电泳进行分析(结果见图5)，5-RACE PCR呈现出一条特异性条带，大小约750bp左右，切胶回收后，连接pUCm载体，转化高效化学感受态细胞DH5α，送往生工生物工程(上海)股份有限公司进行测序。
4、3-RACE-PCR实验
3-RACE模板采用GeneRacerTM Kit(Invitrogen，货号：L1500-01)进行合成。
(1)引物设计及序列
利用客户提供的序列，采用Primer Premier 6.0软件进行3-RACE引物的设计，由生工生物工程(上海)股份有限公司进行合成。引物序列见表6。
表6 3-RACE引物序列
(2)3-RACE PCR反应体系及条件见表7～表10。
表7 3-RACE PCR第一轮反应体系
表8 3-RACE PCR第一轮反应条件
表9 3-RACE PCR第二轮反应体系
表10 3-RACE PCR第二轮反应条件
(3)3-RACE PCR电泳结果
PCR结束后，采用1.5％琼脂糖凝胶电泳进行分析(结果见图6)，3-RACE PCR呈现出一条特异性条带，大小约580bp左右，切胶回收后，连接pUCm载体，转化高效化学感受态细胞DH5α，送往生工生物工程(上海)股份有限公司进行测序。
5、紫云英LHY基因3端第一步移基因克隆
目的基因PCR扩增及回收。
(1)模板：紫云英3-RACE为模板。
(2)引物序列见表11。
表11引物序列
(3)PCR扩增体系及条件见表12～表13。
表12 PCR扩增反应体系
注：Invitrogen，货号：12532-016。
表13 PCR扩增反应条件
(4)PCR扩增结果
PCR结束后，采用1.0％琼脂糖凝胶电泳进行分析(结果见图7)，PCR产物呈现出一条特异性条带，大小约2000bp左右，切胶回收后，连接pUCm-T载体，转化高效化学感受态细胞DH5α，送往生工生物工程(上海)股份有限公司进行测序。
紫云英LHY基因的生物信息分析：
紫云英LHY基因编码蛋白的氨基酸序列与其它九个物种(Cicer arietinum,Medicago truncatula,Glycine soja,Glycine max,Phaseolus vulgaris,Castaneasativa,Prunus persica,Vitis vinifera,Citrus sinensis)序列同源性比对结果见图1。
使用ProtParam进行蛋白质氨基酸含量和理化性质分析。LHY基因编码的蛋白质含有752个氨基酸，其分子量为82800.03道尔顿，理论等电点为5.88，其化学组分见表14。该蛋白含有95个带负电荷的氨基酸残基和84个带正电荷的氨基酸残基，它的N端以蛋氨酸起始。该蛋白共含有11486个原子，其中碳原子3578个，氢原子5672个，氮原子1030个，氧原子1182个，硫原子24个，其脂肪族指数为62.97，疏水性指数为-0.752，不稳定性指数为56.29。综合以上数据，该蛋白被归类为不稳定性蛋白。
表14 LHY基因编码蛋白的化学组分
应用CLC Genomics Workbench进行LHY蛋白的二级结构预测，该蛋白由752个氨基酸组成，包含30个α螺旋和8个β折叠。氨基酸序列中的酰胺化位点、蛋白激酶C磷酸化位点和N-糖基化位点见图2。利用NetPhos预测蛋白质中丝氨酸、苏氨酸和络氨酸的磷酸化位点，在LHY蛋白质中共预测出39个丝氨酸磷酸化位点，17个苏氨酸磷酸化位点和5个络氨酸磷酸化位点(表15)。
表15磷酸化位点预测结果
应用BLASTP程序对紫云英的LHY蛋白与其他9个物种同源蛋白进行了比较分析，结果显示物种之间有一定的相似性。其中，紫云英与Medicago truncatula的LHY蛋白的相似度达到78.96％，与Cicer arietinum和Glycine soja LHY蛋白的相似度分别达到了76.66％和75.17％，这一结果表明紫云英LHY蛋白与这三个物种有一定的同源性。用LHY蛋白构建的进化树显示紫云英与Cicer arietinum和Medicago truncatula聚为一类(图3)。
紫云英LHY蛋白与其他物种LHY蛋白的比较分析结果见表16。包括氨基酸数量，相似度，E值，等电点和分子量等指标。
表16紫云英LHY蛋白与其他物种LHY蛋白的比较分析
利用MEGA 5.2软件构建LHY基因氨基酸序列的NJ系统进化树，bootstrep设置为1000，采用Jones-Thorton-Taylor模型构建NJ树。建树所涉及到的基因的序列号为：Cicerarietinum:NP_001296649.1,Medicago truncatula:AES82836.2,Glycine soja:KHN16397.1,Glycine max:XP_006604920.1,Phaseolus vulgaris:CAD12767.2,Castaneasativa:AAU20773.1,Prunus persica:XP_007218929.1,Vitis vinifera:XP_010661511.1,Citrus sinensis:XP_006491389.1。
