CN114214340A - 水稻粒型粒重相关基因、蛋白、分子标记及应用 - Google Patents
水稻粒型粒重相关基因、蛋白、分子标记及应用 Download PDFInfo
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- CN114214340A CN114214340A CN202111606443.1A CN202111606443A CN114214340A CN 114214340 A CN114214340 A CN 114214340A CN 202111606443 A CN202111606443 A CN 202111606443A CN 114214340 A CN114214340 A CN 114214340A
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Abstract
本发明公开了基因在调控植物粒型、粒重中的应用，所述基因具有如(a)、(b)、(c)所示的序列：(a)Seq ID No：1所示的基因组核苷酸序列；(b)Seq ID No：2所示的cDNA核苷酸序列；(c)在(a)、(b)所示的核苷酸序列的中添加和/或取代和/或缺失一个或几个核苷酸而生成的可编码具有调控粒型粒重功能的蛋白质的突变基因、等位基因或衍生物。所述基因正调控植物(水稻)粒型、粒重。
Description
技术领域
本发明涉及植物基因工程领域，具体地，涉及一种水稻粒型粒重相关基因、蛋白、分子标记及应用。
背景技术
水稻不仅是单子叶模式植物，而且是我国重要的粮食作物。水稻籽粒的大小是影响稻米产量和品质的重要因素，在长期的水稻育种过程中一直备受关注。水稻粒型和粒重是影响产量的直接因素，千粒重是稻谷产量三要素之一，而粒重主要由粒型决定。同时，粒型对稻米品质也有重要影响，尤其是稻米的外观品质和碾磨加工品质等。控制粒型的4个关键因素分别为粒长、粒宽、粒厚和长宽比。
但是，籽粒大小是由多个遗传位点控制的复杂数量性状，至今，科学家们利用图位克隆方法已成功克隆到一系列调控水稻粒型变异的数量性状遗传位点(QTL)。这些QTL主要通过调控水稻颖壳的大小来影响籽粒的形成，其中与粒长和粒重相关的基因有如下所示：
一、GS3
Fan C,Xing Y,Mao H,Lu T,Han B,Xu C,Li X,Zhang Q.GS3,a major QTL forgrain length and weight and minor QTL for grain width and thickness in rice,encodes a putative transmembrane protein.Theor Appl Genet.2006,112(6):1164-1171(Fan Chuchuan,Xing Yongzhong,Mao Hailiang,Lu Tingting,Han Bin,Xu Caiguo,Li Xianghua,Zhang Qifa.GS3基因，水稻粒长粒重主效QTL、粒宽粒厚微效QTL，编码一推测的跨膜蛋白.理论和应用遗传学.2006,112(6):1164-1171)。
二、qGL3/qGL3.1
Hu Z,He H,Zhang S,Sun F,Xin X,Wang W,Qian X,Yang J,Luo X.A Kelchmotif-containing serine/threonine protein phosphatase determines the largegrain QTL trait in rice.J Integr Plant Biol.2012,54(12):979-990(Hu Zejun,HeHaohua,Zhang Shiyong,Sun Fan,Xin Xiaoyun,Wang Wenxiang,Qian Xi,Yang Jingshui,Luo Xiaojin.一包含Kelch结构域的丝氨酸/苏氨酸蛋白磷酸酶决定水稻大粒QTL性状.植物学报.2012,54(12):979-990)；
Qi P,Lin YS,Song XJ,Shen JB,Huang W,Shan JX,Zhu MZ,Jiang L,Gao JP,LinHX.The novel quantitative trait locus GL3.1 controls rice grain size andyield by regulating Cyclin-T1；3.Cell Res.2012,22(12):1666-1680(Qi Peng,LinYou-Shun,Song Xian-Jun,Shen Jin-Bo,Huang Wei,Shan Jun-Xiang,Zhu Mei-Zhen,Jiang Liwen,Gao Ji-Ping,Lin Hong-Xuan.新的数量性状位点GL3.1通过调控细胞周期蛋白-T1；3控制水稻籽粒大小和产量.细胞研究.2012,22(12):1666-1680)；
Zhang X,Wang J,Huang J,Lan H,Wang C,Yin C,Wu Y,Tang H,Qian Q,Li J,Zhang H.Rare allele of OsPPKL1 associated with grain length causes extra-large grain and a significant yield increase in rice.Proc Natl Acad SciUSA.2012,109(52):21534-21539(Zhang Xiaojun,Wang Jianfei,Huang Ji,Lan Hongxia,Wang Cailin,Yin Congfei,Wu Yunyu,Tang Haijuan,Qian Qian,Li Jiayang,ZhangHongsheng.与粒长相关的OsPPKL1基因的稀有等位基因产生了特大谷粒的增产水稻.美国科学院院刊.2012,109(52):21534-21539)。
三、GL7
Wang Y,Xiong G,Hu J,Jiang L,Yu H,Xu J,Fang Y,Zeng L,Xu E,Xu J,Ye W,Meng X,Liu R,Chen H,Jing Y,Wang Y,Zhu X,Li J,Qian Q.Copy number variation atthe GL7 locus contributes to grain size diversity in rice.Nat Genet.2015,47(8):944-948(Wang Yuexing,Xiong Guosheng,Hu Jiang,Jiang Liang,Yu Hong,Xu Jie,Fang Yunxia,Zeng Longjun,Xu Erbo,Xu Jing,Ye Weijun,Meng Xiangbing,LiuRuifang,Chen Hongqi,Jing Yanhui,Wang Yonghong,Zhu Xudong,Li Jiayang,QianQian.GL7位点的拷贝数变异引起了水稻籽粒大小的多样性.自然·遗传.2015,47(8):944-948)。
四、GLW7
Si L,Chen J,Huang X,Gong H,Luo J,Hou Q,Zhou T,Lu T,Zhu J,Shangguan Y,Chen E,Gong C,Zhao Q,Jing Y,Zhao Y,Li Y,Cui L,Fan D,Lu Y,Weng Q,Wang Y,ZhanQ,Liu K,Wei X,An K,An G,Han B.OsSPL13 controls grain size in cultivatedrice.Nat Genet.2016,48(4):447-456(Si Lizhen,Chen Jiaying,Huang Xuehui,GongHao,Luo Jianghong,Hou Qingqing,Zhou Taoying,Lu Tingting,Zhu Jingjie,ShangguanYingying,Chen Erwang,Gong Chengxiang,Zhao Qiang,Jing Yufeng,Zhao Yan,Li Yan,Cui Lingling,Fan Danlin,Lu Yiqi,Weng Qijun,Wang Yongchun,Zhan Qilin,LiuKunyan,Wei Xinghua,An Kyungsook,An Gynheung,Han Bin.OsSPL13基因控制栽培稻的籽粒大小.自然·遗传.2016,48(4):447-456)。
五、qTGW3
Hu Z,Lu SJ,Wang MJ,He H,Sun L,Wang H,Liu XH,Jiang L,Sun JL,Xin X,KongW,Chu C,Xue HW,Yang J,Luo X,Liu JX.A novel QTL qTGW3 encodes the GSK3/SHAGGY-like kinase OsGSK5/OsSK41 that interacts with OsARF4 to negatively regulategrain size and weight in rice.Mol Plant.2018,11(5):736-749(Hu Zejun,Lu Sun-Jie,Wang Mei-Jing,He Haohua,Sun Le,Wang Hongru,Liu Xue-Huan,Jiang Ling,SunJing-Liang,Xin Xiaoyun,Kong Wei,Chu Chengcai,Xue Hong-Wei,Yang Jinshui,LuoXiaojin,Liu Jian-Xiang.一新的QTL qTGW3编码类GSK3/SHAGGY激酶OsGSK5/OsSK41与OsARF4互作负调控水稻籽粒大小和粒重.分子植物.2018,11(5):736-749)。
因此，克隆水稻粒型基因，开发相应的分子标记，通过分子育种改良水稻粒型，可以同时提高水稻产量和改良水稻品质。
水稻GL4基因编码蛋白含有3个Kelch重复结构域。Kelch结构域的一级结构存在8个关键保守位点，包括4个疏水氨基酸、2个紧接连续的甘氨酸和隔开一段序列后的2个有固定间隔的芳香族氨基酸(Goebel SJ,Johnson GP,Perkus ME,Davis SW,Winslow JP,Paoletti E.The complete DNA sequence of vaccinia virus.Virology.1990,179(1):247-266)。该GL4蛋白与二穗短柄草的含Kelch结构域蛋白4同源性高(88％相似度)。但其功能至今尚不清楚。
