CN113862279B - 抑制水稻幼苗生长的基因OsACO及其应用 - Google Patents
抑制水稻幼苗生长的基因OsACO及其应用 Download PDFInfo
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Abstract
本申请公开了一种抑制水稻幼苗生长发育的基因OsACO及其应用，所述基因OsACO编码的蛋白序列如SEQ ID NO.3所示。所述应用包括下调所述水稻中基因OsACO的表达。
Description
技术领域
本申请涉及水稻育种领域，尤其涉及一种抑制水稻幼苗生长的基因OsACO及其应用。
背景技术
水稻是我国重要的粮食作物之一，约一半的人口以水稻为主食(Yong etal.2014)。中国以全球7％的耕地面积，却养活了全球20％的人口，故水稻一直是植物分子生物学研究领域的热点(Xie et al.1997)。
植物激素是影响植物生长和发育的重要因素。植物激素是植物体产生的、极其微量并对植物生长发育起至关重要作用的信号分子，包括：吲哚乙酸(IAA)、脱落酸(ABA)、赤霉素(GA)、细胞分裂素(CTK)、乙烯(ethylene，ETH)和油菜素甾醇(brassinosteroid，BR)等。其中，乙烯作为一种重要的植物内源激素，对幼苗建成具有非常重要的生理作用。例如，在乙烯的处理下，能够抑制水稻的幼苗生长，导致植物矮化(Biao et al.2009)。相反，当乙烯信号通路的正调控因子发生突变时，促进植物的生长。例如，EIN2是乙烯信号转导的中心成分，ein2突变体通过增强细胞的扩张，促进植物的生长(Feng et al.2015)。除此之外，乙烯对植物的叶片衰老(Li et al.2018)、果实成熟(Barry et al.2007)和非生物胁迫(Jiang et al.2013；Yang et al.2015)中都起着至关重要的作用。
研究发现，水稻生长周期长，会大大增加农药和化肥的使用，提高了成本并且对坏境和土壤造成了不可修复的伤害(Wang et al.2018；Gao et al.2002)。所以提高水稻的生长速率，缩短生长周期，对发展双季稻和提高水稻产量都具有重要的作用(Cheng etal.2015)。因此，本领域的技术人员致力于开发一种促进水稻幼苗生长的基因及其应用。
发明内容
有鉴于现有技术的上述缺陷，本申请所要解决的技术问题是如何促进水稻的幼苗生长。
为实现上述目的，本申请提供了一种抑制水稻幼苗生长发育的基因OsACO，其特征在于，所述基因OsACO编码的蛋白序列如SEQ ID NO.3所示。
在某些实施方式中，所述基因OsACO的编码序列如SEQ ID NO.2所示。
在某些实施方式中，所述基因OsACO的序列如SEQ ID NO.1所示。
另一方面，本申请还提够了基因OsACO在促进水稻幼苗生长中的应用，其特征在于，包括下调所述水稻中基因OsACO的表达，所述基因OsACO编码的蛋白序列如SEQ ID NO.3所示。
在某些实施方式中，所述基因OsACO的编码序列如SEQ ID NO.2所示。
在某些实施方式中，所述基因OsACO的序列如SEQ ID NO.1所示。
在某些实施方式中，所述下调水稻基因OsACO的表达包基因敲除、基因突变。
在某些实施方式中，基因突变包括碱基置换突变、移码突变、缺失突变和/或插入突变。
在某些实施方式中，所述碱基置换突变包括位于所述基因OsACO的编码序列335位和/或345位的碱基由T突变成C。
另一方面，本申请还提供了一种水稻育种的方法，其特征在于，包括：提取总RNA，利用如SEQ ID NO.6和7所示的引物进行定量PCR反应，比较基因OsACO的表达量，选出其中基因OsACO表达量低的个体。
本申请中通过对OsACO基因的研究发现，OsACO基因发生突变之后，促进水稻幼苗的生长，证明了OsACO基因抑制水稻幼苗的生长，解决了水稻生长周期长的问题，为发展双季稻奠定了基础，为后续水稻高产、优产提供了新途径。
以下将结合附图对本申请的构思、具体结构及产生的技术效果作进一步说明，以充分地了解本申请的目的、特征和效果。
附图说明
图1显示的是本申请中不同激素处理下水稻乙烯合成基因OsACO的表达模式分析结果图；
图2显示的是本申请中水稻乙烯合成基因OsACO在水稻不同部位的表达谱分析结果图；
图3显示的是本申请中osaco突变体和野生型水稻的cDNA序列比对图；
图4显示的是本申请中osaco突变体和野生型水稻中乙烯合成基因OsACO表达变化分析结果图；
图5显示的是本申请中水稻osaco突变体和野生型幼苗表型对比图；
图6显示的是本申请中水稻乙烯响应基因OsERF2表达分析结果图；
图7显示的是本申请中水稻乙烯响应基因OsETR2表达分析结果图。
具体实施方式
以下将结合实施例对本申请作进一步地说明，应理解这些实施例仅作为例证的目的，本申请可以通过许多不同形式的实施例来得以体现，本申请的保护范围并非仅限于文中提到的实施例。
本申请实施例中出现的野生型(WT)及突变体水稻为中花11号，是京风五号/特特普♀/福锦♂花培获得的粳稻，株高110-115厘米左右，株型较紧凑，较繁茂，叶色浓绿。穗大、码密，穗颈长，有弯腰现象。穗长20厘米，大穗型品种。平均穗粒数115-120粒，空秕粒率为15-20％，较高。谷粒椭圆形，颖及颖尖杆黄色，千短顶芒，粒重26-27克。米质优，垩白少，透明度好，食味佳。全生育期160天左右，属中熟品种。分蘖力强，耐肥，抗倒伏性和抗病性较强，抗寒，抗盐碱。秧令弹性较大。
