CN112877340A - 一种水稻基因gsnl4及其编码的蛋白质的应用 - Google Patents
一种水稻基因gsnl4及其编码的蛋白质的应用 Download PDFInfo
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- CN112877340A CN112877340A CN202110224405.3A CN202110224405A CN112877340A CN 112877340 A CN112877340 A CN 112877340A CN 202110224405 A CN202110224405 A CN 202110224405A CN 112877340 A CN112877340 A CN 112877340A
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Abstract
本发明公开了一种水稻基因GSNL4(Seq ID No：1、2)及其编码的蛋白质(Seq ID No：3)的应用，所述基因和蛋白质用于调控水稻的粒型和叶型，还用于提高水稻粒重及高产育种。本发明采用图位克隆技术首次在水稻中克隆到GSNL4基因，并通过转基因敲除实验鉴定了该基因的功能；本发明通过对GSNL4基因的功能解读，为进一步阐明该基因在水稻产量形成的遗传机制及其作用机理，创建水稻高产新种质打下基础，并拓展了其在水稻高产育种中的前景探索。
Description
技术领域
本发明涉及植物基因工程领域，特别是涉及一种水稻基因GSNL4(grain size andnarrow leaf4)及其编码的蛋白质的应用。
背景技术
水稻是我国重要的粮食作物之一，也是单子叶植物的理想模式植物。在水稻超高产育种中，粒重是决定产量的三要素之一，一般用千粒重来衡量，其与籽粒的大小呈正相关，而籽粒大小又可分为粒长、粒宽和粒厚三个粒型指标。粒型作为水稻理想株型的重要形状，对水稻产量增加起到了至关重要的作用，因此水稻粒型的研究一直是国内外育种家研究的热点和重点。随着现代分子生物学和RFLP、RAPD及SSR等标记技术的进步发展，越来越多的粒型相关基因被克隆。目前克隆了水稻粒型相关基因有：包括GS3、PGL1、PGL2、APG、GL7/GW7、GL4、TGW3、qGL3、OsGSK3、DEP1、SRS1、SRS2、SRS5、SMG1、PGL1、APG、TGW6、Gn1a、GLW7/OsSPL13、GW8、An-1、BG1、BG2、FUWA、GIF1、OsmiR397、OsFBK12、OsAGSW1、GW6a(OsglHAT1)、GL7、GW8、GS2、SRS5等，它们主要涉及到了G蛋白信号途径、植物激素、泛素－蛋白酶体途径、转录因子、激酶、蛋白分子互作、物质运输及microRNA等。对控制水稻粒型基因的克隆有助于了解水稻籽粒的发育过程和相关的调控机制，从而促进高产优质水稻新品种的培育。
为了提高水稻产量，育种家通常会采用多基因聚合来寻求突破，但聚合多个产量基因往往会带来负效应。因此除了在现有资源的基础上，需进一步发掘新的粒型控制基因，开展相关基因的克隆与功能解析，明确这些基因的分子作用机制及其调控机理，从而针对性的开展产量相关基因的分子设计育种。
发明内容
本发明要解决的技术问题是提供一种能调控水稻粒型的基因及其蛋白质。
为解决上述技术问题，本发明主要是利用图位克隆技术来获得所述基因及蛋白质。具体本发明采用如下技术方案：
一、突变体gsnl4的分离和遗传分析：
本发明的水稻粒型叶型突变体gsnl4来自粳稻品种武运粳21(W21)EMS诱变。通过与野生型的正反交实验，证明该突变体受隐性单基因控制，其中，水稻突变体相对于野生型，叶片变窄，籽粒宽度变窄，籽粒长度减小。
二、图位克隆GSNL4：
1)GSNL4的初步定位：
为了分离GSNL4基因，本发明首先组建了一个定位群体，由gsnl4与籼稻品种TN1杂交配组获得F2定位群体，再通过图位克隆的方法，利用STS、SSR等分子标记对GSNL4位点进行初步定位，将其初步定位在第4染色体上，并介于P1和P2两标记之间。
2)GSNL4的精细定位：
通过对P1和P2两个标记之间的BAC序列分析，发展新的SSR、STS标记将GSNL4精确定位于M6和M7标记之间139kb范围之内。通过分析此区段开放阅读框(ORF)推测候选基因。
3)GSNL4基因的鉴定和功能分析：
本发明通过基因编辑手段获得两个等位的gsnl4突变体，并且粒型叶型变窄，籽粒长度减少、粒重或千粒重减少。
基于上述研究结果，本发明开发了其相应的应用。
一方面，本发明提供了一种水稻基因GSNL4的应用，所述基因用于调控水稻的粒型和叶型，所述基因如(a)、(b)或(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)或(b)所示的核苷酸序列中添加和/或取代和/或缺失一个或几个核苷酸而生成的可编码调控水稻粒型和叶型的蛋白质的突变基因、等位基因或衍生物。
进一步地，利用所述基因转化水稻细胞，再将转化后的水稻细胞培育成植株。
进一步地，所述粒型为粒宽和粒长，所述叶型为叶宽。
进一步地，所述基因用于提高水稻粒重及高产育种。
另一方面，本发明提供了一种水稻基因GSNL4编码的蛋白质的应用，所述蛋白质用于调控水稻的粒型和叶型；所述蛋白质如(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。
进一步地，所述粒型为粒宽和粒长，所述叶型为叶宽。
进一步地，所述蛋白质用于提高水稻粒重及高产育种。
本发明利用一个引起水稻籽粒和叶片变窄的突变体，通过图位克隆技术首次在水稻中克隆到了GSNL4基因，并利用转基因敲除实验鉴定了该基因的功能。通过对GSNL4基因的功能解读，为进一步阐明该基因在水稻产量形成的遗传机制及其作用机理，创建水稻高产新种质打下基础，并拓展了其在水稻高产育种中的前景探索。
附图说明
上述仅是本发明技术方案的概述，为了能够更清楚了解本发明的技术手段，以下结合附图与具体实施方式对本发明作进一步的详细说明。
图1是突变体gsnl4与野生型材料的表型，其中(a)为整个植株表型对比图；(b)为叶片表型对比图；(c)为籽粒宽度对比图；(d)为籽粒长度对比图；
图2是GSNL4基因在水稻第4染色体上的初步定位图；
图3是GSNL4基因的精细定位图；
图4是pCAMBIA1300-CAS9-GSNL4载体图谱；
图5是PCA1301-GSNL4载体图谱。
具体实施方式
实施例1：
1、水稻材料：
水稻(Oryza sativa L.)粒型叶型突变体gsnl4(grain size and narrow leaf4)的原始野生型为粳稻品种武运粳21(W21)。如图1所示，为突变体gsnl4与野生型材料的表型，从图中可以看出，水稻突变体相对于野生型，叶片变窄，籽粒宽度变窄，籽粒长度减小。
水稻(Oryza sativa L.)粒型叶型突变体gsnl4的获得过程具体如下：
粳稻品种武运粳21的种子经过EMS诱变后，种植于大田，收取种子继续种植，从M2代中分离出来的一个窄粒窄叶突变体，暂时将其命名为gsnl4(grain size and narrowleaf4)。将该突变体进行多代自交，获得能稳定遗传的突变体植株。
将突变体材料gsnl4和野生型材料武运粳21进行正反交，即以突变体gsnl4为母本，武运粳21为父本进行杂交；同时以武运粳21为母本，gsnl4为父本进行杂交。获得的F1带植株均变现为野生型性状，说明控制该性状的基因受隐性核基因控制。正反交获得的F1代植株自交获得F2群体，表现出野生型表型的单株与突变体表型的单株数的遗传分离比符合3:1，说明该表型是由一对隐性核基因控制的。
2、分析和定位群体：
纯合的gsnl4突变体和籼稻品种TN1进行杂交，F1代自交，得到2965株F2群体；并从2965株F2群体中选出720株具有gsnl4突变表型的个体(即表现为窄粒窄叶)作为定位群体。在成熟期每株收取全部突变体表型的叶片，用来提取总DNA。
3、SSR和STS标记定位GSNL4基因
采用CTAB法提取用于基因定位的水稻叶片基因组DNA。取大约0.2g水稻叶片置于2ml EP管内，直接加入CTAB和钢珠，利用植物组织研磨器将组织破碎，氯仿抽提，乙醇沉淀，ddH2O2溶解，最后获得基因组DNA。每一个PCR反应用1μl DNA样品，体系为10ul。
GSNL4基因的初步定位：从gsnl4与TN1杂交组合的F2群体720株隐性单株中随机选取21株隐性单株，组成混池，利用已公布的234对近似均匀分布于各条染色体上B套引物，获得连锁位置，然后利用196株隐性单株，利用混池定位确定的连锁且具有多态性的引物，根据已知的反应条件进行PCR扩增，具体如下：
与目标基因连锁的STS引物为：
P1F：AAGTGCGGCTGTTTGATTT
P1R：CACCCACAGAGTTCTTCCA
PCR反应体系为：水稻基因组DNA 1ul,2×PCR Mix 5ul，10cM F/R引物各1ul，ddH2O2 2ul，总体系10ul。
PCR扩增条件具体为：94℃预变性4分钟；94℃变性30秒，58℃退火30秒，72℃延伸30秒，35个循环；72℃补齐10分钟；
经4％琼脂糖凝胶电泳分离和Gel-Red染色，检测PCR产物的多态性，将GSNL4初步定位在第4号染色体长臂上STS标记P1和P2之间，如图2所示。
GSNL4基因的精细定位：利用gsnl4与TN1组合的F2群体中共计720(720-196＝324)株隐性个体，在初定位的基础上继续设计STS标记，最终将GSNL4精确定位于M6和M7标记之间139kb范围之内，如图3所示。
STS标记引物序列为：
M6F:5’-TGAGCTGTACAAGCAAACGC-3’
M6R:5’-GGGAGAAATCCTCGAATTGG-3’；
M7F:5’-CGGTACATCACGGTATCAAATCG-3’