发明内容
本发明要解决的技术问题在于提供一种调控水稻粒型和粒重的水稻粒型粒重基因GL4及其编码的蛋白质，并基于此开发其相关应用。
为解决上述技术问题，本发明提供基因在调控植物粒型、粒重中的应用，所述基因具有如(a)、(b)、(c)所示的序列：
(a)Seq ID No：1所示的基因组核苷酸序列；
(b)Seq ID No：2所示的cDNA核苷酸序列；
(c)在(a)、(b)所示的核苷酸序列的中添加和/或取代和/或缺失一个或几个核苷酸而生成的可编码具有调控粒型粒重功能的蛋白质的突变基因、等位基因或衍生物；
所述粒型为粒长。
作为本发明应用的改进：正调控植物(水稻)粒型、粒重，即，使得粒长增加、千粒重增加。
本发明还同时提供了上述基因编码的蛋白在调控植物粒型粒重中的应用，所述蛋白如(A)或(B)所示的序列：
(A)Seq ID No：3所示的氨基酸序列；
(B)在(A)所限定的氨基酸序列中添加和/或取代和/或缺失一个或几个氨基酸且具有相同功能的由(A)衍生的蛋白质。
其中的Seq ID No：3所示的蛋白质具有662个氨基酸，属于富含Kelch结构域(即Seq ID No：3中的第128～172位、第180～233位、第236～288位)的新蛋白。
其中Seq ID No：1所示的93-11基因组核苷酸序列共有6164个核苷酸，Seq ID No：2所示的93-11cDNA序列共有1989个核苷酸(包括终止子TAG)。
为了验证候选基因GL4的功能，构建了基因互补载体；所述基因互补载体为pCAMBIA1300载体；所述用于构建互补载体的基因引物为：
com-F：aataagcttACTAATTACATGGAATGCGTGTAAATTG；
com-R：agaagcttCATTTCGTACGGAGGAGAACAACTG；
进一步地，由于上述基因属于正调控基因，因此可通过过表达来提高水稻粒长粒重；所述基因过表达载体为pCAMBIA1300S载体；所述用于构建过表达载体的基因引物为：
over-F：aagggtaccATGGGGAAGAAGCAGAAGAAGCC；
over-R：tgctctagaTCGAACCAAAGCTATCTCATGCC；
此处的应用，主要是将基因过表达通过转基因的方法转入水稻中以提高水稻的粒长粒重。
另一方面，本发明提供了一种与水稻粒型粒重基因紧密连锁的分子标记，所述分子标记为P4-1、P4-2、P4-3、P4-4、P4-5、P4-6、P4-7、4-8或P4-9；所述各分子标记对应的引物序列分别为：
P4-1：
F,5’-TGGGTCTTCAAAAAATGTTCAGTGG-3’
R,5’-ACCCCGCCTAAACTCCATGAATC-3’；
P4-2：
F,5’-TATAGATTCATCGTACTAAGGC-3’
R,5’-TGAGATTTATTGTTTTGTGTG-3’；
P4-3：
F,5’-AGAAGTAGTGCAGAGTACAGTC-3’
R,5’-AGTACTCCTATCCTTTAATAATATG-3’；
P4-4：
F,5’-ATACGTAGCGTTTGGTTATAGC-3’
R,5’-TTCGGTTTTGAACTCAACTTC-3’；
P4-5：
F,5’-TGCTTGAAGAGGAGAATGGTGG-3’
R,5’-AGCTCCTGAGTTCCTTGCGTC-3’；
P4-6：
F,5’-TTACATTATCGAATTATGCACGATAC-3’
R,5’-TGATACCCGAACTTCCTGACTG-3’；
P4-7：
F,5’-TGTGGCAGAATTGTGGGACAC-3’
R,5’-ACTTTATATCTGATTTAGGCACGTTTAC-3’；
P4-8：
F,5’-TAGATTGGTTTTTATGAAACG-3’
R,5’-TGCTGTCACAGTTTATCACAC-3’；
P4-9：
F,5’-TCCTACGATTTCTCAATCCTG-3’
R,5’-TGAATTCCTTCAATTTTAGAGC-3’。
再一方面，本发明提供了上述与水稻粒型粒重基因紧密连锁的分子标记在分子标记辅助选择育种中的应用，所述应用为辅助选择与水稻粒型粒重相关性状。
上述分子标记是在进行水稻粒型粒重基因定位克隆过程中得到的与其紧密连锁的分子标记，因此，可通过上述获得的分子标记进行分子标记辅助选择育种，如筛选、鉴定与水稻粒型粒重相关性状。
实现本发明的具体技术步骤如下：
一、GL4基因的精细定位和候选基因确定：
利用粒长和粒重上存在显著差异的亲本培矮64s与93-11(图1，图2)构建了大规模的重组自交系(RIL)群体。结合RIL核心群体的高密度遗传图谱，我们检测到水稻第4号染色体上新的控制稻谷粒长和粒重的QTL——qGL4(图3a)。利用培矮64s为轮回亲本的大规模BC4F2群体将该QTL进一步精细定位于两个插入缺失(INDEL)标记P4-7和P4-8之间约9.3kb的物理距离内(图3b)，其中只有1个开放阅读框(ORF)，编码产物与含Kelch结构域的未知功能蛋白同源性高，暂命名为GL4。DNA测序发现，亲本培矮64s和93-11的GL4编码区1989bp内存在2个引起氨基酸改变的单核苷酸多态(SNP)(图3c)。
二、GL4基因的鉴定和功能分析：
通过转基因技术，结果表明本发明获得了粒长和千粒重显著增加的转基因互补培矮64s水稻植株和过表达培矮64s水稻植株(图5–图8)，并且过表达培矮64s水稻植株的基因转录水平表达量显著升高(图9)，证明了本发明正确克隆了GL4基因。氨基酸序列分析表明GL4编码包含Kelch结构域蛋白，对93-11背景的GL4基因为培矮64s类型的近等基因系NIL-GL4培矮64s和93-11的籽粒观察发现，粒长和千粒重也显著减小(图10；图11)，证明该基因编码蛋白调控水稻粒型。
本发明采用图位克隆技术利用水稻BC4F2群体首次在水稻中克隆到粒型粒重基因GL4并通过转基因互补实验和过表达实验以及近等基因系鉴定了基因的功能。GL4基因的克隆和应用，可有效调控水稻粒型粒重和产量，GL4基因也可应用于其它单子叶植物中用于调控粒型粒重和产量。同时本发明有助于水稻粒型粒重调控机理的阐明，并为水稻高产优质育种奠定扎实的理论基础。
综上，公开了水稻粒型粒重基因GL4(Seq ID No：1、2)、其编码的蛋白质(Seq IDNo：3)及上述基因或蛋白质在调控植物粒型和粒重中的应用。本发明还公开了与上述基因紧密连锁的分子标记及其应用。GL4基因的克隆和应用，可通过调控水稻粒型粒重，有效提高水稻的产量。
附图说明
下面结合附图对本发明的具体实施方式作进一步详细说明。
图1是水稻品种培矮64s和93-11的粒型表型；其中a为培矮64s的籽粒，b为93-11的籽粒，c-1cm标尺；
图2是水稻品种培矮64s和93-11的粒长和千粒重比较；a平均值±标准差(n＝100)，b平均值±标准差(n＝3)；
图3GL4基因的定位图；其中竖线上方标注分子标记，竖线下方数字代表交换个体数，ATG和TAG分别表示GL4基因的起始子和终止子；
图4pCAMBIA1300-GL4互补载体图谱(a)和pCAMBIA1300S-GL4过表达载体图谱(b)；
图5是培矮64s和GL4互补T0代转基因水稻的粒型表型；其中a为培矮64s的籽粒，b、c、d为转GL4互补株系的籽粒，e-1cm标尺；
图6是培矮64s和GL4过表达T0代转基因水稻的粒型表型；其中a为培矮64s的籽粒，b、c、d为转GL4过表达株系的籽粒，c-1cm标尺；
图7是培矮64s和转GL4互补载体株系的粒长和千粒重比较；a平均值±标准差(n＝100)，b平均值±标准差(n＝3)；
图8是培矮64s与转GL4过表达载体株系的粒长和千粒重比较；a平均值±标准差(n＝100)，b平均值±标准差(n＝3)；
图9是培矮64s与转GL4过表达载体株系的GL4转录水平的表达比较；平均值±标准差(n＝3)；
图10是93-11和近等基因系NIL-GL4培矮64s的粒型表型；其中a为93-11的籽粒，b为NIL-GL4培矮64s的籽粒，c-1cm标尺；
图11是93-11和近等基因系NIL-GL4培矮64s的粒长和千粒重比较；a平均值±标准差(n＝100)，b平均值±标准差(n＝3)。
具体实施方式
下面结合具体实施例对本发明进行进一步描述，但本发明的保护范围并不仅限于此：
实施例1：
1、水稻材料
籼稻品种为(Oryza sativa L.indica)“93-11”和“培矮64s”，93-11背景的GL4基因替换为培矮64s类型的近等基因系NIL-GL4培矮64s。
近等基因系NIL-GL4培矮64s(93-11遗传背景下INDEL标记P4-6～P4-8之间被培矮64s片段代换)由染色体片段代换系CSSL-qGL4与93-11连续回交4代并自交结合INDEL标记P4-1～P4-9辅助筛选得到。染色体片段代换系CSSL-qGL4是由培矮64s与93-11连续回交5代并自交的BC5F2筛选得到，步骤详见Zhang B,Shang L,Ruan B,Zhang A,Yang S,Jiang H,Liu C,Hong K,Lin H,Gao Z,Hu J,Zeng D,Guo L,Qian Q.Development of three sets ofhigh-throughput genotyped rice chromosome segment substitution lines and QTLmapping for eleven traits.Rice.2019,12(1):33。
2、INDEL标记精细定位GL4基因
采用水稻微量DNA的快速提取方法从水稻叶片中提取用于基因定位的基因组DNA。取0.2g水稻叶片，液氮冷冻研磨成粉状，转移至1.5ml离心管里提取DNA，获得的DNA沉淀溶解于150μl超纯水中。每一PCR反应用2μl DNA样品。
GL4基因的初定位：将培矮64s与93-11杂交，F1代开始自交单粒传13代产生RIL群体。结合RIL核心群体的高密度遗传图谱，检测到水稻第4号染色体上新的控制稻谷粒型和粒重的QTL——qGL4(图3a)。
GL4基因的精细定位：利用培矮64s为轮回亲本的大规模BC4F2群体，同时在目标区间根据培矮64s和93-11的参考序列设计引物，筛选出具有多态的引物(表1)，将该QTL进一步精细定位于两个INDEL标记P4-7和P4-8之间约9.3kb的物理距离内(图3b)。
3、基因预测和比较分析：
