CN115197920B - 调控细胞分裂素氧化酶基因TaCKX5的物质在提高小麦产量中的应用 - Google Patents
调控细胞分裂素氧化酶基因TaCKX5的物质在提高小麦产量中的应用 Download PDFInfo
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- CN115197920B CN115197920B CN202110377264.9A CN202110377264A CN115197920B CN 115197920 B CN115197920 B CN 115197920B CN 202110377264 A CN202110377264 A CN 202110377264A CN 115197920 B CN115197920 B CN 115197920B
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Abstract
本发明公开了TaCKX5基因在调控小麦产量中的应用，所述基因为细胞分裂素氧化酶基因TaCKX5，编码TaCKX5蛋白。本发明采用CRISPR/Cas9技术敲除了小麦TaCKX5基因，获得了低磷条件下和正常氮磷条件下产量明显提高的小麦新种质。本发明提高了小麦产量，创新了小麦种质资源，对新品种培育、环境保护和粮食安全具有重要意义，具有重大的应用推广价值。
Description
技术领域
本发明涉及生物技术领域中调控细胞分裂素氧化酶基因TaCKX5的物质在提高小麦产量中的应用。
背景技术
小麦是主要粮食作物，提高小麦产量对于保障粮食安全具有极其重要的作用。磷肥是小麦获得高产所需的大量元素肥料之一。生产磷肥的磷矿资源为不可再生资源，因此如能提高小麦在少施磷肥下的产量对于促进农业可持续发展具有重要意义。
细胞分裂素是一类重要的植物激素，通过控制细胞分裂、诱导促进芽萌发生长等方面控制植物发育。在植物体内细胞分裂素含量受到合成和降解通路基因的调控，其中细胞分裂素氧化酶(CKX)是降解细胞分裂素的关键酶，通过调控细胞分裂素氧化酶活性可以改变植物体内细胞分裂素含量，进而影响植物的生长发育。自1971年在烟草中首次发现CKX以来，相继在拟南芥、玉米、小麦、水稻等植物中被发现。植物生长不同阶段和不同部位CKX基因的表达模式存在较大差异，因此CKX基因作为下调植物内源细胞分裂素水平的重要基因，在定向改良作物性状、培育优良品种方面具有发掘的研究潜力。在拟南芥中敲除CKX3和CKX5显著增加了籽粒产量(Bartrina et al.2011)。水稻中已知CKX基因家族共有11个成员，日本的科学家发现OsCKX2基因的一个突变体导致细胞分裂素氧化酶降低，增加了水稻穗中细胞分裂素的含量和穗粒数，是在水稻中控制穗粒数的主效基因(Ashikari etal.2005)。利用CRISPR-Cas9技术敲除OsCKX2亦显著增加了水稻的穗粒数(Li etal.2016)。最近浙江大学的研究者利用CRISPR-Cas9基因编辑技术创制了水稻11个CKX成员的突变体，发现其中osckx11突变体不仅表现出叶片适度延缓衰老表型，而且分蘖数和穗粒数均显著增加。体外重组OsCKX11蛋白催化多种类型细胞分裂素的降解，但对反式玉米素(trans-zeatin)和顺式玉米素(cis-zeatin)表现出较强的偏好性。与野生型(WT)相比，osckx11突变体旗叶的细胞分裂素水平显著升高(Zhang et al.,2020)。但是由于组织表达特异性的不同，或者是因为基因冗余，并不是任何一个CKX成员被敲除后都能表现出农艺性状改良。比如利用CRISPR-Cas9技术敲除水稻的OsCKX9基因，增加了穗数，却显著降低了株高、穗长和穗粒数(Duan et al.,2019)。利用RNAi降低水稻中OsCKX4的表达量，显著抑制了根系生长，但可能是基因冗余的原因并未显著影响地上部的生长(Gao et al.,2015)。