M7R:5’-TAAATGCTGGAGCGATGCTAACC-3’.
PCR反应体系为：水稻基因组DNA 1ul，2×PCR Mix 5ul，,10uM F/R引物各1ul，ddH2O2 2ul，总体系10ul。
PCR扩增条件具体为：94℃预变性4分钟；94℃变性30秒，58℃退火30秒，72℃延伸30秒，40个循环；72℃补齐10分钟。
产物检测：在含有Gel-Red的4.0％的琼脂糖凝胶电泳，紫外灯下观察并拍照记录结果。
4、基因预测和比较分析：
根据精细定位的结果，在139kb范围内根据RAP-DB(http://rapdb.dna.affrc.go.jp/)的预测，发现在此区间内共有9个候选基因，根据网站中基因功能的预测，我们首先设计了一个基因的测序引物，采用PCR方法分别从gsnl4和野生型品种基因组中扩增出这一个候选基因进行测序分析。具体如下：
目标基因测序引物的序列：
S1F:TTCAAGTCTGGGCAATGCAC
S1R:CCACCGCGCCATAAACTTTA
S2F:TAAAGTTTATGGCGCGGTGG
S2R:TGCGCAGAATAGTTCAGTCG
S3F:CGACTGAACTATTCTGCGCA
S3R:ATAATCCCTTGTGGCGAGCA
S4F:TGCTCGCCACAAGGGATTAT
S4R:CGTCACCTCAACCTTCACAC
S5F:GTGTGAAGGTTGAGGTGACG
S5R:CTGCAATGGAAGGACTGGAA
S6F:TTCCAGTCCTTCCATTGCAG
S6R:TGCTCCTCCCCAAACAGATT
PCR扩增体系:水稻基因组DNA 5ul，2×KOD Buffer 25ul，2mM dNTP 10ul，10uMF/R引物个3.0ul,KOD FX DNA聚合酶1ul，ddH2O 3ul，总体系50ul。
PCR扩增条件:94℃预变性4分钟；98℃变性1分钟，60℃退火30秒，68℃延伸1分钟，32个循环；68℃补齐10分钟。
发现其中该基因的基因组DNA片段中，突变体gsnl4扩增的产物与野生型品种比较有1个碱基突变。在该位点前后各150bp左右设计前后引物，利用该对引物，继续扩增野生型和突变体DNA，进行测序比较，并重复三次，发现突变体gsnl4与野生型比较在该位点均存在单碱基突变。根据RAP-DB的关于该基因的注释，该基因编码的是一个AGO蛋白。
检测引物为：
M-F：CAGTCCTTCCATTGCAGCTG
M-R：GTTCGTGAGGGTTTGCAACT
实施例2：
敲除载体构建：如图4，分别在其基因外显子上设计两个靶位点，首先构建到中间载体gRNA上，然后构建到pCAMBIA1300-Cas9敲除载体上。
基因编辑靶位点为：
gRNA1:AGAACTGGGTCTGGCAGCAC
gRNA2:ACATGCTCAGACCGCAGGGC
基因编辑检测引物为：
C1F：TCTCCTCTTTGCGCACCATT
C1R：TGGTATTGGACATGTGGGGC
C2F：CACCTGTAGCTGGAACTTGCT
C2R：CCCCCACCTGAAAAGTAGGAC
GFP载体的构建(如图5所示)。GFP载体引物序列为：
G-F:5’-TACAATTACAGTCGACATGGTGAAGAAGAAAAGAACTG-3’
G-R:5’-ATCCTCTAGAGTCGACGCAGTAAAACATGACACG-3’；
通过电击的方法将质粒转入农杆菌(Agrobacterium tumefaciens)株系EHA105中，然后通过农杆菌介导转化水稻愈伤。我们利用武粳稻21幼胚诱导的愈伤组织，经过诱导培养基培养3周后，挑选生长旺盛的愈伤用作转化的受体。用含有双元质粒载体的EHA105菌株侵染水稻愈伤，在黑暗、25℃条件下共培养3天后，在含有40mg/LHygromycin的筛选培养基上培养。筛选抗性愈伤在含有50mg/L预分化培养基上培养10天左右。将预分化的愈伤转至分化培养基上，在光照条件下培养。一个月左右得到抗性转基因植株。通过上述基因编辑手段获得一个与gsnl4等位的突变体(与gsnl4为同一基因，不同突变位点)，并且粒型叶型均变窄，籽粒长度减少、粒重或千粒重减少。证明本发明正确克隆到了相关基因。
以上列举的仅是本发明的若干个具体实施例。有必要指出，本发明不局限于以上实施例，对于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> 中农常乐（深圳）生物育种技术有限公司
<120> 一种水稻基因GSNL4及其编码的蛋白质的应用
<160> 3
<170> SIPOSequenceListing 1.0
<210> 1
<211> 14650
<212> DNA
<213> 稻属水稻(Oryza sativa)
<400> 1
atggcgctgc agttggagaa tggccgtccc catcatcatc aaggtatgcc tgcccatgcc 60
gtcgcccccc cacctccctc ggctctctcc cgttttcggc aaccctttgc cttttgaggc 120
gattctatcg ttttcttccc ctttttttcc tcccctcttc gtcctgtccc atcagatcgt 180
atacagttgg cgtcgaggcc gcgtccacac acgacgcgtc agtgcttgcg cgcgtgaggg 240
cgtgacacgg gttttaactg ttggtgcctg cgataattgt tcgacgcctg tgtgcttctg 300
ggtgagtttt ctcgtcgacc ctgtgtttgg ctgtcaccat gcggccccgg gcctgagtgt 360
ttagctgaca gagtgacagc ctacaggggt ttagccttgg cccctcgaga tcttttttca 420
ggttaggtta gtttgccatg ctgcctctgt taaatagagt cagctcgtta cccagcaagg 480
attagatcat cagctctttg ccaaacgccc caaaccgcta caacctgtaa acatacaggc 540
ctacattgat cagtcagtct ggagccacgg cacgagcgaa gccgatcgca cagtgctcca 600
ctgcgcgccc atgactgacg ccgctggtgc tatagctaca tggcatgttg gcatttgatc 660
cttgctgccg ttgatttcac tccgttgatt ttactcctcc tgagcaagcg gccgatcaga 720
atcatggagg aaaacaggag cagccatggc ggggcatcgc acgacggcta gctaaagttt 780
atggcgcggt gggggccata ggaatttgtg gaagcaaaaa cccatgtgcg gggtgcggcg 840
gcctccacag cattatggac gaggacgacg acgatatgga tgaggaggca gcactgccac 900
agcacggcgc ggtgcgtgcg ccacagtgcg gtgagctcgc tcgctgtgca cgcctccctc 960
ctccgccata tccaccacca gtcagtcgcc tcgtggaggc gtggatcccg gcgcctcccc 1020
ccctcgtgta gtgggacacg tttcgagcca ccagcggccc gacacacgtc gtaggcccgt 1080
gggagccgcg cgacgcgtgg tggtggtggt gggaggcggc cctcttctct cttggcccgc 1140
gctgcagatt cacgggcgct tgcactctcg gcctgcgggg cgtgggggtt acgggcccgc 1200
gcgtcagggg cgcgggacgg cggcgttggc tcggctcgct ggctggacct cccctgggcg 1260
catgtgcgcg ttcgcgaacg gtcacgtcgc gcgaccgtgg gtgggtccgt ccgtccggta 1320
tggagaccga gcgggtccct ctatagttct gatcatctca gggggaaaaa gaacgttttc 1380
ttcccttgca gtcgcatttt caacgcatga ttttctttac ggctgaaatg gattctgtaa 1440
attaaatcat gtggaattct ggctgtggtt taggacatta ggagatgagt aaactgactt 1500
aaaaaggaag cattagtcac tgtagttact accccttgac agatttagag gaaaaatgtc 1560
gataggaaat gaaaagttga tactccgttt gaaaaaagtc gatagaaaat gaaaagcatt 1620
agtcactgta gaccaaataa cacttggtct gttcggtgta gctaaactgc agctgcacaa 1680
cagtagccac taccgtgcat gataaagaaa aatcgataca ctggctgtac aacgcaaccg 1740
gttacagctg catagcaatg ttgccgaata gggccactta tgattgaagg aaaagtacat 1800
ataatttagc ttagagcctt agggtgtgtt cgctaggaga tgtcattaac caggaacagt 1860