通过RGAP网站(http://rice.plantbiology.msu.edu/)分析，该区间内存在1个ORF，基因组测序发现，亲本培矮64s的如Seq ID No：4的基因组核苷酸序列与93-11的如SeqID No：1的基因组核苷酸序列间共存在11个SNP和1个INDEL，包括如Seq ID No：2的编码区1989bp内的2个引起氨基酸改变的SNP：SNP1为93-11的A175变为培矮64s的G175(由异亮氨酸变为缬氨酸)，SNP2为93-11的C841变为培矮64s的T841(由精氨酸变为半胱氨酸)(图3c)。因此，该ORF可能是该性状的候选基因。
表1定位发展的分子标记
*F：正向引物；R：反向引物。
本发明的序列表中：
Seq ID No：1为93-11的GL4基因的基因组核苷酸序列(6164bp)；
Seq ID No：2为93-11的GL4基因的cDNA核苷酸序列(1989bp)；
Seq ID No：3为93-11的GL4基因编码的蛋白质的氨基酸序列(662aa)；
Seq ID No：4为培矮64s的GL4基因的基因组核苷酸序列(6156bp)；
Seq ID No：5为93-11的GL4基因起始子ATG前的包含启动子的序列(2237bp)；
Seq ID No：6为93-11的GL4基因终止子TAG后的序列(1318bp)。
实施例2：
植物转化和功能分析：
为了验证该ORF为候选基因，用PCR扩增93-11的包含Seq ID No：5所示的起始子ATG前2237个核苷酸、Seq ID No：1所示的93-11基因组核苷酸序列共6164个核苷酸、Seq IDNo：6所示的终止子TAG后1318个核苷酸共9719个核苷酸，将该序列连接到常规双元表达载体pCAMBIA1300中(图4a)，所得命名为pCAMBIA1300-GL4；
用PCR扩增93-11的如Seq ID No：2所示的cDNA核苷酸序列(从起始子ATG至终止子TAG共1989bp)，然后将该序列连接到常规双元表达载体pCAMBIA1300S中(图4b)，所得命名为pCAMBIA1300S-GL4。
通过农杆菌介导的植物转化方法将此载体(即，上述pCAMBIA1300-GL4、pCAMBIA1300S-GL4)分别转入培矮64s，各得到转基因植株3株，pCAMBIA1300-GL4载体对应所得称为基因互补系植株，pCAMBIA1300S-GL4载体对应所得称为基因过表达系植株。可委托武汉伯远生物科技有限公司完成。
按照常规水稻栽培方式在转基因圃内进行种植直至收获。所有转基因互补T0代植株和过表达T0代植株的表型均为粒长和千粒重显著增加(图5，图6，图7，图8)，并且过表达T0代植株的基因转录水平的表达量显著升高(图9)。由此说明，该ORF就是控制粒型和粒重性状的候选基因GL4。
最后，还需要注意的是，以上列举的仅是本发明的若干个具体实施例。显然，本发明不限于以上实施例，还可以有许多变形。本领域的普通技术人员能从本发明公开的内容直接导出或联想到的所有变形，均应认为是本发明的保护范围。
序列表
<110> 中国水稻研究所
<120> 水稻粒型粒重相关基因、蛋白、分子标记及应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 6164
<212> DNA
<213> 水稻(Oryza sativa)
<400> 1
atggggaaga agcagaagaa gcccaggaag gggaaggaga agacggagcg gaagacggcc 60
aagggcgagg agaagcgcgc ccgccgcgag gcccggaagg tcggcgagga ggacgacatc 120
gacgccatcc tcgtacgtgt gctccctccc tcccgccccc ctcctctgcg tcagctcttc 180
acgctcgctc agtgcgctct agctcgatcg gcgtctccat gtgcggtttt gtttgctcac 240
caccaccttc tgcgtctcgt gatgcactcc ggtggctgaa aaattggaag cgattttcgc 300
actggctcac ctttttccct tcacatttcg ttgcgtagca gctatggatt ttagatgagt 360
ttggtgttgc tatgtgtgct gtttgaactt tttttttttg ctgatttatc tatttgctct 420
ggttattttt cattgcagag gagcatacaa aaggaggagg ctaagaagaa ggaggtacat 480
atagatgaga atgtccctgc accatctccc cggtccaatt gctcggtaag acattttaga 540
gcaagtgcca gctaaagaag ttaatctttc ggtattcttg tgattatgta cttgaagtgg 600
actaggtatc atttttatgc ttgtggtggt catgtttgca tgcttagaat ttatatactc 660
cagtaggatt aataaatttc ttcaggcaag atggattttt ttatgaagta atttgatcaa 720
atatgatgat cttttggtga tactgaaccg attggttttt cagttccgga taggtgatgt 780
caaattacaa ttcaggggca aaatgtgtta aaagatagat gttcgttttg tttttttttt 840
tgttaactgt tggaaaaagt ttttgatgtt gtacagagat cctccctcta aatgttaact 900
ataacaaata aacctgcata tgatcttcca gctgaccaag ttaattcctt cttaaatgca 960
gcttacaata aatcccctga aagatacaga attggttctg tatggaggag agttctacaa 1020
tggcagcaag gtgggagaca tatccttcac gatttcactg ctggttgaat gagatatgtt 1080
catctagtgt tctttttgcc tctattctgc ttctagaggt ttttgagtac tgaaatagta 1140
tttcttttcc tcgccacaat ctatttctgc agacctttgt ttatggtgat ctttatcgct 1200
acgatgtaga gaaaaatgag tggaagttgg tatctagtcc taacagtcct cctccacgaa 1260
gtgctcacca aacagttgcc tggaagaata atatatacat gtttggtaac ataacttaac 1320
tttgggaagg cattctatga gttaaaatgc tttttcagta tgtatgattt agttttttat 1380
tctgtgttga tgtttcaggt ggggaattca cttcgccaaa ccaagaacgt tttcatcatt 1440
acaaggtaga agactaattt tgtcagtcta tttcttgtga ctgttttgag tatcttctct 1500
tggcacaagg catttgcaca attcgtatat aagcagtagc ttcataagca atacatatct 1560
ggcatgattt tttttcaatt ttaaaccaag agataggtac tgattcccat gttcttacat 1620
aattataagt tgaaatatga ctaatggaga ctaatatgtg cagttcttct gctatttatt 1680
gatttaagat taagatgtag aggatgcggg gttcacctgc ccttttattt ggatttgcac 1740
atataacatc actttttatt cctgattgcc tttttttttc ctttctgttg tgatattaat 1800
ggggttattt cgagattagc tatcttagat ggaaaactca ttattatggt tctattcaaa 1860
tttctggtct gattttgtac tgtaggactt ttggtcattg gatctaaaaa caaatcaatg 1920
ggagcaaatt cttgcgaagg gttgtccaag tgcacgttca gggcacagga tggttagtgg 1980
ttttacatta aaatcagtca tcaactattc tgctcccctc ttttcacctt aatatttctg 2040
tattatgagc agtagtaaat tgtgttcttt ttcctacagg tcctctataa gcacaagatc 2100
gtgctatttg gtggttttta tgacactctt agggaagtga ggttagtaca gttacatttt 2160
atatgactct accctggtaa tcttgttatt agagtaacat ttatatttgg accatgtcta 2220
gaagtagtgg agtcattatg tccaaactac aaattatgca ctacctgaat aacagtgtag 2280
cactcttaca gctgatcatc cgcaaagaat gaaatggtgt ggcgtagagt acgttctgaa 2340
taaatagtgt gacatgacat gatctgatct gcattttttt tataatatcc ttctgcagat 2400
actacaatga cttacatgtt tttgatttag ataatttcaa ggtgagtaca ccatgttaat 2460
attttgttta atactgttag tagtaacaca tgaagtcatt tattttaata ctcttactgg 2520
gaatatttgt atttcagtgg gaggagatca agcctcgccc tgggtgcttg tggccaagtc 2580
caagaagtgg ctttcagcta atggtatacc aagatcaggt aggtcttttt ggatttaaag 2640