紫云英LHY和质型多角体病毒RNA聚合酶分别有752和1225个氨基酸组成。质型多角体病毒RNA聚合酶蛋白在PDB数据库里的ID为3ja4。紫云英LHY蛋白的三维结构是以3ja4为模板在I-TASSER软件中预测得到的(图4)。预测的蛋白结构C值为-0.36，表示其与3ja4的A链在折叠和二级结构方面非常类似。显示目标蛋白和模板蛋白的结构相似性的TM值为0.968，RMSD值为
实施例2 LHY基因在紫云英各组织的相对表达水平检测
1、总RNA提取
紫云英均为温室种植的植物材料，是用普通紫云英植株材料采集种子培养而成的新鲜紫云英组织样本。
2、荧光定量PCR引物设计和合成
采用Primer Premier 6.0和Beacon designer 7.8软件进行定量PCR引物设计，然后由生工生物工程(上海)股份有限公司合成。所用内参基因为18S rDNA(GenBank：AF359603.1)。
3、Real-Time PCR(Q-PCR)基因表达差异统计分析
每个样品重复三次，各个基因的相对表达水平以2(Ct内参基因－Ct目的基因)进行统计分析。LHY基因Q-PCR表达量分析结果见表17。
表17 LHY基因Q-PCR表达量分析
以上结果表明，LHY基因在各器官中均有表达，表达量由高到低依次为叶、花、花芽、茎、叶芽、荚果。可见，LHY基因的表达具有组织特异性，可能在紫云英花发育过程中起到重要的作用。
实施例3转基因拟南芥的培养及其表型分析
1、含本发明紫云英LHY基因(SEQ ID NO:1)的重组质粒的构建。
2、转化农杆菌感受态
将测序正确的重组质粒转化农杆菌感受态。菌落PCR鉴定表明载体质粒已成功转入到农杆菌中。
3、拟南芥转化流程(花序浸染法)
(1)种植：选择吸水性好，土质松软的至石配合营养土(1：1/2)作为拟南芥种植土壤。直径9cm的花盆，每盆播种100-150颗。播种以后在花盆上罩上薄膜，给植株的生长提供一个湿润的环境。
(2)移栽：播种10-15天，待拟南芥幼苗长至四叶时期开始移栽，每盆4-5棵。
(3)去顶：拟南芥初次开花时将花蕾剪掉，可以促进侧枝更多的花枝的增生。适合转化植株的花卉并没有成熟，也没有产生已受精的角果。
(4)配制浸染液：在5％的蔗糖溶液中重悬转化后的农杆菌使OD＝0.8，为了保持蔗糖溶液的新鲜，可以现配现用，无需灭菌。100-200ml浸染2-3个小盆植株，400-500ml浸染，2-3个花盆(9cm)植株。在浸染之前加入表面活性剂至浓度0.05％(500ul/L)。
(5)浸染：将盛花期拟南芥的花表面部分浸泡在转化后的农杆菌悬浮液中20-30s，同时轻轻旋转。
(6)暗培养：将浸染后植株套袋保持高度的湿润状态暗室培养24h。
(7)浸染后培养：隔天浇水，保证水分充足即可。
(8)种子收集：种子成熟，角果自然开裂后可以收种子。
(9)转基因种子筛选：在含有潮霉素抗生素的平板上培养浸染后所得种子。40mg的种子大约200颗于含潮霉素10-50μg/ml的0.5×MS培养基上春化2天，之后在持续光照条件下培养7-10天。根据生长状况判断是否为转基因种子。成功转入重组质粒的种子能够在抗性培养及上正常生长出4片以上真叶。非转基因种子不能正常生长，仅能长出2片子叶，根的生长也受到严重抑制，一般萌发10天后死亡。
(10)转基因植株转土栽培。转基因种子在含潮霉素的MS平板上萌发2周以后，将阳性植株转入土壤继续培养。
(11)PCR鉴定：取阳性植株叶片进行基因组DNA的提取并使用目的基因序列和载体35S启动子序列引物进行PCR验证，引物序列如下(5′-3′)：
P1235F:CGGGATCCATGGACGCTTATTCCTCTGG
P1235R:CCGCTCGAGAGTATCTCCCTCCAAGCG
检测转基因阳性植株。
植物筛选标记潮霉素基因检测：
潮霉素F：GAGCATATACGCCCGGAGTC，潮霉素R：GTCTCCGACCTGATGCAGCTCTCGG。
将LHY基因转化到农杆菌C5C81，利用花序法侵染拟南芥。将待检测的拟南芥种子在抗性培养基上种植，转化成功的植株能够正常生长，而未转化成功的植株是黄色死亡植株LHY基因未转化成功。将初筛得到的幼苗移栽到培养钵中继续培养，当生长至10～12片时，剪取叶片进行基因组DNA的提取，并使用目的片段扩增引物及载体35S启动子序列引物进行PCR检测，电泳结果出现目的条带，表明LHY基因已经整合到拟南芥基因组中。
4、转基因拟南芥表型分析