agcacgcaaa acatagcggt ctattagcgc gtgattaatt aagtattagc ttttttttta 1920
aaatggatta ttttgacttt ttaagcaact ttcgtataga aactttttgc aaaagacgta 1980
ccgtttagca gtttaaaaag cgtgcacgcg gaaaacgagg gacagggttt gggaagagga 2040
atacaattgt aaaacagagg attgcaaaac acaggaatgg ccgtttgatt ggaccacagg 2100
aaaaacgcag gaatcagatg agagagatag actcagagga aatgttcaaa gaggttagac 2160
ctcttgctaa ctttcctcca aaatgtgcat aggattaccc attccatagg aattttaaag 2220
gattggatag gattcaatcc tttgtctcaa aggccttcat aggatttttt tccataggat 2280
tgaaatcctc caaaattcct atatttttcc tacaaatcaa aggggcccta aagtttttca 2340
aatcctatga aattcctatg gaatgtcaca ttgcatgtgt attttggaga aaatttagca 2400
agagctctaa cctcttggaa aatttccttt gagtctatct ctctcatccg attcctgcgc 2460
tccaattaaa cgaccattcc tgtgtttttc ctatgttttg caatcctctg ttttacactt 2520
taatcccttt cagaatcctg tgttttttct attcctccgt tttttctacc ctgctattca 2580
aagggaccct taatcctttt gaatcaaatg accaaatagg aaaattttct ataggattta 2640
aatcctatga aattcttata taaatcattt gattcaaagg aacccttaga ccatggggtt 2700
gaaagtgtta aggtcgagct tagttcctaa tattttcttc aaactttcaa cttttctatc 2760
acatcaaaac ttttctacac atacaaactt tttcgtcaca tcattccaat ttcaatcaaa 2820
gttttatttt tggcgtgaac taaacacacc ctaataaaca caccctaagt cctgccattg 2880
taggagcacg aaacacacat ttgagttgga ctttatgtaa ccgtaatcaa tgcaacggat 2940
gtgagagcgc atgtatatta cctatgcgta cgtgactcct tgtttttttt tttttgcagg 3000
gagagtatgt gaggccgagt ttagttttaa actttttctt caaacttcca acttttccat 3060
cacatcaaaa ttttcctaca cataaacttt caacttttcc gtcacatcgt ttcaatttca 3120
atcaaacttt caattttagc gtgtactaaa cacaccctga gtcctttgca ttgaccttat 3180
agtacgtctt ttctttgttg ctctgcatat tcttttctaa aatttctatt agttgcagtt 3240
gtactccttc catccaaaaa aaatacagtt gtataaaaaa tgtcccatct actagattaa 3300
gtttttttaa ggacggagag aactgaggga gtatgttact agtatgtaaa gtaatttgca 3360
aatgccccaa agtattaagg tgttcatggt acgatgcatg aatccacgac cgcgttattg 3420
agtgatccac gaatcaacat cttctttttc ctagtaaaac attttaatgt acggagtagt 3480
tacattgatt ttttttttgg gatgtttttc tttctacata tagttagaca tgatcttttg 3540
cctcttgtcg ctctgtgttg gcttattagc tgtaaccaac atttgaatcc taagaattaa 3600
aattgatttt gaaattgagg cttttgcact atagtctatc ttttagcttt ggcttaaaag 3660
acacaaataa tacgtacata aaaaaattac tcataaatca tacattttac tatctattac 3720
ccccgcttta tttctcattg ggatagcaag gtacctctac gtgtcctttt tttttcccta 3780
gcagaatgat ccagtatttc cggactgtgt agtccatcct agcgactgaa ctattctgcg 3840
catcgctatc ctaccaaaag cttctagggt ggagtagaac gcttccatca aacaaaattc 3900
tatagtactt catctgtctc aaaataaatg cagcgtctta tttaaaaaag attatgatta 3960
gtatttttat tgttattaga tgataaaaca tgaatagtac tttatgtgtg actaaatttt 4020
ttaatatttt ttataaattt tttaaataag acggatagtc aaaacgctaa acacgaatat 4080
ctatggcttc acttattttg ggacgaggta ctactactcc tatataagca ctggctactt 4140
tattaattta tcaatgtagc aggagcacaa aaggaggagc atcatacagt tggctcgtag 4200
tgtacacctc caatactgcc tgcagctctg cagctgcatt gctgcaagcg agagcgatag 4260
ccggtatagc tgcatcgatg gccgaagggg cttttgcctt tgtgaaggcc ggggcgattc 4320
tagtcggggg gaaaaggccc gaggcagccc agcgaccgac ctggcgctac gatgcgcgaa 4380
aaagggcccc cacacccaca cacctgcctc gtgggcccca cgggccgcag gcctcggtcc 4440
gcgggcccag cagagcgact cctggctgcg ttctcggtac ggccctacag gtgggcccct 4500
cgcgtctgtt cagtgtcctg tatacacagg gcaaaacatc cggtcacggc cgtgcgggcg 4560
tggtgtgtga ctgagcggtg ggccggagga tattgggagg cccagatgtc attcggtgag 4620
agcgggggag aagggtgagg ccgtgggtgc tgggcccacg cactggcgcg tgcgccccat 4680
aagcggaacc aatctgaacc atcgattccg gtagggtgtg tcgctgtgct gccgttggtg 4740
caattgcgag tgctgcacgc tgcgtcaccg ctgtgacctg tttgctgcat cgagcggcgc 4800
ccattgaccc gtttccctat cctttttacg ggtcggagtg gcctaaccaa aacgggacgg 4860
cctcgacagc gacagcgacg gcgacccacc cgccgtcctc atccgtttgc gccattattt 4920
cgtccacctg cacggcttca ccgctttgta gctgtagtag cagtagcaca agggcagcca 4980
tttccccagc catgttcagc cagcccagct cttggatttt gatgacggca ttggattagg 5040
cacgtactag gagtgctgat ctgcatggtt ccggttgatc gcgtggtgcg tacgggacac 5100
aaggcgatac tgatccaatt cacacacacg agagagagag aggaaaaaaa aaaagaaagg 5160
caagagtgat ccaatcagca gccgaaacgt ccctgggccc tgggggaatg gggagcaacg 5220
gagcgcgagg cgcagttacc aaacactctg acatcccggg cccggccgtc cgatccttaa 5280
tggtcgatta gtcgccatct tgaacccacc cgggccatca gcgacgacgc ccgtatcccc 5340
gcacgggccc cacgccgtca tcgacacagc cgcgtgccct ctcgttccgt acagccactg 5400