细胞分裂素氧化酶基因对小麦农艺性状的调控：普通小麦每个亚基因组携有11-13个CKX基因(Chen et al.,2020)，其中绝大部分的功能未被研究。通过遗传连锁分析，发现位于3D染色体上的TaCKX6a02与籽粒大小、千粒重和灌浆速率显著关联，加性效应来自于小麦品种京411(Lu et al.,2015)。中国农科院作科所贾继增研究员研究发现TaCKX6-D1在小麦种质资源中存在5种等位变异，TaCKX6-D1a与高千粒重关联，并在小麦育种中受到选择(Zhang et al.,2012)。小麦中3A染色体上的TaCKX4存在拷贝数变异，这种变异影响了千粒重和旗叶叶绿素含量(Chang et al.,2015)。利用RNAi技术降低TaCKX2的表达量，显著增加了小麦的穗粒数和产量(Li et al.,2018)。对小麦全基因组CKX基因与农艺性状进行关联分析，发现小麦中的CKX基因主要与千粒重和株高关联(Shoaib et al.,2020)。大麦是小麦的近缘物种，利用RNAi技术降低HvCKX1的表达显著增加了大麦的产量(Zalewski et al.,2010)。由于CKX在调控小麦农艺性状方面的重要作用，科学家认为调控小麦中CKX酶活性可以创制小麦理想株型。但目前未见对小麦中CKX5基因功能的报道，也未见报道利用基因组编辑技术调控小麦CKX基因功能以提高小麦产量和养分利用效率。
发明内容
本发明所要解决的技术问题是如何提高小麦产量。
为了解决以上技术问题，本发明提供了调控细胞分裂素氧化酶基因TaCKX5的物质在提高小麦产量中的应用，所述TaCKX5编码TaCKX5蛋白，所述TaCKX5蛋白是如下A1、A2或A3的蛋白质：
A1、氨基酸序列是序列表中序列2、序列4和序列6中任一种所示的氨基酸序列的蛋白质；
A2、将序列表中序列2、序列4和序列6中任一种所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的与A1)所示的蛋白质具有80％以上的同一性且具有细胞分裂素氧化酶基因活性的蛋白质；
A3、在A1)或A2)的N末端或/和C末端连接蛋白标签得到的融合蛋白质。
上述应用中，序列表中的序列2由530个氨基酸残基组成，序列表中的序列4由531个氨基酸残基组成，序列表中的序列6由531个氨基酸残基组成。
上述应用中，同一性是指氨基酸序列的同一性。可使用国际互联网上的同源性检索站点测定氨基酸序列的同一性，如NCBI主页网站的BLAST网页。例如，可在高级BLAST2.1中，通过使用blastp作为程序，将Expect值设置为10，将所有Filter设置为OFF，使用BLOSUM62作为Matrix，将Gap existence cost，Perresidue gap cost和Lambda ratio分别设置为11，1和0.85(缺省值)并进行检索一对氨基酸序列的同一性进行计算，然后即可获得同一性的值(％)。
上述应用中，所述80％以上的同一性可为至少81％、85％、90％、91％、92％、95％、96％、98％、99％或100％的同一性。
上述应用中，所述TaCKX5蛋白可来源于小麦。
上述应用中，所述TaCKX5基因具体可为如下D1或D2所示的基因：
D1、编码链的编码序列是序列表中序列1、序列3、序列5中任一种的DNA分子；
D2、核苷酸序列是序列表中序列1、序列3、序列5中任一种的DNA分子。
上述应用中，所述调控可通过抑制或降低所述TaCKX5基因的表达进行。具体可为通过CRISPR-Cas9敲除所述TaCKX5基因实现。