播种转基因拟南芥T2代，记录转基因株系的开花情况。从播种后开始计算抽薹、开花所需天数，以及开花数，结果见表18和图8。
表18
可见，与野生型拟南芥植株相比，转基因拟南芥的系表现出晚花表型。对转基因拟南芥及野生型拟南芥的抽薹天数、开花天数及开花数进行统计。结果显示，超表达LHY基因拟南芥株系的抽薹天数比野生型平均晚18.36天，开花天数比野生型平均晚20.72天。这表明LHY基因的超表达对拟南芥的抽薹、花期均有影响。LHY基因的表达可以延迟植物开花。
虽然，上文中已经用一般性说明及具体实施方案对本发明作了详尽的描述，但在本发明基础上，可以对之做一些修改或改进，这对本领域技术人员而言是显而易见的。因此，在不偏离本发明精神的基础上所做的这些修改或改进，均属于本发明要求保护的范围。
序列表
<110> 浙江省农业科学院
<120> 紫云英LHY基因及其应用
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atcttcttct ggcctacttc tacttgtgct tctctcctcc tcctgctact gctgctactg 60
ctgctactgc tactgctact gctctctctc tctcttccaa tttgattctt cctgctttca 120
atctgaaagg ctcacatgag tccgccggtg aattcctgcg cttcacacac atgatcgcac 180
tttcaacctt cctctttttc ttcaacatgc ctttatcccc tggttttggc gctgaggact 240
tcgttttgta gcaccgccat cacctgatca tcagcttctg cttactttca ccacaacggc 300
tttcctattt gctctgcttc atgtcacaaa aaaagaaagg aggttttcct tttacccact 360
cctagtcagg gaagatctca agcagcgcta gctgcacatc tcctctcatg gacgcttatt 420
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aacgatggac cgaggaggaa cacaatagat ttcttgaagc cctgaaacta tacgggcgag 540
catggcagcg catagaagag cacataggaa caaagactgc tgtgcaaatt cggagccatg 600
cacagaaatt cttttcaaag ttggagaagg aagccctcgt aaaaggtgtt cctgtaggtc 660
aaactcttga catagacatt ccccctccac ggcccaaaag aaaaccaagc aatccttatc 720
ctcggaagac ctatgttggt accccaacat tacctagtgg agaaaagtat ggaaagcctc 780
ttcctgtggt tgcatcttca tatggtaaac aagcaatgga cttcgacaaa gaaccacttc 840
cagagaagaa caatgatgat gaaaggccaa caactgcaaa agaaaataag gatgataatt 900
gctcaaaatt atttacgatc ctccaggagg caccatgtgc atctgtatct tcagcaaaca 960
agagttcaat cacaatgtca gtgccgcaga gaaattcctg caccttaagg gagtttatac 1020
catcagtgaa agaggtaata actcaaggtg aaacaaatca atcttttgtc actactgaga 1080
ttgaaaatca gaagttggag atagatgatg gcaaacacac acaagagagt aatggtactt 1140
gtacagcctc taagttggag aactcttgtt ctcccaaatc agttcaaact gagaaaacag 1200
atgggcttac ttgtgcgtta acaatagatg agatgcaagt taatcagaat tacccaagac 1260
acatcactgt gcacgttgtt gatgggaacc ttggaaatac tactcaaaat gcatcacaag 1320
ataagttgat tcaagactcc atatttcagc caataggggt taatggccaa cctaattttt 1380
ttgccaactc agctgcatca aacacaagtg gaagtcaaaa taacacagca cgatcctcta 1440