ctaggacatt gatacttcat aaaaagagtt taaattaact ataaaccaac cttgtcttcg 2700
actacttttg tttacaagag tattaatggc ccttatttct gtagatatat ctgtatggcg 2760
gatattttaa agaagtagtt tcttctgaca aatctgcatc agaaaaagga acagttcatg 2820
cagatatgtg gactcttgat cctcgtactt gggagtggaa taaggtgatc tcttgcaatt 2880
ttttagaaca ttgtatcaac ttccatcatg atagtgtatc gagttttact ttaagccata 2940
tatccactga gtgatttgca tattattacc ttcacttgat ttcttaatag gttaagaaaa 3000
ctgggatgcc acctggcccc agagctgggt tttctatgtg cgttcacaag aaaagggctg 3060
ttcttttcgg tggtgtggta gatatggaaa ttgaaggtta ttttcagctc aattttgctc 3120
tgtgcatagc tacttaggtt attttactaa gtatttgaaa taccacgcgt gtcaagtttg 3180
ttccttttct gtagaagttc tcaaggccta actgtagaac ccaattttgt gattgcaggg 3240
gatgtcatta tgagcatgtt tatgaatgag ctctatggtt tccagctgga caaccatcgc 3300
tggtatactt caatactcca tttgagtatc ttgtgttttt aagtaacaca ggctagtttt 3360
atccctgctt attttgttcc ttgctatttt tgtgtattgt tgcaactttc tttattatta 3420
tttaacagtg cagctaacta atgtcacttc ctattccgat atgcaaactg cttctaacta 3480
ggtttaaaat tattaaaggt ctaaatcttt ctcggctgat tgatggttat gttcatctat 3540
cgtcttagga tgaactttgt ttgtaatctt gtggttatct agataacata actactttga 3600
gaattgttca gtgatattat tgtttactct tgggatccct tcgtagcata ttattgttta 3660
cttcagcgga attgttcagc gatgttcatt ttggttgaaa ctactggtcc acggctcaca 3720
tattctccaa tttcaattgt ccacactggt gcatagatga gaaatgttat gctttcttat 3780
ttagcttcat ttttgtgtgt ctatgttcag attattttcc ttgctgtttc aggtatcctt 3840
tagagctcag gaaagacaag cctgctaaaa ataaggtgat tcctcgaatt caatcatatg 3900
accagccatc tcatgtttac agtttgtatt ttttaggtac ttgatacaca gtctactctg 3960
caggaatcta tttcattagt tcagacataa agactaagca tgtagtatag ggtatgcctg 4020
tttttctttt taataagcat atacgatatg tacatattct catatactat tatactgtta 4080
ttttcttaat aagcatatgc gatctgtcga catattcaca ttgactatta tatagtaata 4140
tttggatcat tgcttaaaag cagtttgtgc ttcttttttt ttttttgggt gccatgtgta 4200
gacaaaggac atcaaaagaa aagaaccatc gaacaatgtg gaagataatc ttggtaatga 4260
ggaggatgag atcatggagg actcagaaac tactggaggg caatccgaag tccatggggt 4320
ttcgaatcac ttgaccaaga gtctaacctt aaataaagct ggctcaggca atagctctga 4380
tattctctct gattcgacaa cacaagaagt actcccagag gtattgcagc tgttctttta 4440
gatgttgaca tttacattct aatgatcttt tgtttctcat tagcatttgc tgcttacagg 4500
cagtgaaacc cggtggtcgg atcaatgcat gcttggctgt agggaaagat acactctatt 4560
tatatggagg aatgatggaa ttgaaagata gagaaattac tcttgatgat atgtattcac 4620
ttaaccttag caaactagat gagtggaagt gtatcatacc ggtcagttgc agattggccc 4680
cttctttttg ccattttgtt gtttaactga tagtgttgtt tatttcaatc agagataaca 4740
gaaaattatc tgttatcatt ttttactatt cacattggtt tctgaacttg ccttactcac 4800
ctttcttatg caggcatctg aatctgaatg gctagaaatt tctgaagatg aggatgatga 4860
agatgatgat gatgatgata atgagaatga tagcgaggat gacgctaatc agaccgatga 4920
agatgatgaa gaggtatgca aaattatttt aggtttggtc acacattttt gggatttata 4980
tcttgctaag ttcatgatta atggctgtac tagatagaat ctttctaagt tcgcgtgggc 5040
gaggacttta tatctgattt aggcacgttt acattttctc tacaaattag aacagatttt 5100
caaaaaatgt tttttaagaa aatggggaag ataatgttga cttgatgtgt cccacaattc 5160
tgccacaaac caaacatcct gtctctggcc tgtttgtttc tagttgaatc ttggtgtttg 5220
accaaaatac tgcatgatgg ttcatcttct attacggata ctgtatacgt taatatgaag 5280
tccatggttc tcatggcatc cttctgagat ttatagctat tgtgtatctt tcatttcctc 5340
tcaatcatac tgtgtggtta ttaatctgta atcctaaaac tgttttcata gtctgatgaa 5400
gatgccgaga agaatgtcga tatgtccact gctgtatcgc taataaaggg tgaacgtaag 5460
aacttgcgaa gaaaagagaa gcgtgctcgg atagagcaaa ttcgggttat gctcggtctt 5520
tctgattctc aaaggactcc aatggtgatg ttgtaatcaa catttttttt gttctaaatt 5580
tgtttgaagt tgttccgaca aagtacatat actttgttta ctcagaggaa ctcttggctg 5640
ataatttgtt acacacagtt aacaattaaa accatatatc actaattccc atattcacac 5700
ttttaaagcc aggagagtca ctaaaagatt tctacaagag aacggatatg tactggcaga 5760
tggctgcata tgagcacact caacacactg gaaaggttag tttctgctcc ttaagtatct 5820
tcacccgtca tacctgttat catattctct aggttgctgg cagtatgagt ttgctgtatt 5880
tattcgtgct catccaatgc caggagctcc gcaaagatgg ttttgatctt gccgaaactc 5940
gatataagga actgaaaccc atactcgacg aggtaaaatt gtcatgttgt gtcccctttg 6000
agacaaaacg gtatttctga cttggtacat attaactgac tcttacacgc cctcttcagc 6060
tggctgtgct cgaggctgaa cagaaagctg aggaagaggc tagtgcttcc actagttcca 6120
agaaagacac gaagaaaagc aagcagaaga gtggcatgag atag 6164
<210> 2
<211> 1989
<212> DNA
<213> 水稻(Oryza sativa)
<400> 2
atggggaaga agcagaagaa gcccaggaag gggaaggaga agacggagcg gaagacggcc 60
aagggcgagg agaagcgcgc ccgccgcgag gcccggaagg tcggcgagga ggacgacatc 120
gacgccatcc tcaggagcat acaaaaggag gaggctaaga agaaggaggt acatatagat 180
gagaatgtcc ctgcaccatc tccccggtcc aattgctcgc ttacaataaa tcccctgaaa 240
gatacagaat tggttctgta tggaggagag ttctacaatg gcagcaagac ctttgtttat 300