acgggtccgg cgcgacccga cccgcgcccc gcgacctgac acgagcgcac ccgtccttcc 5460
tctcctctgc actggcgctc gcttcggctg tttccccagc gtgtgcctca ccgctgctgc 5520
taattaaccg caagcgctcg tcgtctttcc ccttcctcaa aaaaggggga gggggggtgg 5580
tggaggcgga ggcggaggca gcagcagcag tgcggtagtg caagcgctag tggaggagtt 5640
gggaggaggc cccctagggt ttcccgagac cgcctccccc cgcgcctgcg ccgccgctcg 5700
ccgagcgcgc gctccggtaa tgcctcccgc tctctagatc tgtgtgtgtc tccccccgtc 5760
tgtcttcgct gattctgccg cgggggcgtg cggccgataa gttcgatcgg ttcaggggag 5820
ggtccggctc tggcatcgcc gcgtggttat ggtgatggta cggccatgag agagcgcggg 5880
ttggtttgga cggggtttgt cggtggattt gggcggatct agttctcggc gagctgatcc 5940
gatagggggc tttggcgatt cgctcgtgtt ctgtgtgatc gcgtttggat ttttggattt 6000
agtactaatg gtgcgtgcga tacgagttgg tgcatcgcat gagaaaactc ttttcctttg 6060
tgtggtttga agtgtgtaca tttgggcgaa aatattttct gaacatgttt ttcccccttc 6120
tgctgctata gcgtgtgatt gcgtgatgac atcacgctaa ggtacggtga aagtttcgtt 6180
cactctgttt ctgtgactga tttaagtttg gaagggttgc tgcttttcct gtcgtccagc 6240
aactaaacga atgcctgatg gtttttcaaa tgcatcatgg agccaggagt ggaattggat 6300
ccgcactcaa gaatgtggtt gagttctagc ttctttacct acgttcgagc taaatgttcc 6360
cacttagctt gaactcagca ccttcattgg tagctaggaa catatattga cttttgcaaa 6420
ataaattagt gggattttga tcattacgaa atattgactt ttgcaaaata aagtagcggg 6480
attttgatca tcatgtgctg tatatgtagt gttccgtttt caagttcttc atatttgttt 6540
ttgattctat gtagcactgt agcagagttt ttttgttgtg ctccatacct ttctttagga 6600
agcttctgat cttgcgtatt gacatgcttt tccattttca cctcttcagc catgttaccg 6660
agtaatatgt ctgctggaag tagttcaatt gctcacatga tattctggtg cgggttgcac 6720
gtgacctgct cacatacttc aattgctgga agtagtacat gcattttcag tgtctataac 6780
cttttctgct cgccacaagg gattattgat taattctgtt tactgcccat tggctcatgc 6840
tctttaggct atcacatgca aatgaatgac atataatcat ttgtattagc tatggaaatg 6900
agccagccat cccttataca tgctatgctt ttatgttttc attgattgtt gattccttcc 6960
gtttgatcgt gatcatatac catatggtgc ttgcgtatta gcagttcatg tcttacattt 7020
aggttgtctc gcaaatgcat aaaatgcttt taggccacta caggaaaatg aaagccaccg 7080
tttggaaaat agtcaaatac actttagttc tttcataaat gttgcttact gatgtgctta 7140
tcaatccttt tcttaaacat tgtttactaa tttacagttg attgccgaca tgtgcaaaca 7200
ctttcacttt ttattagccg tgtcgcagac tcgcagtgac ttagtttatt tcttattgca 7260
gaaattctgt gtactctatt actcttacca taggttcatg ctccactaaa actaccttga 7320
tggaacttat tacattttct atttacattg aaactcttcc taattttgtg tgcgcgggtg 7380
tgaacgtgaa gtcctgagca ggtattttta tgctatgcta gtcactgtgt gtgtgcgcgt 7440
gtgttgcggg ctctgttact gttaaccata cataggatta ttctccattg aaactacctt 7500
gatgcaactt attaaaattc caatttacat tgaaacactc tgttttccta attctgtatg 7560
tggtgatgtg aagtgctgag gagttagtta ttcaaatttc attgtacatt gaaacattgt 7620
gtttgtccta attttatctg cgcagatgtg aagttctgag cagttagcca tcttttattt 7680
tttttaaaaa aaattctctg tggttctttt gcctgtttgt ttttacactc tgctaacctc 7740
tgtctgtctg tctgtgttgt atccctccaa atcgtgtctc ctctttgcgc accatttctt 7800
caaatgattt ggattggact aattgtttca attgtgtcat tgtttagtaa gtttttcttg 7860
ctactgctga tgatgatgga ggttaaaagt aacattatca cttccacaat gagttaagga 7920
tgttagaatc tactgtaggt cctgcaattc tgtggatgga ttggcctagt tttcagtgtg 7980
gaacaatccc attccttttt tttcccttat tcagaatatt cattttccat ttttcttatc 8040
aagttttgat agatgtgatt tgtggtctta cagttctgtg ttcctttctt tccagtgccc 8100
atcatggtga agaagaaaag aactgggtct ggcagcaccg gtgagagttc tggagaggct 8160
ccaggagctc ctggccatgg ttcttcacag cgagctgaga gaggtcctca acagcatggg 8220
ggaggacgtg gttgggtgcc tcaacatggt ggccgtggtg gtgggcaata ccagggccgt 8280
ggtggacatt atcagggccg tggagggcaa ggttcacacc atccaggtgg agggcctcct 8340
gagtatcagg gtcgtggagg gccaggttca catcatccag gtggtgggcc tcctgactat 8400
cagggccgtg gaggatcagg ttcacatcac ccaggtggtg ggcctcccga gtatcaaccg 8460
cgtgactatc aaggacgtgg tggtccacgc cccagaggtg gaatgccaca gccatactat 8520
ggcggaccta gggggagtgg cggacgtagt gttccttcag gttcatcaag aacagttccc 8580
gagctgcacc aagccccaca tgtccaatac caagccccga tggtttcacc aaccccatcg 8640
ggagctggct catcctctca gcctgcggcg gaggtgagca gtggacaagt ccaacaacag 8700
tttcagcaac ttgccacccg tgatcaaagt tcgaccagcc aagccattca aatagcacca 8760
ccgtcaagca aatcagttag attcccgttg cgccctggca agggtacata tggggacagg 8820
tgcattgtga aggcgaacca tttctttgct gaacttcctg ataaagacct tcaccaatac 8880
gacgtaaggc ttttgtaagt cctatttcct tgctgtagct ttcattttgt gattttgatc 8940