下列实例中未注明具体的实验方法，均可按照常规方法进行。如Sambrook等分子克隆：实验手册(New York:Cold Spring Harbor Laboratory Press,1989)中所述条件，或按照制造生产厂商的使用说明；本申请中的野生型及突变体水稻种子均可自上海师范大学获取。
实施例1水稻基因OsACO的获得
1.水稻品种中花11在培养箱(SPX-250-GB,Shanghai,China)中培养：生长条件为光周期16h/8h(L/D)，28℃。
1.DNA提取：取500mg左右新鲜的水稻植物组织材料，加入80μLSDS-提取缓冲溶液Lysis Buffer，用研磨棒将植物组织研碎，加入120μL ddH2O。12000rpm离心15min后将上清转移至新的离心管，测OD值，电泳检测。
3.基因的克隆。以提取的水稻DNA为模板，以如SEQ ID NO.4和5所示的序列作为引物，通过PCR反应获得基因全长，全长序列信息如SEQ ID NO.1所示，编码序列如SEQ IDNO.2所示，编码的氨基酸序列如SEQ ID NO.3所示。
实施例2在不同激素处理下水稻乙烯合成基因OsACO的表达模式分析
1.野生型的种子放入在30℃培养箱中培养24h；
2.无菌水中漂洗后，放于37℃的培养箱培养24h；
3.将种子放入28℃培养箱中培养两周；
4.使用不同激素处理，黑暗条件下28℃培养72h；
5.提取水稻两周幼苗总RNA，利用反转录试剂盒将以上得到的总RNA反转录成cDNA，利用如SEQ ID NO.6和SEQ ID NO.7所示的引物序列进行Real-time PCR检测。结果如图1所示，野生型水稻在赤霉素(GA)、吲哚乙酸(IAA)、脱落酸(ABA)激素作用下，OsACO表达量显著下调；而在乙烯(ET)处理下，OsACO表达量显著上调，CK为未作处理的对照组。
实施例3乙烯合成基因在水稻不同部位的表达谱分析
1.野生型的种子放入在30℃培养箱中培养24h；
2.无菌水中漂洗后，放于37℃的培养箱培养24h；
3.将种子放入28℃培养箱中培养两周；
4.提取水稻幼苗根、茎、叶总的RNA，利用反转录试剂盒将以上得到的总RNA反转录成cDNA，利用如SEQ ID NO.6和SEQ ID NO.7所示的引物序列进行Real-time PCR检测。结果如图2所示，野生型水稻的茎和叶中，OsACO表达量显著高于根。
实施例4突变体osaco的分子鉴定
(1)中花11号的突变体株系与野生型的种子浸泡在水中24h，让种子充分吸涨，随后将种子玻璃培养皿内，放在水稻恒温培养箱内37℃催芽至露白，每天换水，防止长霉，选取萌发一致的种子，将其转移至种子架上，置于基础营养液中，28℃培养箱培养。
(2)分别提取突变体osaco以及野生型对照组中的总DNA，利用如SEQ ID NO.8和SEQ ID NO.9所示的引物进行PCR检测。结果显示，突变体osaco位于编码区335位和345位碱基由T突变成C，如图3所示。筛选出纯合突变体，并进行繁殖。
实施例5野生型和突变体水稻中乙烯合成基因的表达模式分析
1.突变体与野生型的种子放入在30℃培养箱中培养24h；
2.无菌水中漂洗后，放于37℃的培养箱培养24h；
3.将种子放入28℃培养箱中培养两周；
4.提取水稻幼苗总RNA，利用反转录试剂盒将以上得到的总RNA反转录成cDNA，利用如SEQ ID NO.6和SEQ ID NO.7所示的引物序列进行Real-time PCR检测。结果如图4所示，OsACO基因在osaco突变体中的表达显著降低，本领域已知OsACO基因能够促进乙烯的合成，该结果表明会影响乙烯的合成。
实施例6OsACO基因对幼苗生长发育的影响
(1)从种子中各挑选50粒饱满种子作为实验材料，突变体与野生型的种子放入在30℃培养箱中培养24h，于无菌水中漂洗后，放于37℃的培养箱培养培养24h，最后，将种子放入28℃培养箱中培养两周；
(2)osaco突变体与野生型水稻幼苗分别在茎和根进行观察，结果如图5所示，图A为株高表型图，图B为株高数据结果统计图。与野生型(图A及图B中ZH11组)相比，osaco突变体(图A和图B中osaco组)幼苗的株高和根都显著增长，即OsACO基因负调控水稻幼苗的发育。本实施例中的osaco突变体以野生型中花11号为背景进行突变，因此在具有相同的遗传背景下，该结果证明了促进幼苗的生长是由OsACO基因突变直接导致的。
实施例7水稻乙烯响应基因的表达模式分析
1.突变体与野生型的种子放入在30℃培养箱中培养24h；
2.无菌水中漂洗后，放于37℃的培养箱培养24h；
3.将种子放入28℃培养箱中培养两周；
4.提取水稻幼苗总RNA，利用反转录试剂盒将以上得到的总RNA反转录成cDNA，利用引物SEQ ID NO.10、SEQ ID NO.11、SEQ ID NO.12、SEQ ID NO.13，进行Real-time PCR检测。结果显示，OsERF2和OsETR2在osaco突变体中的表达都显著降低，如图6和7所示。osaco突变体中的OsACO基因突变会下调乙烯的信号响应，表明了OsACO基因突变通过下调对乙烯信号的响应从而促进了幼苗的生长发育。