ttcatcaatc attttcttct agtcctccct ttgcgcaaca taatcacgag gattaccaat 1500
cttttcttca catgtcctct acattttcaa gtcttattgt gtctaacttg ctgcaaaacc 1560
ctgcagccca tgctgcagca agttttgctg caacattttg gccctatgca aatgtagaaa 1620
cttcagcgga ttctcctgct tgctcccaag gaggttttac atctagacaa attgctcccc 1680
ctccaagtgt ggcagctatt gcagctgcta cggtagctgc tgcaactgca tggtggacag 1740
ctcatggact gcttccgctg tgtgctccac tccatgcacc ttttgcatgt cctcctgcat 1800
caacaactgc agttccatca atggatgttg gtgaagcaca gccaaagaca gagcaaggaa 1860
atattacact acaaaatcct cctttacaag atcagatact agatccaaaa gacgcggaag 1920
ctctgcgagc tcaacgtccg gtttccaagt caccaactgt ttcttcatct gaatctgagg 1980
agaggggaga tgccaagtta aatacttcac caaaggccac tactaataat gagataaacc 2040
aagcaatttc tgagaacccc gattccaaca tattgaaggg cagaaaactt gttgaccgtt 2100
cctcatgtgg ttctaacaca ccctccagca gcgaagagac tgatgcacta gcaaagaatg 2160
agaaagaaaa ggaagaattc agaacacctg atgcaaacca gttagctact gagcctagta 2220
atcgtagaaa tagaagtatt agcaacctta ttgattcttg gaaggaagtc tctgaagagg 2280
gacgaattgc ctttcatgct ctattctcca gggaggtttt gccccaaagc ttttcacctc 2340
ctcatgattt gataattaag gaaaatcaga tggacagcat gaaggataac gagcaaaaaa 2400
cagactacca agattacctt gagggcaaga aatgtagttc taattttaat gggatgcaga 2460
aagacgtaca atttgtagaa aataacagtg aggaggaagg actgttaacc atgagtctgg 2520
gacaaggaaa gctaaaaact cgtcgaacag gatttaaacc ttacaaaaga tgtctggtag 2580
aggcgaagga aaacaggatg ggaacggcct gcaaccaagt tgaagagaca gttccaaaga 2640
gaatacgctt ggagggagat acttgacggt tgatatgata tattcatgca aatgcaatca 2700
cttaatgtat tgagtttaag tctctccttg ctcggtgtct gtattgttta atttctaatt 2760
ctgtagctca cgaacatgac gtgtaagatg tgtgtaccat agtaaattat taccaactct 2820
cttttacttt gttaattgga ctcatgttac tataaaaaaa aaaaa 2865
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Gln Asn Asn Thr Ala Arg Ser Ser Ile His Gln Ser Phe Ser Ser Ser
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Pro Ala Ala His Ala Ala Ala Ser Phe Ala Ala Thr Phe Trp Pro Tyr
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Ser Thr Thr Ala Val Pro Ser Met Asp Val Gly Glu Ala Gln Pro Lys