acctatcttg ttccttcagg tatctattac tcctgaggtt acttcacgtg gcgtgaatcg 9000
tgctgttatg tttgagttag taacgctgta tagatattcc catttgggcg ggcgtctacc 9060
tgcctatgat ggaaggaaga gtctttacac agctggacca ttgccatttg cttctaggac 9120
atttgaaatt actcttcaag atgaggaaga tagtcttggt ggtggccaag gcacccaaag 9180
gtatgctatt gctattttat ctttagttaa atatctatta aaaacttgtt actgacattc 9240
cttctatttt aaggcgtgag agactattta gggtggtgat caagtttgct gcccgtgctg 9300
atcttcacca tttggctatg tttctagctg gaaggcaagc agatgctcct caagaagccc 9360
ttcaagtcct tgacattgtg ttacgtgaat tgcctaccac aaggtaatat ctgatctagc 9420
catctattgt ttattgattt tcttgtgaca atggctttat ttcctttttt ttttaggtac 9480
tcaccagttg gtcggtcatt ttattctccc aatttaggga gacgccagca acttggtgag 9540
ggtttggaaa gttggcgtgg tttttaccaa agcataaggc ctacccagat gggtctctca 9600
ctgaatattg gttagatact gttgcacttc tcctgatttg tcattgtgta tctagatgca 9660
aaaaacattt ttttggtata atcagattca ccattggtgt catctggcgt actgaaattg 9720
cttatttgtt gtttcagata tgtcatcaac tgcatttatt gagcctctac ctgtgattga 9780
ctttgttgct cagcttctga acagagacat ctcagttaga ccattatctg attctgatcg 9840
tgtgaaggtt tggttatatt acctcaccac ctttgttgac aatacctccg tatgtgctta 9900
agaaaatgtt ttttttaacc gtcattgtcc tttttctcac agataaagaa agctctaaga 9960
ggtgtgaagg ttgaggtgac gcatagagga aacatgcgta gaaaatatcg tatatctgga 10020
ctcacttcac aggcaacaag ggagttatcg tatgcacttc ttccctagct tatatgagaa 10080
tctattgcac tcctgcagat gggtatttga aaggattgtg cactgatatg atttggtccg 10140
ttctcctgtg atagattccc tgtcgatgat cgtggtactg tgaagactgt ggtgcaatat 10200
tttctggaga catatggttt tagtattcag cacaccactt tgccttgcct tcaagtgggc 10260
aatcagcaaa ggcccaatta tctgcctatg gaggtcagta tgtttgctgt gctcaattat 10320
agtgatgtat catgctgttt ttgtacgaaa atattttcca aatgctaaat ccagcttcag 10380
catgttatca agtatttacc ttgcctttgg aattgagttc aggtttgtaa gatcgttgag 10440
ggacagcgtt actcgaagcg gcttaacgag aaacagatta ctgcgctatt gaaagtgact 10500
tgccagcgac ctcaagagcg tgaactggat attttgcggg taactgttga tcatattttg 10560
tgatgacatt tgttttgata gtgctgtatt atcggcccca tcttttcact tataaatgca 10620
cttatctgaa ccacttacta ctaactaaaa aataatttat gggtaaaact tgtatatatg 10680
tgttcttagc aattcaaacg caaatgttgt aaaataaact tcgatgagaa agccacaaaa 10740
tcaactccaa aattaagctt taaaattcaa attttggttt ataagcataa gcataagcga 10800
aacgatgggg ctgataatct gatgaatcca tgagttgtat gtttcatgtc cattagcatg 10860
ctgctgtagt taaaacttct aggatgatct ttagcctttt gatttctgct ctctgtactt 10920
tcacatttac tttgtgtgtt tgaagaggaa aatccttggt tgtaggcgat ctctaagacg 10980
cttaattatg ttggtttctt tctttctttc ttcttttttt tttaaaaaaa aatattttgg 11040
ctgttgctag acttctgatg ttacaacaca aagtcgtcct tttttgtata ttttgtcgat 11100
ctaccagaat agtgttatat gttatggtta tgtactatga aaaaacataa atatggtatt 11160
gcttttggtt gtatttattt tctccaagat taaaacagct atattgaggg gttgattctc 11220
atgcattttg ccacctcttt tgttccagct atttgtgagt gtagtggaat ctgtcatgaa 11280
tgtataagag aatatggcaa acttccgatg gagcagtttt tgtttatttt aattatctac 11340
ccttcactga gatactgagt tcagggatct aaatctttgt ttttccttgt tttgatcaga 11400
ctgtatctca caatgcatac catgaagatc agtatgcgca ggaatttggc ataaaaattg 11460
atgagcgtct tgcatctgtt gaagctcgtg ttctgcctcc cccaagggta aatcaatttt 11520
cagatgtggt ttgacagact cacagcagtt gatttccata ttgggcattc gatattcaca 11580
tctattgatt gcttttctat ctctttatta gcttaaatac catgatagtg ggagagaaaa 11640
ggatgtattg ccgagagttg gccagtggaa catgatgaat aaggtacacc tttcaaaagg 11700
agaatcatta tgaaatgtct cttcctctta attcctttgg gcatatccta tgttcatctt 11760
ttatattaag aagggtgaac tgtaccaaaa cagagtcaat attgtacgta ggtatgtgca 11820
aaataaagaa cccaatgttt aatgtatcat taaccagtgg ttttaaaata actgcgaggg 11880
cgcgatatat ggtctagttt ttaagctgta cttctgttca tcacatgatc agtacagtaa 11940
taaaactaat atttatacgg tgtacaaacg tcattctcat gatagaattt cattactgtt 12000
atgaagctcc attctcatgt catgtgtcct acgtacagaa actgttttgg agggatttgg 12060
agtatttaat ttgaggatcc tttataaacc acagagttct ctggcacttc cctccaactt 12120
tcctttgctt ctactcccat cttcactgtg gtagccatag gaccaatatt gtcattttgg 12180
ttaggttact aatcttgata taatctttca cctgtagctg gaacttgctt actgcctctt 12240
ttatgtgtgt aattttatat tgcttgttta catatatgta ttatttattt ggttgtttgt 12300
tttgtagaaa atggtcaatg gtgggagagt caacaactgg gcatgtatta acttctctag 12360
aaatgtgcaa gatagtgctg ccaggggctt ctgtcatgag ctggctatca tgtgccaaat 12420
atctggaatg gtatttacaa gtcatttcag tagcagttca tttttcaggg ttttcttttt 12480