以上详细描述了本申请的较佳具体实施例。应当理解，本领域的普通技术无需创造性劳动就可以根据本申请的构思作出诸多修改和变化。因此，凡本技术领域中技术人员依本申请的构思在现有技术的基础上通过逻辑分析、推理或者有限的实验可以得到的技术方案，皆应在由权利要求书所确定的保护范围内。
序列表
<110> 上海师范大学
<120> 抑制水稻幼苗生长的基因OsACO及其应用
<130> CN015-20017PICN
<160> 13
<170> PatentIn version 3.5
<210> 1
<211> 1391
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 基因OsACO全长序列
<400> 1
cttttaatca tctactccca ccctaattta atttagacgg acctagttaa tggccatcct 60
ggagagagca gaggaagcca acatcggcga aggctcaggc tcgtcggagt gggagctggg 120
cgtccggcag ctgtgcgaca gcggcatcac caccctgccc gcccgctacg tcctcccgcc 180
cgccgaccgc cccgcccgct acgtcacacc acctgctctg ctccccgtcg tcgaccttgc 240
cgccctccgt gctcgagacc cctgtcagct cgccgcgctc cacgccgcct gccgggacta 300
cggcttcttc cagcttctca accacggcgt gccccccgac gccatgctgt acgccgctcg 360
ccgcttcttc ttcgaccttc ccctgcccgc ccgtaagcgc tacatgtccg ccgacatccg 420
cgccgccgtc cgctacggca ccagcttcaa ccagctcaac gacgccgtcc tctcctggcg 480
cgacttcctc aagctactca tccgcgacac gcgccgcctc gccgacgtcc tcccctcctg 540
gcccgacgcc cccgacgacc tcaggccggc cgccgcggcg tacgccacgg cgtgccagag 600
gctgttccgg gagctcatgg aggcggcgct ggatgcgctg ggcatcgtgc ggtgccgccg 660
ccagctgctg gaggagtgcg acgccgggtc gcagatgatg atggtcaact gcttcccggc 720
gtgcccggag ccggagctga cgctgggcat gccgccgcac tccgactacg gcctgctaac 780
catcctcctg caggacgagg tgagagggct ggaggtgagc tacggcgacg gcggcgggtg 840
ggcggtggtg gagccgcttc ccggcgcggt ggtggtgaac gtgggcgacc acctggagat 900
actgagcaac gggctgtacc ggagcgtgct gcaccgtgtg cgcgtgaacg gtcggcgggc 960
gcgcgtgtcg gtggcgtcgc tgcacagcct ggcggcggag cgtgtgatcg ggccggcggc 1020
ggagctggtg gacgagcagc ggggcaggcc gcggcggtac atggacaccg acatggccgc 1080
gttccttgcc tacctcgcct ccgcagaggg caaccacaag tccttcctcc actcccgcag 1140
gatcaacacc atttcttctt ccggactgac ccaacccagc aactaaatta attagctgga 1200
cgtacggact ctctatctaa ttactagtcg ccaaattaag agtacgtcgt cccttctagc 1260
tagttcctat acaaacatat ccaccctggt tgcaactgta gtgttaagct agcacggaat 1320
ccatcactgt acaagcatgt gtgtacgtgc agtgatttaa ttaattaatt tgctgaaaaa 1380
taatccaaat t 1391
<210> 2
<211> 1137
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 基因OsACO编码序列
<400> 2
atggccatcc tggagagagc agaggaagcc aacatcggcg aaggctcagg ctcgtcggag 60
tgggagctgg gcgtccggca gctgtgcgac agcggcatca ccaccctgcc cgcccgctac 120