ggtgatcttt atcgctacga tgtagagaaa aatgagtgga agttggtatc tagtcctaac 360
agtcctcctc cacgaagtgc tcaccaaaca gttgcctgga agaataatat atacatgttt 420
ggtggggaat tcacttcgcc aaaccaagaa cgttttcatc attacaagga cttttggtca 480
ttggatctaa aaacaaatca atgggagcaa attcttgcga agggttgtcc aagtgcacgt 540
tcagggcaca ggatggtcct ctataagcac aagatcgtgc tatttggtgg tttttatgac 600
actcttaggg aagtgagata ctacaatgac ttacatgttt ttgatttaga taatttcaag 660
tgggaggaga tcaagcctcg ccctgggtgc ttgtggccaa gtccaagaag tggctttcag 720
ctaatggtat accaagatca gatatatctg tatggcggat attttaaaga agtagtttct 780
tctgacaaat ctgcatcaga aaaaggaaca gttcatgcag atatgtggac tcttgatcct 840
cgtacttggg agtggaataa ggttaagaaa actgggatgc cacctggccc cagagctggg 900
ttttctatgt gcgttcacaa gaaaagggct gttcttttcg gtggtgtggt agatatggaa 960
attgaagggg atgtcattat gagcatgttt atgaatgagc tctatggttt ccagctggac 1020
aaccatcgct ggtatccttt agagctcagg aaagacaagc ctgctaaaaa taagacaaag 1080
gacatcaaaa gaaaagaacc atcgaacaat gtggaagata atcttggtaa tgaggaggat 1140
gagatcatgg aggactcaga aactactgga gggcaatccg aagtccatgg ggtttcgaat 1200
cacttgacca agagtctaac cttaaataaa gctggctcag gcaatagctc tgatattctc 1260
tctgattcga caacacaaga agtactccca gaggcagtga aacccggtgg tcggatcaat 1320
gcatgcttgg ctgtagggaa agatacactc tatttatatg gaggaatgat ggaattgaaa 1380
gatagagaaa ttactcttga tgatatgtat tcacttaacc ttagcaaact agatgagtgg 1440
aagtgtatca taccggcatc tgaatctgaa tggctagaaa tttctgaaga tgaggatgat 1500
gaagatgatg atgatgatga taatgagaat gatagcgagg atgacgctaa tcagaccgat 1560
gaagatgatg aagagtctga tgaagatgcc gagaagaatg tcgatatgtc cactgctgta 1620
tcgctaataa agggtgaacg taagaacttg cgaagaaaag agaagcgtgc tcggatagag 1680
caaattcggg ttatgctcgg tctttctgat tctcaaagga ctccaatgcc aggagagtca 1740
ctaaaagatt tctacaagag aacggatatg tactggcaga tggctgcata tgagcacact 1800
caacacactg gaaaggagct ccgcaaagat ggttttgatc ttgccgaaac tcgatataag 1860
gaactgaaac ccatactcga cgagctggct gtgctcgagg ctgaacagaa agctgaggaa 1920
gaggctagtg cttccactag ttccaagaaa gacacgaaga aaagcaagca gaagagtggc 1980
atgagatag 1989
<210> 3
<211> 662
<212> PRT
<213> 水稻(Oryza sativa)
<400> 3
Met Gly Lys Lys Gln Lys Lys Pro Arg Lys Gly Lys Glu Lys Thr Glu
1 5 10 15
Arg Lys Thr Ala Lys Gly Glu Glu Lys Arg Ala Arg Arg Glu Ala Arg
20 25 30
Lys Val Gly Glu Glu Asp Asp Ile Asp Ala Ile Leu Arg Ser Ile Gln
35 40 45
Lys Glu Glu Ala Lys Lys Lys Glu Val His Ile Asp Glu Asn Val Pro
50 55 60
Ala Pro Ser Pro Arg Ser Asn Cys Ser Leu Thr Ile Asn Pro Leu Lys
65 70 75 80
Asp Thr Glu Leu Val Leu Tyr Gly Gly Glu Phe Tyr Asn Gly Ser Lys
85 90 95
Thr Phe Val Tyr Gly Asp Leu Tyr Arg Tyr Asp Val Glu Lys Asn Glu
100 105 110
Trp Lys Leu Val Ser Ser Pro Asn Ser Pro Pro Pro Arg Ser Ala His
115 120 125
Gln Thr Val Ala Trp Lys Asn Asn Ile Tyr Met Phe Gly Gly Glu Phe
130 135 140
Thr Ser Pro Asn Gln Glu Arg Phe His His Tyr Lys Asp Phe Trp Ser
145 150 155 160
Leu Asp Leu Lys Thr Asn Gln Trp Glu Gln Ile Leu Ala Lys Gly Cys
165 170 175
Pro Ser Ala Arg Ser Gly His Arg Met Val Leu Tyr Lys His Lys Ile
180 185 190
Val Leu Phe Gly Gly Phe Tyr Asp Thr Leu Arg Glu Val Arg Tyr Tyr
195 200 205
Asn Asp Leu His Val Phe Asp Leu Asp Asn Phe Lys Trp Glu Glu Ile
210 215 220
Lys Pro Arg Pro Gly Cys Leu Trp Pro Ser Pro Arg Ser Gly Phe Gln
225 230 235 240
Leu Met Val Tyr Gln Asp Gln Ile Tyr Leu Tyr Gly Gly Tyr Phe Lys
245 250 255
Glu Val Val Ser Ser Asp Lys Ser Ala Ser Glu Lys Gly Thr Val His
260 265 270
Ala Asp Met Trp Thr Leu Asp Pro Arg Thr Trp Glu Trp Asn Lys Val
275 280 285
Lys Lys Thr Gly Met Pro Pro Gly Pro Arg Ala Gly Phe Ser Met Cys
290 295 300
Val His Lys Lys Arg Ala Val Leu Phe Gly Gly Val Val Asp Met Glu
305 310 315 320
Ile Glu Gly Asp Val Ile Met Ser Met Phe Met Asn Glu Leu Tyr Gly
325 330 335
Phe Gln Leu Asp Asn His Arg Trp Tyr Pro Leu Glu Leu Arg Lys Asp
340 345 350
Lys Pro Ala Lys Asn Lys Thr Lys Asp Ile Lys Arg Lys Glu Pro Ser
355 360 365
Asn Asn Val Glu Asp Asn Leu Gly Asn Glu Glu Asp Glu Ile Met Glu
370 375 380
Asp Ser Glu Thr Thr Gly Gly Gln Ser Glu Val His Gly Val Ser Asn
385 390 395 400
His Leu Thr Lys Ser Leu Thr Leu Asn Lys Ala Gly Ser Gly Asn Ser
405 410 415
Ser Asp Ile Leu Ser Asp Ser Thr Thr Gln Glu Val Leu Pro Glu Ala
420 425 430
Val Lys Pro Gly Gly Arg Ile Asn Ala Cys Leu Ala Val Gly Lys Asp
435 440 445
Thr Leu Tyr Leu Tyr Gly Gly Met Met Glu Leu Lys Asp Arg Glu Ile
450 455 460
Thr Leu Asp Asp Met Tyr Ser Leu Asn Leu Ser Lys Leu Asp Glu Trp
465 470 475 480
Lys Cys Ile Ile Pro Ala Ser Glu Ser Glu Trp Leu Glu Ile Ser Glu
485 490 495
Asp Glu Asp Asp Glu Asp Asp Asp Asp Asp Asp Asn Glu Asn Asp Ser
500 505 510
Glu Asp Asp Ala Asn Gln Thr Asp Glu Asp Asp Glu Glu Ser Asp Glu