tctattagtt gtttcaacct atgcattttt ttttctttct ataggatttt gcactggaac 12540
ctgtgctgcc cccacttact gctagacctg aacatgtgga aagagcactg aaggcacgct 12600
atcaagatgc aatgaacatg ctcagaccgc agggcaggga acttgattta ctgattgtaa 12660
tactgcctga caataatggt tctctttatg gtatgctctg ttcctaaaga cacttgacca 12720
ttatgcggtg actacctttt cttaacataa ttcttttcat tcctcagggg atctcaaaag 12780
aatctgtgag actgatcttg gattggtctc ccaatgttgt ttgacaaaac atgtttttaa 12840
aatgagcaag cagtatcttg caaatgttgc ccttaaaata aacgttaagg tatgtgttgc 12900
acgccaacta tactttcttg acctttcacc tgaactctat ttctaacttt acattggtcc 12960
tacttttcag gtggggggaa ggaatactgt acttgtggat gctttgacaa ggaggattcc 13020
ccttgtcagt gacagaccaa ctatcatatt tggtgcggat gttactcatc ctcatcctgg 13080
agaagattcc agtccttcca ttgcagctgt aagtgcaatt acgatgaaga ttggccagaa 13140
attctaccaa gttacaatgt aagtttggct agtttgtaac tgttctccct tttaggtggt 13200
tgcttctcaa gactggcctg aagtcactaa gtatgctgga ttggtgagtg cccaagccca 13260
tcgtcaagaa ttgatacaag atcttttcaa agtatggcaa gacccgcata gaggaactgt 13320
tactggtggc atgatcaagt atggacttat tgagatgata catttttact tccctatgtt 13380
tgtacgtcac tgtgcataaa atatgttgaa tgtgcaggga gcttctcatt tctttcaaga 13440
gggctactgg acagaaacct cagaggataa tattttacag gtttttatcc ttgtacagaa 13500
atcttagagg acaacatttt gcaggctttt atccctgtat ggacatcttc ctgaccataa 13560
ttgtatgtga cttcaacacc tgtcatttca gggatggtgt cagcgagggg cagttttatc 13620
aagttttgtt gtatgagctt gatgccatta gaaaggtaca catgttttga cctgaatttg 13680
atcttcaaaa tttttctctt tgatattaac atctactaat ttctggatgc aggcttgtgc 13740
atccctggaa cccaactatc agcctccagt tacctttgtg gtggtccaga agcggcatca 13800
cacaaggttg tttgctaata atcacaacga ccagcgtact gttgatagaa gtggaaacat 13860
tctgcctggt tagttgttga tgcacattca ttttactttg ggcttaggtg atctattctg 13920
actgacattt attgtacctg tttttctttt tgcctaattt ctaggaactg ttgttgactc 13980
aaagatttgc catccaaccg agtttgattt ctacctgtgt agccatgctg gcatacaggt 14040
tggtttaact tgtttgcaat ttcttcactt aatggagtgg tatggatgta tatgattgct 14100
gacttgaatt aattttcttt tctagggaac aagccgtcct gctcattatc atgttctgtg 14160
ggatgagaac aaatttactg cagacgagtt gcaaaccctc acgaacaact tgtgctacac 14220
gtaatttact attccaccag tatggctttt atattcactt tttacaggta tattaaatga 14280
tatttctact gttgtaggta tgcaaggtgc actcgctctg tatcaattgg taagccatct 14340
ttgaaatcac ccccttcggt ttcctggctc ctaaatccag tgcattgtac aactcttgta 14400
aatcactatg ttaacctaca ccacttggtt tcttgcagtg cctcctgcgt actatgctca 14460
tctggcagcc ttccgagctc gcttttacat ggagccagag acatctgaca gtggatcaat 14520
ggcgagtgga gctgcaacga gccgtggcct tccaccaggt gtgcgcagcg ccagggttgc 14580
tggaaatgta gccgtcaggc ctctacctgc tctcaaggaa aacgtgaagc gtgtcatgtt 14640
ttactgctaa 14650
<210> 2
<211> 3357
<212> DNA
<213> 稻属水稻(Oryza sativa)
<400> 2
atggcgctgc agttggagaa tggccgtccc catcatcatc aagtgcccat catggtgaag 60
aagaaaagaa ctgggtctgg cagcaccggt gagagttctg gagaggctcc aggagctcct 120
ggccatggtt cttcacagcg agctgagaga ggtcctcaac agcatggggg aggacgtggt 180
tgggtgcctc aacatggtgg ccgtggtggt gggcaatacc agggccgtgg tggacattat 240
cagggccgtg gagggcaagg ttcacaccat ccaggtggag ggcctcctga gtatcagggt 300
cgtggagggc caggttcaca tcatccaggt ggtgggcctc ctgactatca gggccgtgga 360
ggatcaggtt cacatcaccc aggtggtggg cctcccgagt atcaaccgcg tgactatcaa 420
ggacgtggtg gtccacgccc cagaggtgga atgccacagc catactatgg cggacctagg 480
gggagtggcg gacgtagtgt tccttcaggt tcatcaagaa cagttcccga gctgcaccaa 540
gccccacatg tccaatacca agccccgatg gtttcaccaa ccccatcggg agctggctca 600
tcctctcagc ctgcggcgga ggtgagcagt ggacaagtcc aacaacagtt tcagcaactt 660
gccacccgtg atcaaagttc gaccagccaa gccattcaaa tagcaccacc gtcaagcaaa 720
tcagttagat tcccgttgcg ccctggcaag ggtacatatg gggacaggtg cattgtgaag 780
gcgaaccatt tctttgctga acttcctgat aaagaccttc accaatacga cgtatctatt 840
actcctgagg ttacttcacg tggcgtgaat cgtgctgtta tgtttgagtt agtaacgctg 900
tatagatatt cccatttggg cgggcgtcta cctgcctatg atggaaggaa gagtctttac 960
acagctggac cattgccatt tgcttctagg acatttgaaa ttactcttca agatgaggaa 1020
gatagtcttg gtggtggcca aggcacccaa aggcgtgaga gactatttag ggtggtgatc 1080
aagtttgctg cccgtgctga tcttcaccat ttggctatgt ttctagctgg aaggcaagca 1140
gatgctcctc aagaagccct tcaagtcctt gacattgtgt tacgtgaatt gcctaccaca 1200
aggtactcac cagttggtcg gtcattttat tctcccaatt tagggagacg ccagcaactt 1260
ggtgagggtt tggaaagttg gcgtggtttt taccaaagca taaggcctac ccagatgggt 1320