465 470 475 480
Thr Glu Gln Gly Asn Ile Thr Leu Gln Asn Pro Pro Leu Gln Asp Gln
485 490 495
Ile Leu Asp Pro Lys Asp Ala Glu Ala Leu Arg Ala Gln Arg Pro Val
500 505 510
Ser Lys Ser Pro Thr Val Ser Ser Ser Glu Ser Glu Glu Arg Gly Asp
515 520 525
Ala Lys Leu Asn Thr Ser Pro Lys Ala Thr Thr Asn Asn Glu Ile Asn
530 535 540
Gln Ala Ile Ser Glu Asn Pro Asp Ser Asn Ile Leu Lys Gly Arg Lys
545 550 555 560
Leu Val Asp Arg Ser Ser Cys Gly Ser Asn Thr Pro Ser Ser Ser Glu
565 570 575
Glu Thr Asp Ala Leu Ala Lys Asn Glu Lys Glu Lys Glu Glu Phe Arg
580 585 590
Thr Pro Asp Ala Asn Gln Leu Ala Thr Glu Pro Ser Asn Arg Arg Asn
595 600 605
Arg Ser Ile Ser Asn Leu Ile Asp Ser Trp Lys Glu Val Ser Glu Glu
610 615 620
Gly Arg Ile Ala Phe His Ala Leu Phe Ser Arg Glu Val Leu Pro Gln
625 630 635 640
Ser Phe Ser Pro Pro His Asp Leu Ile Ile Lys Glu Asn Gln Met Asp
645 650 655
Ser Met Lys Asp Asn Glu Gln Lys Thr Asp Tyr Gln Asp Tyr Leu Glu
660 665 670
Gly Lys Lys Cys Ser Ser Asn Phe Asn Gly Met Gln Lys Asp Val Gln
675 680 685
Phe Val Glu Asn Asn Ser Glu Glu Glu Gly Leu Leu Thr Met Ser Leu
690 695 700
Gly Gln Gly Lys Leu Lys Thr Arg Arg Thr Gly Phe Lys Pro Tyr Lys
705 710 715 720
Arg Cys Leu Val Glu Ala Lys Glu Asn Arg Met Gly Thr Ala Cys Asn
725 730 735
Gln Val Glu Glu Thr Val Pro Lys Arg Ile Arg Leu Glu Gly Asp Thr
740 745 750
Claims (10)
1.紫云英LHY基因，其特征在于，LHY基因为编码如下蛋白质(a)或(b)的基因：
(a)由SEQ ID NO:2所示的氨基酸序列组成的蛋白质；
(b)SEQ ID NO:2所示序列经取代、缺失或添加一个或几个氨基酸且具有同等功能的由(a)衍生的蛋白质。
2.含有权利要求1所述LHY基因的生物材料，所述生物材料为重组DNA、表达盒、转座子、质粒载体、噬菌体载体、病毒载体或工程菌或非可再生的植物部分。
3.权利要求1所述LHY基因或含有该基因的生物材料在调控植物开花时间中的应用。
4.根据权利要求3所述的应用，其特征在于，所述调控是指延迟开花时间。
5.根据权利要求4所述的应用，其特征在于，所述应用包括：
1)使植物包含LHY基因；或者，
2)使植物过表达LHY基因。
6.根据权利要求5所述的应用，其特征在于，所述应用包括转基因、杂交、回交、自交或无性繁殖。
7.根据权利要求5所述的应用，其特征在于，过表达LHY基因的方法选自以下1)～4)，或任选的组合：
1)通过向植物中导入具有LHY基因的质粒；
2)通过增加植物染色体上LHY基因的拷贝数；
3)通过将强启动子与LHY基因可操作地连接；
4)通过导入增强子。
8.根据权利要求3-7任一项所述的应用，其特征在于，所述植物为拟南芥、紫云英。
9.权利要求1所述LHY基因或含有该基因的生物材料在植物育种中的应用。
10.根据权利要求9所述的应用，其特征在于，育种目的为延迟开花时间。
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