515 520 525
Asp Ala Glu Lys Asn Val Asp Met Ser Thr Ala Val Ser Leu Ile Lys
530 535 540
Gly Glu Arg Lys Asn Leu Arg Arg Lys Glu Lys Arg Ala Arg Ile Glu
545 550 555 560
Gln Ile Arg Val Met Leu Gly Leu Ser Asp Ser Gln Arg Thr Pro Met
565 570 575
Pro Gly Glu Ser Leu Lys Asp Phe Tyr Lys Arg Thr Asp Met Tyr Trp
580 585 590
Gln Met Ala Ala Tyr Glu His Thr Gln His Thr Gly Lys Glu Leu Arg
595 600 605
Lys Asp Gly Phe Asp Leu Ala Glu Thr Arg Tyr Lys Glu Leu Lys Pro
610 615 620
Ile Leu Asp Glu Leu Ala Val Leu Glu Ala Glu Gln Lys Ala Glu Glu
625 630 635 640
Glu Ala Ser Ala Ser Thr Ser Ser Lys Lys Asp Thr Lys Lys Ser Lys
645 650 655
Gln Lys Ser Gly Met Arg
660
<210> 4
<211> 6156
<212> DNA
<213> 水稻(Oryza sativa)
<400> 4
atggggaaga agcagaagaa gcccaggaag gggaaggaga agacggagcg gaagacggcc 60
aagggcgagg agaagcgcgc ccgccgcgag gcccggaagg tcggcgagga ggacgacatc 120
gacgccatcc tcgtacgtgt gctccctccc tcccgccccc ctcctctgcg tcagctcttc 180
acgctcgctc agtgcgctct agctcgatcg gcgtctccat gtgcggtttt gtttgctcac 240
caccaccttc tgcgtctcgt gatgcactcc ggtggctgaa aaattggaag cgattttcgc 300
actggctcac ctttttccct tcacatttcg ttgcgtagca gctatggatt ttagatgagt 360
ttggtgttgc tatgtgtgct gtttgaactt tttttttttg ctgatttatc tatttgctct 420
ggttattttt cattgcagag gagcatacaa aaggaggagg ctaagaagaa ggaggtacat 480
gtagatgaga atgtccctgc accatctccc cggtccaatt gctcggtaag acattttaga 540
gcaagtgcca gctaaagaag ttaatctttc ggtattcttg tgattatgta cttgaagtgg 600
actaggtatc atttttatgc ttgtggtggt catgtttgca tgcttagaat ttatatactc 660
cagtaggatt aataaatttc ttcaggcaag atggattttt ttatgaagta atttgatcaa 720
atatgatgat cttttggtga tactgaaccg attggttttt cagttccgga taggtgatgt 780
caaattacaa ttcaggggca aaatgtgtta aaagatagat gttcgttttg tttttttttt 840
tgttaactgt tggaaaaagt ttttgatgtt gtacagagat cctccctcta aatgttaact 900
ataacaaata aacctgcata tgatcttcca gctgaccaag ttaattcctt cttaaatgca 960
gcttacaata aatcccctga aagatacaga attggttctg tatggaggag agttctacaa 1020
tggcagcaag gtgggagaca tatccttcac gatttcactg ctggttgaat gagatatgtt 1080
catctagtgt tctttttgcc tctattctgc ttctagaggt ttttgagtac tgaaatagta 1140
tttcttttcc tcgccacaat ctatttctgc agacctttgt ttatggtgat ctttatcgct 1200
acgatgtaga gaaaaatgag tggaagttgg tatctagtcc taacagtcct cctccacgaa 1260
gtgctcacca aacagttgcc tggaagaata atatatacat gtttggtaac ataacttaac 1320
tttgggaagg cattctatga gttaaaatgc tttttcagta tgtatgattt agttttttat 1380
tctgtgttga tgtttcaggt ggggaattca cttcgccaaa ccaagaacgt tttcatcatt 1440
acaaggtaga agactaattt tgtcagtcta tttcttgtga ctgttttgag tatcttctct 1500
tggcacaagg catttgcaca attcgtatat aagcagtagc ttcataagca atacatatct 1560
ggcatgattt tttttcaatt ttaaaccaag agataggtac tgattcccat gttcttacat 1620
aattataagt tgaaatatga ctaatggaga ctaatatgtg cagttcttct gctatttatt 1680
gatttaagat taagatgtag aggatgcggg gttcacctgc ccttttattt ggatttgcac 1740
atataacatc actttttatt cctgattgcc tttttttttc ctttctgttg tgatattaat 1800
ggggttattt cgagattagc tatcttagat ggaaaactca ttattatggt tctattcaaa 1860
tttctggtct gattttgtac tgtaggactt ttggtcattg gatctaaaaa caaatcaatg 1920
ggagcaaatt cttgcgaagg gttgtccaag tgcacgttca gggcacagga tggttagtgg 1980
ttttacatta aaatcagtca tcaactattc tgctcccctc ttttcacctt aatatttctg 2040
tattatgagc agtagtaaat tgtgttcttt ttcctacagg tcctctataa gcacaagatc 2100
gtgctatttg gtggttttta tgacactctt agggaagtga ggttagtaca gttacatttt 2160
atatgactct accctggtaa tcttgttatt agagtaacat ttatatttgg accatgtcta 2220
gaagtagtgg agtcattatg tccaaactac aaattatgca ctacctgaat aacagtgtag 2280
cactcttaca gctgatcatc cgcaaagaat gaaatggtgt ggcgtagagt acgttctgaa 2340
taaatagtgt gacatgacat gatctgatct gcattttttt tataatatcc ttctgcagat 2400
actacaatga cttacatgtt tttgatttag ataatttcaa ggtgagtaca ccatgttaat 2460
attttgttta atactgttag tagtaacaca tgaagtcatt tattttaata ctcttactgg 2520
gaatatttgt atttcagtgg gaggagatca agcctcgccc tgggtgcttg tggccaagtc 2580
caagaagtgg ctttcagcta atggtatacc aagatcaggt aggtcttttt ggatttaaag 2640
ctaggacatt gatacttcat aaaaagagtt taaattaact ataaaccaac cttgtcttcg 2700
actacttttg tttacaagag tattaatggc ccttatttct gtagatatat ctgtatggcg 2760
gatattttaa agaagtagtt tcttctgaca aatctgcatc agaaaaagga acagttcatg 2820
cagatatgtg gactcttgat ccttgtactt gggagtggaa taaggtgatc tcttgcaatt 2880
ttttagaaca ttgtatcaac ttccatcatg atagtgtatc gagttttact ttaagccata 2940
tatccactga gtgatttgca tattattacc ttcacttgat ttcttaatag gttaagaaaa 3000
ctgggatgcc acctggcccc agagctgggt tttctatgtg cgttcacaag aaaagggctg 3060
ttcttttcgg tggtgtggta gatatggaaa ttgaaggtta ttttcagctc aattttgctc 3120
tgtgcatagc tacttaggtt attttactaa gtatttgaaa taccacgtgt gtcaagtttg 3180
ttccttttct gtagaagttc tcaaggccta actgtagaac ccaattttgt gattgcaggg 3240