为了解决上述技术问题，本发明提供了提高小麦产量的试剂，所述试剂的活性成分为抑制编码所述TaCKX5蛋白的基因的表达、降低所述TaCKX5蛋白的丰度、和/或敲除编码所述TaCKX5蛋白的基因的物质。
上述试剂中，所述物质含有下述F1、F2或F3：
F1、靶向所述基因的sgRNA、siRNA、shRNA、miRNA或反义RNA；
F2、产生靶向所述基因的sgRNA的DNA分子、产生靶向所述基因的siRNA的DNA分子、产生靶向所述基因的shRNA的DNA分子、产生靶向所述基因的miRNA的DNA分子或产生靶向所述基因的反义RNA的DNA分子；
F3、产生靶向所述基因的sgRNA的表达载体、产生靶向所述基因的siRNA的表达载体、产生靶向所述基因的shRNA的表达载体、产生靶向所述基因的miRNA的表达载体或产生靶向所述基因的反义RNA的表达载体。
上述试剂的活性成分还可含有其他生物成分或/和非生物成分，上述药剂的其他活性成分本领域技术人员可根据植物的产量效果确定。
为了解决上述技术问题，本发明还提供一种提高小麦产量的方法，包括如下步骤：抑制受体小麦中所述TaCKX5蛋白的表达、降低所述TaCKX5蛋白的丰度、和/或敲除编码所述TaCKX5蛋白的基因，得到产量高于所述受体小麦的目的小麦。
上述方法中，所述抑制受体小麦所述TaCKX5蛋白的表达、降低所述TaCKX5蛋白的丰度、和/或敲除编码所述TaCKX5蛋白的基因可通过CRISPR/Cas9系统实现，所述CRISPR/Cas9系统包括表达含有Cas9和sgRNA质粒，所述sgRNA的靶序列可为序列表中序列1的第552-第574位所示的核苷酸序列。
上述方法中，所述目的小麦为满足如下条件的小麦：A基因组、B基因组和D基因组其中一个或者两个或者三个在靶点区域发生突变。
本发明还提供蛋白质，为TaCKX5蛋白，是如下A1、A2或A3的蛋白质：
A1、氨基酸序列是序列表中序列2、序列4和序列6中任一种所示的氨基酸序列的蛋白质；
A2、将序列表中序列2、序列4和序列6中任一种所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的与A1)所示的蛋白质具有80％以上的同一性且具有细胞分裂素氧化酶基因活性的蛋白质；
A3、在A1)或A2)的N末端或/和C末端连接蛋白标签得到的融合蛋白质。
本发明还提供所述蛋白质的核酸分子，所述核酸分子具体为如下D1或D2所示的核酸分子：
D1、编码链的编码序列是序列表中序列1、序列3、序列5中任一种的DNA分子；
D2、核苷酸序列是序列表中序列1、序列3、序列5中任一种的DNA分子。
本发明提供调控基因表达的物质在调控小麦产量的应用或在制备调控小麦产量产品中的应用，所述基因编码TaCKX5蛋白，为细胞分裂素氧化酶基因TaCKX5。本发明采用CRISPR/Cas9技术敲除了小麦TaCKX5基因，获得了产量明显提高的小麦新种质。本发明提高了小麦正常氮磷(氮肥施用量纯氮12kg/亩，磷肥施用量为纯磷6kg/亩)和低磷条件下(缺磷土壤，不施用磷肥)的小麦产量，创新了小麦种质资源，对新品种培育、环境保护和粮食安全具有重要意义，具有重大的应用推广价值。
附图说明
图1为本发明实施例1中水培小麦苗期地上部和根系中TaCKX5表达量对氮磷水平的响应结果图。其中，CK为正常氮磷水平(2mM N+0.2mM P)，LN为低氮处理(0.2mM N+0.2mMP)，LP为低磷处理(2mM N+0.01mM P)。Shoot表示地上部；Root表示根系。TaCKX5表达量为相对于内参TaActin的相对表达量，数据为3次生物学重复的Mean±SE。*表示与CK相比差异显著性达到P<0.05，**表示与CK相比差异显著性达到P<0.01。
图2为本发明实施例1中大田试验中TaCKX5在小麦开花后14天不同组织的特异性表达结果图，数据为4次生物学重复的Mean±SE。