ctctcactga atattgatat gtcatcaact gcatttattg agcctctacc tgtgattgac 1380
tttgttgctc agcttctgaa cagagacatc tcagttagac cattatctga ttctgatcgt 1440
gtgaagataa agaaagctct aagaggtgtg aaggttgagg tgacgcatag aggaaacatg 1500
cgtagaaaat atcgtatatc tggactcact tcacaggcaa caagggagtt atcattccct 1560
gtcgatgatc gtggtactgt gaagactgtg gtgcaatatt ttctggagac atatggtttt 1620
agtattcagc acaccacttt gccttgcctt caagtgggca atcagcaaag gcccaattat 1680
ctgcctatgg aggtttgtaa gatcgttgag ggacagcgtt actcgaagcg gcttaacgag 1740
aaacagatta ctgcgctatt gaaagtgact tgccagcgac ctcaagagcg tgaactggat 1800
attttgcgga ctgtatctca caatgcatac catgaagatc agtatgcgca ggaatttggc 1860
ataaaaattg atgagcgtct tgcatctgtt gaagctcgtg ttctgcctcc cccaaggctt 1920
aaataccatg atagtgggag agaaaaggat gtattgccga gagttggcca gtggaacatg 1980
atgaataaga aaatggtcaa tggtgggaga gtcaacaact gggcatgtat taacttctct 2040
agaaatgtgc aagatagtgc tgccaggggc ttctgtcatg agctggctat catgtgccaa 2100
atatctggaa tggattttgc actggaacct gtgctgcccc cacttactgc tagacctgaa 2160
catgtggaaa gagcactgaa ggcacgctat caagatgcaa tgaacatgct cagaccgcag 2220
ggcagggaac ttgatttact gattgtaata ctgcctgaca ataatggttc tctttatggg 2280
gatctcaaaa gaatctgtga gactgatctt ggattggtct cccaatgttg tttgacaaaa 2340
catgttttta aaatgagcaa gcagtatctt gcaaatgttg cccttaaaat aaacgttaag 2400
gtggggggaa ggaatactgt acttgtggat gctttgacaa ggaggattcc ccttgtcagt 2460
gacagaccaa ctatcatatt tggtgcggat gttactcatc ctcatcctgg agaagattcc 2520
agtccttcca ttgcagctgt ggttgcttct caagactggc ctgaagtcac taagtatgct 2580
ggattggtga gtgcccaagc ccatcgtcaa gaattgatac aagatctttt caaagtatgg 2640
caagacccgc atagaggaac tgttactggt ggcatgatca aggagcttct catttctttc 2700
aagagggcta ctggacagaa acctcagagg ataatatttt acagggatgg tgtcagcgag 2760
gggcagtttt atcaagtttt gttgtatgag cttgatgcca ttagaaaggc ttgtgcatcc 2820
ctggaaccca actatcagcc tccagttacc tttgtggtgg tccagaagcg gcatcacaca 2880
aggttgtttg ctaataatca caacgaccag cgtactgttg atagaagtgg aaacattctg 2940
cctggaactg ttgttgactc aaagatttgc catccaaccg agtttgattt ctacctgtgt 3000
agccatgctg gcatacaggg aacaagccgt cctgctcatt atcatgttct gtgggatgag 3060
aacaaattta ctgcagacga gttgcaaacc ctcacgaaca acttgtgcta cacgtatgca 3120
aggtgcactc gctctgtatc aattgtgcct cctgcgtact atgctcatct ggcagccttc 3180
cgagctcgct tttacatgga gccagagaca tctgacagtg gatcaatggc gagtggagct 3240
gcaacgagcc gtggccttcc accaggtgtg cgcagcgcca gggttgctgg aaatgtagcc 3300
gtcaggcctc tacctgctct caaggaaaac gtgaagcgtg tcatgtttta ctgctaa 3357
<210> 3
<211> 1118
<212> PRT
<213> 稻属水稻(Oryza sativa)
<400> 3
Met Ala Leu Gln Leu Glu Asn Gly Arg Pro His His His Gln Val Pro
1 5 10 15
Ile Met Val Lys Lys Lys Arg Thr Gly Ser Gly Ser Thr Gly Glu Ser
20 25 30
Ser Gly Glu Ala Pro Gly Ala Pro Gly His Gly Ser Ser Gln Arg Ala
35 40 45
Glu Arg Gly Pro Gln Gln His Gly Gly Gly Arg Gly Trp Val Pro Gln
50 55 60
His Gly Gly Arg Gly Gly Gly Gln Tyr Gln Gly Arg Gly Gly His Tyr
65 70 75 80
Gln Gly Arg Gly Gly Gln Gly Ser His His Pro Gly Gly Gly Pro Pro
85 90 95
Glu Tyr Gln Gly Arg Gly Gly Pro Gly Ser His His Pro Gly Gly Gly
100 105 110
Pro Pro Asp Tyr Gln Gly Arg Gly Gly Ser Gly Ser His His Pro Gly
115 120 125
Gly Gly Pro Pro Glu Tyr Gln Pro Arg Asp Tyr Gln Gly Arg Gly Gly
130 135 140
Pro Arg Pro Arg Gly Gly Met Pro Gln Pro Tyr Tyr Gly Gly Pro Arg
145 150 155 160
Gly Ser Gly Gly Arg Ser Val Pro Ser Gly Ser Ser Arg Thr Val Pro
165 170 175
Glu Leu His Gln Ala Pro His Val Gln Tyr Gln Ala Pro Met Val Ser
180 185 190
Pro Thr Pro Ser Gly Ala Gly Ser Ser Ser Gln Pro Ala Ala Glu Val
195 200 205
Ser Ser Gly Gln Val Gln Gln Gln Phe Gln Gln Leu Ala Thr Arg Asp
210 215 220
Gln Ser Ser Thr Ser Gln Ala Ile Gln Ile Ala Pro Pro Ser Ser Lys
225 230 235 240
Ser Val Arg Phe Pro Leu Arg Pro Gly Lys Gly Thr Tyr Gly Asp Arg
245 250 255
Cys Ile Val Lys Ala Asn His Phe Phe Ala Glu Leu Pro Asp Lys Asp
260 265 270
Leu His Gln Tyr Asp Val Ser Ile Thr Pro Glu Val Thr Ser Arg Gly
275 280 285
Val Asn Arg Ala Val Met Phe Glu Leu Val Thr Leu Tyr Arg Tyr Ser
290 295 300
His Leu Gly Gly Arg Leu Pro Ala Tyr Asp Gly Arg Lys Ser Leu Tyr
305 310 315 320
Thr Ala Gly Pro Leu Pro Phe Ala Ser Arg Thr Phe Glu Ile Thr Leu
325 330 335
Gln Asp Glu Glu Asp Ser Leu Gly Gly Gly Gln Gly Thr Gln Arg Arg
340 345 350
Glu Arg Leu Phe Arg Val Val Ile Lys Phe Ala Ala Arg Ala Asp Leu
355 360 365