gatgtcatta tgagcatgtt tatgaatgag ctctatggtt tccagctgga caaccatcgc 3300
tggtatactt caatactcca tttgagtatc ttgtgttttt aagtaacaca ggctagtttt 3360
atccctgctt attttgttcc ttgctatttt tgtgtattgt tgcaactttc tttattatta 3420
tttaacagtg cagctaacta atgtcacttc ctattccgac atgcaaactg cttctaacta 3480
ggtttaaaat tattaaaggt ctaaatcttt ctcggctgat tgatggttat gttcatctat 3540
cgtcttagga tgaactttgt ttgtaatctt gtggttatct agataacata actactttga 3600
gaattgttca gtgatattat tgtttactct tgggatccct tcgtagcata ttattgttta 3660
cttcagcgga attgttcagc gatgttggtt ttggttgaaa ctactggtcc acggctcaca 3720
tattctccaa tttcaattgt ccacactggt gcatagatga gaaatgttat gctttcttat 3780
ttagcttcat ttttgtgtgt ctatgttcag attattttcc ttgctgtttc aggtatcctt 3840
tagagctcag gaaagacaag cctgctaaaa ataaggtgat tcctcgaatt caatcatatg 3900
accagccatc tcatgtttat agtttgtatt ttttaggtac ttgatacaca gtctactctg 3960
caggaatcta tttcattagt tcagacataa agactaagca tgtagtatag ggtatgcctg 4020
tttttctttt taataagcat atatgatatg tacatattct catatactat tatactgtta 4080
ttttcttaat aagcatatgc gatctgtcga catattcaca ttgactatta tatagtaata 4140
tttggatcat tgcttaaaag cagtttgtgc ttcttttttt ttttttgggt gccatgtgta 4200
gacaaaggac atcaaaagaa aagaaccatc gaacaatgtg gaagataatc ttggtaatga 4260
ggaggatgag atcatggagg actcagaaac tactggaggg caatccgaag tccatggggt 4320
ttcgaatcac ttgaccaaga gtctaacctt aaataaagct ggctcaggca atagctctga 4380
tattctctct gattcgacaa cacaagaagt actcccagag gtattgcagc tgttctttta 4440
gatgttgaca tttacattct aatgatcttt tgtttctcat tagcatttgc tgcttacagg 4500
cagtgaaacc cggtggtcgg atcaatgcat gcttggctgt agggaaagat acactctatt 4560
tatatggagg aatgatggaa ttgaaagata gagaaattac tcttgatgat atgtattcac 4620
ttaaccttag caaactagat gagtggaagt gtatcatacc ggtcagttgc agattggccc 4680
cttctttttg ccattttgtt gtttaactaa tagtgttgtt tatttcaatc agagataaca 4740
gaaaattatc tgttatcatt tttgactatt cacatttgtt tctgaacttg ccttactcac 4800
ctttcttatg caggcatctg aatctgaatg gctagaaatt tctgaagatg aggatgatga 4860
agatgatgat gatgatgata atgagaatga tagcgaggat gacgctaatc agaccgatga 4920
agatgatgaa gaggtatgca aaattatttt aggtttggtc acacattttt gggatttata 4980
tcttgctaag ttcatgatta atggctgtac tagatagaat ctttctaagt tcgcgtgggc 5040
gaggacttta tatctgattt aggcacgttt acattttctc tacaaattag aacagatttt 5100
caaaaaatgt tttttaagaa aatggggaag ataatgatgt gtcccacaat tctgccacaa 5160
accaaacatc ctgtctctgg cctgtttgtt tctagttgaa tcttggtgtt tgaccaaaat 5220
actgcatgat ggttcatctt ctattacgga tactgtatac gttaatatga agtccatggt 5280
tctcatggca tccttctgag atttatagct attgtgtatc tttcatttcc tctcaatcat 5340
actgtgtggt tattaatctg taatcctaaa actgttttca tagtctgatg aagatgccga 5400
gaagaatgtc gatatgtcca ctgctgtatc gctaataaag ggtgaacgta agaacttgcg 5460
aagaaaagag aagcgtgctc ggatagagca aattcgggtt atgctcggtc tttctgattc 5520
tcaaaggact ccaatggtaa tgttgtaatc aacatttttt ttgttctaaa tttgtttgaa 5580
gttgttccga caaagtacat atactttgtt tactcagagg aactcttggc tgataatttg 5640
ttacacacag ttaacaatta aaaccatata tcactaattc ccatattcac acttttaaag 5700
ccaggagagt cactaaaaga tttctacaag agaacggata tgtactggca gatggctgca 5760
tatgagcaca ctcaacacac tggaaaggtt agtttctgct ccttaagtat cttcacccgt 5820
catacctgtt atcatattct ctaggttgct ggcagtatga gtttgctgta tttattcgtg 5880
ctcatccaat gccaggagct ccgcaaagat ggttttgatc ttgccgaaac tcgatataag 5940
gaactgaaac ccatactcga cgaggtaaaa ttgtcatgtt gtgtcccctt tgagacaaaa 6000
cggtatttct gacttggtac atattaactg actcttacac gccctcttca gctggctgtg 6060
ctcgaggctg aacagaaagc tgaggaagag gctagtgctt ccactagttc caagaaagac 6120
acgaagaaaa gcaagcagaa gagtggcatg agatag 6156
<210> 5
<211> 2237
<212> DNA
<213> 水稻(Oryza sativa)
<400> 5
actaattaca tggaatgcgt gtaaattgtg agatgaatct tttaagtcta attgcgccat 60
gatttgacaa tgtggtgaca gtaaacattt gctaatgacg gattaattag gcttaataaa 120
ttcgtctcgc ggtttacaga cagattctgt aatttatttt attattagac tacgtttaat 180
acttcaaatg tgtgtccgta tatccgatgt gacacgccaa aacttttaca cctcttgata 240
taaacacagc gtagctttct tgctaactcg atattttctt accgtagtca catgtcacgt 300
ctccgatacc atctcaataa ttgcttttga agttattctc taatttaata gtcgaagaag 360
tcgatatacc ctatttattg ttgagggata tgaagtaaat ctgacccata gttgaggtag 420
tcaggactgt aggtttcaaa gtacgattta gcccatttag gttgttgggc caaacgctct 480
tccgttttag aagagacgtg gtcactctgc caaaaatagg aaaaagtaca cccaaggtcc 540
ctcaacttgt catagggata aaaaacgtcc tcaaatcaca aaaccagata tacggggtct 600
attaattata taaaaccggt cattagaggt ccttcggcgg tcttgaaccc ggttttatct 660
gacgtagcgg ctaaatcagt gcgggacccg cgtgggcccc acatgtcagc tggccacgtc 720
atcaaactcc tctctctttt cccctcctct ctctcttcct catctctctc ccttctctgc 780
cgccggcagt gcctcggcgg cgggcatcgg cggtgatggc ggcggcggtg gggggagcgg 840