图3为本发明实施例2中载体pJIT163-ubi-2NLS-rCas9-wheat的结构示意图。
图4为本发明实施例2中载体pU6-sgRNA的结构示意图。
图5为本发明实施例2中载体pAHC25的结构示意图。
图6为本发明实施例2中sgRNA序列与小麦品种中国春参考基因组序列的比对结果图。
图7为本发明实施例2中正常氮磷处理下突变系幼穗中的细胞分裂素和IAA含量检测结果。其中，iPR为异戊烯基腺嘌呤核苷，iP为异戊烯基腺嘌，tZ为反式玉米素，tZR为反式玉米素核苷，cZ为顺式玉米素，cZR为顺式玉米素核苷，DHZ为二氢玉米素，IAA为吲哚乙酸。数据为4个生物学重复的Mean±SE。*表示转基因系与受体亲本之间的差异显著性达到P<0.05。
具体实施方式
下面结合具体实施方式对本发明进行进一步的详细描述，给出的实施例仅为了阐明本发明，而不是为了限制本发明的范围。以下提供的实施例可作为本技术领域普通技术人员进行进一步改进的指南，并不以任何方式构成对本发明的限制。
下述实施例中的定量试验，均设置三次重复实验，结果取平均值。
下述实施例中的实验方法，如无特殊说明，均为常规方法。下述实施例中所用的材料、试剂等，如无特殊说明，均可从商业途径得到。
下述实施例中的小麦品种科农199由中国科学院遗传与发育生物学研究所选育，审定编号：国审麦2006017。科农199记载于非专利文献“李俊明、张相岐、张爱民、王志国、安调过、纪军、王静.高产广适小麦新品种—科农199[J].麦类作物学报，2007(02):368.”，公众可以从中国科学院遗传与发育生物学研究所获得，以重复本申请实验，不可作为其它用途使用。
实施例1普通小麦中TaCKX5序列克隆及表达量检测
一、TaCKX5的cDNA序列克隆
提取小麦品种科农199的总RNA，并检测RNA浓度及质量。
依据东洋纺(ReverTra qPCR RT Master Mix with gDNA Remover，FSQ-301)反转录试剂盒进行反转录，得到cDNA，于-20℃储存或稀释20倍用于普通PCR扩增。PCR扩增TaCKX5序列，所用的引物为：
Primer F：5′-TGGCAATGGCGCGGTGCTTGGT-3′；
Primer R：5′-CAATCAGTCTTCTCAAAAATCGGAC-3′。
PCR反应体系和PCR程序如下：
1)PCR反应体系(TOYOBO,KFX-101，含有2×PCR buffer for KOD FX、2mM dNTPs与KOD FX)
cDNA(100ng/μL)：2μL
Primer F(10μM)：1.5μL
Primer R(10μM)：1.5μL
2×PCR buffer for KOD FX：25μL
2mM dNTPs：10μL
KOD FX(1.0U/μL)：1μL
ddH2O：9μL
2)PCR程序
98℃，5min；98℃，30s，62℃，30s，68℃，2min，36cycles；68℃，10min，10℃，保温。
PCR扩增后测序，用1％琼脂糖凝胶电泳检测所得PCR产物，并连接至pEASY-Blunt载体中进行测序，结果显示：
小麦cDNA中：A基因组第3部分同源群染色体(对应A染色体组第3部分同源群染色体)中的基因TaCKX5-3A的编码区如序列表的序列1所示(编码序列表的序列2所示的蛋白质)；B基因组第3部分同源群染色体(对应B染色体组第3部分同源群染色体)中的基因TaCKX5-3B的编码区如序列表的序列3所示(编码序列表的序列4所示的蛋白质)；D基因组第3部分同源群染色体(对应D染色体组第3部分同源群染色体)中的基因TaCKX5-3D的编码区如序列表的序列5所示(编码序列表的序列6所示的蛋白质)。
二、小麦中TaCKX5表达量检测方法
以小麦品种科农199为材料进行如下营养液培养实验和大田实验，检测TaCKX5表达量。
1、营养液培养实验
1.1、营养液培养方法