gtcctcccgc ccgccgaccg ccccgcccgc tacgtcacac cacctgctct gctccccgtc 180
gtcgaccttg ccgccctccg tgctcgagac ccctgtcagc tcgccgcgct ccacgccgcc 240
tgccgggact acggcttctt ccagcttctc aaccacggcg tgccccccga cgccatgctg 300
tacgccgctc gccgcttctt cttcgacctt cccctgcccg cccgtaagcg ctacatgtcc 360
gccgacatcc gcgccgccgt ccgctacggc accagcttca accagctcaa cgacgccgtc 420
ctctcctggc gcgacttcct caagctactc atccgcgaca cgcgccgcct cgccgacgtc 480
ctcccctcct ggcccgacgc ccccgacgac ctcaggccgg ccgccgcggc gtacgccacg 540
gcgtgccaga ggctgttccg ggagctcatg gaggcggcgc tggatgcgct gggcatcgtg 600
cggtgccgcc gccagctgct ggaggagtgc gacgccgggt cgcagatgat gatggtcaac 660
tgcttcccgg cgtgcccgga gccggagctg acgctgggca tgccgccgca ctccgactac 720
ggcctgctaa ccatcctcct gcaggacgag gtgagagggc tggaggtgag ctacggcgac 780
ggcggcgggt gggcggtggt ggagccgctt cccggcgcgg tggtggtgaa cgtgggcgac 840
cacctggaga tactgagcaa cgggctgtac cggagcgtgc tgcaccgtgt gcgcgtgaac 900
ggtcggcggg cgcgcgtgtc ggtggcgtcg ctgcacagcc tggcggcgga gcgtgtgatc 960
gggccggcgg cggagctggt ggacgagcag cggggcaggc cgcggcggta catggacacc 1020
gacatggccg cgttccttgc ctacctcgcc tccgcagagg gcaaccacaa gtccttcctc 1080
cactcccgca ggatcaacac catttcttct tccggactga cccaacccag caactaa 1137
<210> 3
<211> 378
<212> PRT
<213> Artificial Sequence（人工序列）
<220>
<223> OsACO编码氨基酸序列
<400> 3
Met Ala Ile Leu Glu Arg Ala Glu Glu Ala Asn Ile Gly Glu Gly Ser
1 5 10 15
Gly Ser Ser Glu Trp Glu Leu Gly Val Arg Gln Leu Cys Asp Ser Gly
20 25 30
Ile Thr Thr Leu Pro Ala Arg Tyr Val Leu Pro Pro Ala Asp Arg Pro
35 40 45
Ala Arg Tyr Val Thr Pro Pro Ala Leu Leu Pro Val Val Asp Leu Ala
50 55 60
Ala Leu Arg Ala Arg Asp Pro Cys Gln Leu Ala Ala Leu His Ala Ala
65 70 75 80
Cys Arg Asp Tyr Gly Phe Phe Gln Leu Leu Asn His Gly Val Pro Pro
85 90 95
Asp Ala Met Leu Tyr Ala Ala Arg Arg Phe Phe Phe Asp Leu Pro Leu
100 105 110
Pro Ala Arg Lys Arg Tyr Met Ser Ala Asp Ile Arg Ala Ala Val Arg
115 120 125
Tyr Gly Thr Ser Phe Asn Gln Leu Asn Asp Ala Val Leu Ser Trp Arg
130 135 140
Asp Phe Leu Lys Leu Leu Ile Arg Asp Thr Arg Arg Leu Ala Asp Val
145 150 155 160
Leu Pro Ser Trp Pro Asp Ala Pro Asp Asp Leu Arg Pro Ala Ala Ala
165 170 175
Ala Tyr Ala Thr Ala Cys Gln Arg Leu Phe Arg Glu Leu Met Glu Ala
180 185 190
Ala Leu Asp Ala Leu Gly Ile Val Arg Cys Arg Arg Gln Leu Leu Glu