catacgggct ccccggcggt cgccgtccac gcatagctcc tccccccgca gcttgcgccc 900
accgcctccg gtgcgccgcc gcctccccgt cggcgaggac gcggtgcttg aggttggata 960
tgtcgaagca gcagaggaag catgtaggct tggtgcagct ggggatgtag tcgcggagca 1020
tgtcagcgac gccgcggtag cggcggaacg cgaggagaga gcacgcccgg agctcgagag 1080
cggcctccat gcgcggcgac aactctagca ctgcttccac tagtccaagc gccgtcgtgg 1140
ccgctgagtg gtcgccacat tccccattga gcggcgcggc ggcagcgaaa gcagcccgtg 1200
cctcaatgag gtagtcgccg atgatctgct gcacaatgca gagcaccaac aaatcagttg 1260
ctgaagccaa aggcaaaggc aaatctaaat tgggagcaaa accgcccaaa ttgccatcat 1320
tttttgtggc acattagatg aagaattaaa gagggagcaa gaaacaacct ttcggtgtcc 1380
gcggagccaa atccttctct tgtcaggagg ggaagaagac ggcgaggaca ccgtcgccga 1440
ccaccgccgt tcgtccgcaa gccggctgtc gacgagatgt ccccgcgcag cctcgcacac 1500
ctgcagtgcc tcctcacgct cgggctccat cgccgacgag tggcggctgg gacgggctac 1560
tcgacccgct cgaccagaac ctccgccgcg aggtcctccg ctacggcgac ttcgtgcagg 1620
ccgcatacac agcgttccat tccatgctgt cggcggcggc ggcgtcgcag cacatctcgg 1680
gtgggcgcac cggacgctcg tgctccccga cctgcggcgg cgtcgcagca cagccagaag 1740
ggagagagag atgaggaaga gagagaagag gggaaaagag agaggagttt gatgacgtgg 1800
ccagctgaca tgtggggccc acgtgggtcc cgcgccgact cagccgccac gtcggataaa 1860
accggattca gaaccaccga aggacctcgg tgaccggttt tgtataatta agggaccccg 1920
tatatctggt tttgtggttc gaggatgttt ttttatcccc atgacaagtt gagggacctt 1980
cggtgtactt tttccaatgg agggagtagt atccaacccc agcgtccgac tccgactccc 2040
gcttgcacgc gttcgtcacg gcccgttgag gcccaactaa gtccaagatg ggccgtcggc 2100
ccacggtgcg aacgccggcg ccaccgttgt ccctgctgct ccccacgagg gttttaggca 2160
cgcctccgcc tccgcctccg cctccggcag caagtgagcg cggggagaga gacagaagcc 2220
ggcggcggcg gggcgag 2237
<210> 6
<211> 1318
<212> DNA
<213> 水稻(Oryza sativa)
<400> 6
ctttggttcg aaatatcgaa atagagagtg gtttaggcca atgctttaag ccatctggga 60
ttttttcctt tctgagccat tggtcagctc gatcgattaa tccacacgag gtgcctataa 120
ctacctgtgt aggttggctt aatccatggt taaaattttc caacgtactg tatttgcgat 180
tcctcggaag cattcatgga gagattgtac acagttctat tcatcggaaa ttcgttacta 240
cgtacaagtt ttgagttctc tactgttcag tgttcacagc ctgcgatgta acacggtaca 300
ccctaacagt atctgggtgc ggtgtaaatt cactccatcc gtttcaggtt accagacgtt 360
tatagacaaa ctattttaag tttgactaaa tttatagata aatatagtaa tatttataat 420
actaaattag tgtcatcaaa tcaataatcg aatatatttt cataataaat ttgtcttggg 480
tgaaaaatgt tgctattttt ttctataaac ttgatcaaac ttaaatcaat ttgagtttga 540
cgaaggttaa aacgttttat ggttgaaacg gagggagtac tcttctgtac acatattgtt 600
tttttcttct cgagtatatc gatcttgttc cagaaaaaaa aaaagagtat atcgatcaag 660
ttttctcccc atacgctcgt gctactgttc ttaggtcgaa acgtccatct tgtacctttg 720
tacggtggcc catggcgcca ctgctccagt gatgggtact atactacctg cggtgaggtg 780
atgcaaccgt gatggtggtg agcgggtggt ggtgggcgag caaccgatcg tccagcgaca 840
gctaaccaca ataacgaagc gagtacgcgc ttcacctcac caacagcgag ctcccgtttt 900
gttcgcacga aaagaagctg agctcgcgtg cgtgcggtgc gttttgtgtg cgctgcgatg 960
tgcgatggtt gattgtgtgg tgtgcgagga gaggagtaga aatcaggcgg gggggctttt 1020
aatatcctgc cgtgcccgtc atctgtgtgg gactttgacc acaccattta cttcagctca 1080
tcaaacctca acaaccataa ctgcaccacc tgtccgccca acggcccaac ccctgctggt 1140
cccggttcac cacgcctccg gtgaaccata catgcaacct aatgtgttca tggcacctaa 1200
tgtgccgttg gcaccgaaat tgaacggtac gtgcacccaa ggtgaaggtg aggcaaggtg 1260
ctcatggctg cagagtttag actttagagc aagcagttgt tctcctccgt acgaaatg 1318
Claims (7)
1.基因在调控植物粒型、粒重中的应用，其特征在于：所述基因具有如(a)、(b)、(c)所示的序列：
(a)Seq ID No：1所示的基因组核苷酸序列；
(b)Seq ID No：2所示的cDNA核苷酸序列；
(c)在(a)、(b)所示的核苷酸序列的中添加和/或取代和/或缺失一个或几个核苷酸而生成的可编码具有调控粒型粒重功能的蛋白质的突变基因、等位基因或衍生物；
所述粒型为粒长。
2.根据权利要求1所述的应用，其特征在于：正调控植物粒型、粒重。
3.如权利要求1或2所述基因编码的蛋白在调控植物粒型粒重中的应用，其特征在于，所述蛋白具有如(A)或(B)所示的序列：
(A)Seq ID No：3所示的氨基酸序列；
(B)在(A)所限定的氨基酸序列中添加和/或取代和/或缺失一个或几个氨基酸且具有相同功能的由(A)衍生的蛋白质。
4.一种基因互补载体，其特征在于：所述基因互补载体为pCAMBIA1300载体。
5.一种基因过表达载体，其特征在于：所述基因过表达载体为pCAMBIA1300S载体。
6.一种与水稻粒型粒重基因紧密连锁的分子标记，其特征在于，所述分子标记为P4-1、P4-2、P4-3、P4-4、P4-5、P4-6、P4-7、4-8或P4-9；所述各分子标记对应的引物序列分别为：
P4-1：
F,5’-TGGGTCTTCAAAAAATGTTCAGTGG-3’
R,5’-ACCCCGCCTAAACTCCATGAATC-3’；
P4-2：
F,5’-TATAGATTCATCGTACTAAGGC-3’
R,5’-TGAGATTTATTGTTTTGTGTG-3’；
P4-3：
F,5’-AGAAGTAGTGCAGAGTACAGTC-3’
R,5’-AGTACTCCTATCCTTTAATAATATG-3’；
P4-4：
F,5’-ATACGTAGCGTTTGGTTATAGC-3’
R,5’-TTCGGTTTTGAACTCAACTTC-3’；
P4-5：
F,5’-TGCTTGAAGAGGAGAATGGTGG-3’
R,5’-AGCTCCTGAGTTCCTTGCGTC-3’；
P4-6：
F,5’-TTACATTATCGAATTATGCACGATAC-3’
R,5’-TGATACCCGAACTTCCTGACTG-3’；
P4-7：
F,5’-TGTGGCAGAATTGTGGGACAC-3’
R,5’-ACTTTATATCTGATTTAGGCACGTTTAC-3’；
P4-8：
F,5’-TAGATTGGTTTTTATGAAACG-3’
R,5’-TGCTGTCACAGTTTATCACAC-3’；
P4-9：
F,5’-TCCTACGATTTCTCAATCCTG-3’
R,5’-TGAATTCCTTCAATTTTAGAGC-3’。
7.权利要求6所述的与水稻粒型粒重基因紧密连锁的分子标记在分子标记辅助选择育种中的应用，其特征在于：所述应用为辅助选择与植物粒型粒重相关性状。
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