选取饱满一致的种子，10％H2O2灭菌20min，清水冲洗数次，将种子均匀摆放于铺有滤纸的培养皿中，加入适量饱和CaSO4，于23℃培养箱中暗催芽1天后移入带有纱网的培养盘中，自来水培养6天后，挑取长势一致的幼苗去掉胚乳后移入8L水培培养盒中继续培养。营养液培养实验分为正常氮磷水平(CK)、低氮处理(LN)、低磷处理(LP)三组处理(营养液配方见表1)。每个处理设置3个重复，每隔2天更换一次培养液。培养温度：20℃；光周期：15h光照/9h；光照强度：405μmol m-2s-1。培养3周后采集根系和地上部样品分析基因表达量。
表1.小麦水培试验营养液配方
1.2、荧光实时定量PCR方法检测不同小麦组织中TaCKX5的表达量
用荧光实时定量PCR(qRT-PCR)方法检测不同小麦组织中TaCKX5的表达量。
提取开花后14天不同小麦组织的总RNA，并检测RNA浓度及质量。依据东洋纺(ReverTra qPCR RT Master Mix with gDNA Remover，FSQ-301)反转录试剂盒进行反转录，得到cDNA，于-20℃储存或稀释20倍用于实时定量PCR(qRT-PCR)扩增反应。
进行qRT-PCR反应的反应体系和反应条件如下：
每个样品设三个技术重复。2x SYBR Green I Master为(Roche，480SYBR Green I Master，4707516001)。
鉴定TaCKX5基因相对表达量的引物由针对TaCKX5-3A基因的引物对TaCKX5-3AF/TaCKX5-3AR、针对TaCKX5-3B基因的引物对TaCKX5-3BF/TaCKX5-3BR、针对TaCKX5-3D基因的引物对TaCKX5-3AF/TaCKX5-3DR组成：
TaCKX5-3AF:5′-ACCGGCAGATCCTCGACTTCTGCG-3′；
TaCKX5-3AR:5′-CGCGAAGCGCGCCCACCTCTTCT-3′；
TaCKX5-3BF:5′-GAACCGGCAGATCCTCGACTTCTGCA-3′；
TaCKX5-3BR:5′-GGATCGAATTGGGCCTTGAGGCCG-3′；
TaCKX5-3DF:5′-GAACCGGGAGATCCTCGACTTCTGCG-3′；
TaCKX5-3DR:5′-TCTCCTCCCCGAAATGGGCCTC-3′。
内标基因TaActin引物为：
TaActin-F：5′-ACCTTCAGTTGCCCAGCAAT-3′；
TaActin-R：5′-CAGAGTCGAGCACAATACCAGTTG-3′。
qRT-PCR反应程序：
95℃，2min；95℃，15s，60℃，15s，72℃，20s，45cycles；95℃，5s，65℃，1min，40℃，20s。
Real-Time PCR使用的仪器为Roche,LightCycler480 II。
反应结束后确认qRT-PCR的扩增曲线和溶解曲线，可以通过溶解曲线分析，确认PCR反应的特异性。计算Ct平均值。根据相应的内参基因的表达量来确定目的基因的相对表达量。基因的相对表达量计算公式为：
相对表达量＝2-ΔCt，△Ct＝Ct目标基因-Ct内标基因。
结果见图1，表明低氮处理(LN)和低磷处理(LP)均不同程度抑制了根系和地上部中TaCKX5的表达，低磷处理尤为显著，说明TaCKX5的表达受到氮、磷供应水平的调控。TaCKX5-3A、TaCKX5-3B和TaCKX5-3D表达量相比较，TaCKX5-3B的表达量最低。
2、大田试验