His His Leu Ala Met Phe Leu Ala Gly Arg Gln Ala Asp Ala Pro Gln
370 375 380
Glu Ala Leu Gln Val Leu Asp Ile Val Leu Arg Glu Leu Pro Thr Thr
385 390 395 400
Arg Tyr Ser Pro Val Gly Arg Ser Phe Tyr Ser Pro Asn Leu Gly Arg
405 410 415
Arg Gln Gln Leu Gly Glu Gly Leu Glu Ser Trp Arg Gly Phe Tyr Gln
420 425 430
Ser Ile Arg Pro Thr Gln Met Gly Leu Ser Leu Asn Ile Asp Met Ser
435 440 445
Ser Thr Ala Phe Ile Glu Pro Leu Pro Val Ile Asp Phe Val Ala Gln
450 455 460
Leu Leu Asn Arg Asp Ile Ser Val Arg Pro Leu Ser Asp Ser Asp Arg
465 470 475 480
Val Lys Ile Lys Lys Ala Leu Arg Gly Val Lys Val Glu Val Thr His
485 490 495
Arg Gly Asn Met Arg Arg Lys Tyr Arg Ile Ser Gly Leu Thr Ser Gln
500 505 510
Ala Thr Arg Glu Leu Ser Phe Pro Val Asp Asp Arg Gly Thr Val Lys
515 520 525
Thr Val Val Gln Tyr Phe Leu Glu Thr Tyr Gly Phe Ser Ile Gln His
530 535 540
Thr Thr Leu Pro Cys Leu Gln Val Gly Asn Gln Gln Arg Pro Asn Tyr
545 550 555 560
Leu Pro Met Glu Val Cys Lys Ile Val Glu Gly Gln Arg Tyr Ser Lys
565 570 575
Arg Leu Asn Glu Lys Gln Ile Thr Ala Leu Leu Lys Val Thr Cys Gln
580 585 590
Arg Pro Gln Glu Arg Glu Leu Asp Ile Leu Arg Thr Val Ser His Asn
595 600 605
Ala Tyr His Glu Asp Gln Tyr Ala Gln Glu Phe Gly Ile Lys Ile Asp
610 615 620
Glu Arg Leu Ala Ser Val Glu Ala Arg Val Leu Pro Pro Pro Arg Leu
625 630 635 640
Lys Tyr His Asp Ser Gly Arg Glu Lys Asp Val Leu Pro Arg Val Gly
645 650 655
Gln Trp Asn Met Met Asn Lys Lys Met Val Asn Gly Gly Arg Val Asn
660 665 670
Asn Trp Ala Cys Ile Asn Phe Ser Arg Asn Val Gln Asp Ser Ala Ala
675 680 685
Arg Gly Phe Cys His Glu Leu Ala Ile Met Cys Gln Ile Ser Gly Met
690 695 700
Asp Phe Ala Leu Glu Pro Val Leu Pro Pro Leu Thr Ala Arg Pro Glu
705 710 715 720
His Val Glu Arg Ala Leu Lys Ala Arg Tyr Gln Asp Ala Met Asn Met
725 730 735
Leu Arg Pro Gln Gly Arg Glu Leu Asp Leu Leu Ile Val Ile Leu Pro
740 745 750
Asp Asn Asn Gly Ser Leu Tyr Gly Asp Leu Lys Arg Ile Cys Glu Thr
755 760 765
Asp Leu Gly Leu Val Ser Gln Cys Cys Leu Thr Lys His Val Phe Lys
770 775 780
Met Ser Lys Gln Tyr Leu Ala Asn Val Ala Leu Lys Ile Asn Val Lys
785 790 795 800
Val Gly Gly Arg Asn Thr Val Leu Val Asp Ala Leu Thr Arg Arg Ile
805 810 815
Pro Leu Val Ser Asp Arg Pro Thr Ile Ile Phe Gly Ala Asp Val Thr
820 825 830
His Pro His Pro Gly Glu Asp Ser Ser Pro Ser Ile Ala Ala Val Val
835 840 845
Ala Ser Gln Asp Trp Pro Glu Val Thr Lys Tyr Ala Gly Leu Val Ser
850 855 860
Ala Gln Ala His Arg Gln Glu Leu Ile Gln Asp Leu Phe Lys Val Trp
865 870 875 880
Gln Asp Pro His Arg Gly Thr Val Thr Gly Gly Met Ile Lys Glu Leu
885 890 895
Leu Ile Ser Phe Lys Arg Ala Thr Gly Gln Lys Pro Gln Arg Ile Ile
900 905 910
Phe Tyr Arg Asp Gly Val Ser Glu Gly Gln Phe Tyr Gln Val Leu Leu
915 920 925
Tyr Glu Leu Asp Ala Ile Arg Lys Ala Cys Ala Ser Leu Glu Pro Asn
930 935 940
Tyr Gln Pro Pro Val Thr Phe Val Val Val Gln Lys Arg His His Thr
945 950 955 960
Arg Leu Phe Ala Asn Asn His Asn Asp Gln Arg Thr Val Asp Arg Ser
965 970 975
Gly Asn Ile Leu Pro Gly Thr Val Val Asp Ser Lys Ile Cys His Pro
980 985 990
Thr Glu Phe Asp Phe Tyr Leu Cys Ser His Ala Gly Ile Gln Gly Thr
995 1000 1005
Ser Arg Pro Ala His Tyr His Val Leu Trp Asp Glu Asn Lys Phe Thr
1010 1015 1020
Ala Asp Glu Leu Gln Thr Leu Thr Asn Asn Leu Cys Tyr Thr Tyr Ala
1025 1030 1035 1040
Arg Cys Thr Arg Ser Val Ser Ile Val Pro Pro Ala Tyr Tyr Ala His
1045 1050 1055
Leu Ala Ala Phe Arg Ala Arg Phe Tyr Met Glu Pro Glu Thr Ser Asp
1060 1065 1070
Ser Gly Ser Met Ala Ser Gly Ala Ala Thr Ser Arg Gly Leu Pro Pro
1075 1080 1085
Gly Val Arg Ser Ala Arg Val Ala Gly Asn Val Ala Val Arg Pro Leu
1090 1095 1100
Pro Ala Leu Lys Glu Asn Val Lys Arg Val Met Phe Tyr Cys
1105 1110 1115
Claims (7)
1.一种水稻基因GSNL4的应用，其特征在于，所述基因用于调控水稻的粒型和叶型，所述基因如(a)、(b)或(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)或(b)所示的核苷酸序列中添加和/或取代和/或缺失一个或几个核苷酸而生成的可编码调控水稻粒型和叶型的蛋白质的突变基因、等位基因或衍生物。
2.根据权利要求1所述的水稻基因GSNL4的应用，其特征在于，利用所述基因转化水稻细胞，再将转化后的水稻细胞培育成植株。
3.根据权利要求1或2所述的应用，其特征在于，所述粒型为粒宽和粒长，所述叶型为叶宽。
4.根据权利要求1或2所述的水稻基因GSNL4的应用，其特征在于，所述基因用于提高水稻粒重及高产育种。
5.一种水稻基因GSNL4编码的蛋白质的应用，其特征在于，所述蛋白质用于调控水稻的粒型和叶型；所述蛋白质如(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。
6.根据权利要求5所述的水稻基因GSNL4编码的蛋白质的应用，其特征在于，所述粒型为粒宽和粒长，所述叶型为叶宽。
7.根据权利要求5或6所述的水稻基因GSNL4编码的蛋白质的应用，其特征在于，所述蛋白质用于提高水稻粒重及高产育种。
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