195 200 205
Glu Cys Asp Ala Gly Ser Gln Met Met Met Val Asn Cys Phe Pro Ala
210 215 220
Cys Pro Glu Pro Glu Leu Thr Leu Gly Met Pro Pro His Ser Asp Tyr
225 230 235 240
Gly Leu Leu Thr Ile Leu Leu Gln Asp Glu Val Arg Gly Leu Glu Val
245 250 255
Ser Tyr Gly Asp Gly Gly Gly Trp Ala Val Val Glu Pro Leu Pro Gly
260 265 270
Ala Val Val Val Asn Val Gly Asp His Leu Glu Ile Leu Ser Asn Gly
275 280 285
Leu Tyr Arg Ser Val Leu His Arg Val Arg Val Asn Gly Arg Arg Ala
290 295 300
Arg Val Ser Val Ala Ser Leu His Ser Leu Ala Ala Glu Arg Val Ile
305 310 315 320
Gly Pro Ala Ala Glu Leu Val Asp Glu Gln Arg Gly Arg Pro Arg Arg
325 330 335
Tyr Met Asp Thr Asp Met Ala Ala Phe Leu Ala Tyr Leu Ala Ser Ala
340 345 350
Glu Gly Asn His Lys Ser Phe Leu His Ser Arg Arg Ile Asn Thr Ile
355 360 365
Ser Ser Ser Gly Leu Thr Gln Pro Ser Asn
370 375
<210> 4
<211> 25
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 正向引物1
<400> 4
atggccatcc tggagagagc agagg 25
<210> 5
<211> 25
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 反向引物1
<400> 5
ttagttgctg ggttgggtca gtccg 25
<210> 6
<211> 18
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 正向引物2
<400> 6
cgatccacca gcaagagc 18
<210> 7
<211> 18
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 反向引物2
<400> 7
gaactgcctc gggatcag 18
<210> 8
<211> 31
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 正向引物3
<400> 8
gaccttcccc cgcccgcccg caagcgctac a 31
<210> 9
<211> 31
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 反向引物3
<400> 9
tgtagcgctt gcgggcgggc gggggaaggt c 31
<210> 10
<211> 22
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 正向引物4
<400> 10
ccctgttcat ctacctgttc tt 22
<210> 11
<211> 22
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 反向引物4
<400> 11
caactgcaac cctattatgc tc 22
<210> 12
<211> 22
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 正向引物5
<400> 12
cattcaacct tcattcgttg gt 22
<210> 13
<211> 22
<212> DNA
<213> Artificial Sequence（人工序列）
<220>
<223> 反向引物5
<400> 13
aactcaaaat caatcccctt gc 22
Claims (1)
1.基因OsACO在促进水稻幼苗生长中的应用，其特征在于，包括下调所述水稻中基因OsACO的表达，所述基因OsACO编码的蛋白序列如SEQ ID NO. 3所示；所述下调水稻基因OsACO的表达为基因突变，所述基因突变为碱基置换突变，所述碱基置换突变为位于所述基因OsACO的编码序列335位和345位的碱基由T突变成C；所述基因OsACO的编码序列如SEQ IDNO. 2所示。
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