表型鉴定试验地位于河北省石家庄市农林科学研究院北方农业科技园(河北赵县)。试验设正常施用氮磷肥(纯氮12kg/亩、纯磷6kg/亩)、和低磷(纯氮12kg/亩、不施磷肥)两个处理，每个处理4次生物学重复。氮肥为尿素，磷肥为重过磷酸钙。每个重复每个材料2m行长，株距5cm，行距23cm。在拔节期采集正常氮磷处理幼穗测定细胞分裂素和IAA含量。开花后14天检测正常氮磷处理小麦植株各组织的TaCKX5的表达量，检测方法见步骤1.2，数据为4次生物学重复。在成熟期取每个重复每个材料20株的主穗考查穗粒数和千粒重，考察2行的穗数和籽粒产量。
在开花后14天分不同组织取样，检测科农199中TaCKX5表达量的结果见图2，TaCKX5在茎和茎节表达量较高，预示TaCKX5参与调控细胞分裂素在不同组织的分布。在TaCKX5-3A、TaCKX5-3B和TaCKX5-3D中，TaCKX5-3B的表达量最低。
实施例2基因编辑TaCKX5获得的突变系及其TaCKX5表达量检测与性状表现一、基因编辑TaCKX5获得突变系
利用CRISPR/Cas9介导的基因组编辑技术对小麦A、B、D基因组上的编码细胞分裂素氧化酶基因TaCKX5-3A、TaCKX5-3B和TaCKX5-3D设计靶点序列进行基因敲除，所用载体如下：
pJIT163-ubi-2NLS-rCas9-wheat的结构示意图见图3，含Cas9基因线性最小表达框序列总长6917bp，其中启动子为玉米Ubiquitin(UBI)启动子，长度为1987bp，优化后Cas9(rCas9)的CDS序列和核定位序列长度为4206bp，终止子为CaMV term终止子，长度为724bp。
含目的基因编辑靶位点的载体pU6-sgRNA的结构示意图见图4，引入TaU6-sgRNA线性最小表达框序列总长为476bp，其中TaU6启动子长度为363bp。
sgRNA的靶位点序列：5′-CCACCACGGGCCCCAGATCAGCA-3′(如序列表序列1的第557-第579位/序列3的第557-第579位/序列5的第557-第579位所示的核苷酸序列，其中下划线指示的序列为ApaI酶识别位点序列)。
含有筛选标记Bar基因的载体pAHC25结构示意图见图5，引入Bar基因的线性最小表达框序列总长为2832bp，其中启动子为玉米Ubiquitin启动子(Ubi promoter)，长度为1987bp，Bar基因(BAR)的CDS序列长度为552bp，终止子为Nos终止子(NOS)，长度为252bp。
采用基因枪介导的转化方法，将载体pJIT163-ubi-2NLS-rCas9-wheat、载体pU6-sgRNA和载体pAHC25的线性最小表达框共同转化受体小麦科农199的愈伤组织，Cas9蛋白与sgRNA结合形成RNA-蛋白质复合体，共同完成识别并切割TaCKX5的DNA靶序列的功能。其中Cas9蛋白作为核酸酶切割双链DNA，而sgRNA则通过碱基互补配对决定靶序列的特异性，从而突变小麦中的TaCKX5基因靶序列。利用筛选标记Bar基因筛获得T0代转基因植株，并连续自交获得T2代和T3代转基因系。
通过采用PCR、酶切、测序相结合的方法鉴定T2代和T3代转基因系目的基因靶位点的突变情况。具体方法如下：提取转基因系的基因组DNA作为模板，在TaCKX5基因组编辑的靶位点序列附近分别对A、B、D组基因序列设计PCR检测引物(见表2)，引物序列和扩增产物大小如下表，扩增程序为94℃4分钟；94℃30秒，58℃30秒，68℃90秒，36个循环；68℃10分钟。然后用限制性内切酶ApaI(识别位点序列为GGGCCC)对PCR产物进行酶切，PCR及酶切产物用2％琼脂糖胶检测，被切开的说明与野生型受体对照一样，不能被切开说明靶位点序列发生了突变。本实施例中TaCKX5-3D的突变位点不影响ApaI酶切位点，因此不能用ApaI酶切检测TaCKX5-3D的突变，需要用测序的方法鉴定。
表2.检测TaCKX5突变的引物序列
通过对T3代转基因系进行PCR产物测序，Bar和Cas9基因PCR检测，筛选获得突变位点遗传稳定，不含Bar和Cas9基因，且产量性状突出的突变系CKX5-CAS4d(敲除TaCKX5-3D的单突变)、CKX5-CAS10ad(敲除TaCKX5-3A和TaCKX5-3D的双突变)、CKX5-CAS10d(敲除TaCKX5-3D的单突变)。各突变系TaCKX5的序列突变如表3所示。
CKX5-CAS4d在TaCKX5-3D的CDS(序列表序列5)第563位缺失1个碱基C，导致从第186个氨基酸开始突变，并提前终止，推断的突变蛋白序列共198aa(如序列表序列8所示)。
CKX5-CAS10ad在TaCKX5-3A的CDS(序列表序列1)第564位碱基G缺失，导致从第189个氨基酸开始突变，提前终止，推断的突变蛋白序列共198aa(如序列表序列7所示)；同时CKX5-CAS10ad在TaCKX5-3D的CDS(序列表序列5)第563位缺失1个碱基C，导致从第186个氨基酸开始突变，并提前终止，推断的突变蛋白序列共198aa(如序列表序列8所示)。
CKX5-CAS10d在TaCKX5-3D的CDS(序列表序列5)第563位缺失1个碱基C，导致从第186个氨基酸开始突变，并提前终止，推断的突变蛋白序列共198aa(如序列表序列8所示)。
表3.T3代突变株系TaCKX5序列突变
用sgRNA序列CCACCACGGGCCCCAGATCAGCA比对小麦品种中国春参考基因组序列，共比对出4条序列，其中3条为目标靶位点，位于7B上的序列位于基因间区域，且sgRNA序列与7B染色体上的比对序列存在2bp碱基的差异，具体见图6，表明用本实施例所用的sgRNA不会导致脱靶。
二、正常氮磷处理下突变系的细胞分裂素和IAA含量检测
正常氮磷处理下(纯氮12kg/亩、纯磷6kg/亩)种植步骤一获得的突变系CKX5-CAS4d(敲除TaCKX5-3D的单突变)、CKX5-CAS10ad(敲除TaCKX5-3A和TaCKX5-3D的双突变)、CKX5-CAS10d(敲除TaCKX5-3D的单突变)，以科农199作为对照。
在拔节期采集正常氮磷处理幼穗测定了细胞分裂素和IAA含量。测定方法如下：
代谢物提取：取80±5mg样品于2mL离心管中，加入30μL内标溶液，加入1mL乙腈水溶液(1％FA)，震荡混匀2min；4℃避光抽提12h，14000g离心20min，取上清液800μL，氮气吹干，用100μL乙腈水(1:1，v/v)复溶，14000g离心20min，取上清进样分析。
色谱-质谱分析：样品采用Waters I-Class LC超高效液相色谱系统进行分离。流动相:A液为0.05％FA水溶液，B液为0.05％FA乙腈。样品置于4℃自动进样器中，柱温45℃，流速为400μL/min，进样量2μL。相关液相梯度如下：0-10min，B液从2％线性变化到98％；10-10.1min，B液从98％线性变化至2％；11.1-13min，B液维持在2％。样本队列中每间隔一定数量的实验样本设置一个QC样本，用于检测和评价系统的稳定性及重复性。
采用5500QTRAP质谱仪(AB SCIEX)在正/负离子模式下进行质谱分析。5500QTRAPESI源条件如下：source temperature 500℃，ion Source Gas1(Gas1)：45，IonSource Gas2(Gas2)：45，Curtain gas(CUR)：30，ionSapary Voltage Floating(ISVF)-4500V；采用MRM模式检测待测离子对。
数据处理：采用Multiquant软件提取色谱峰面积及保留时间。根据标准曲线，计算样品中测得植物激素的含量。
测定结果如图7所示：敲除TaCKX5-3A和TaCKX5-3D的双突变(aadd，对应突变系CKX5-CAS10ad)幼穗中的iPR和tZR显著高于亲本科农199(WT)，分别增加43.3％和50.1％。敲除TaCKX5-3D的单突变(dd，对应突变系CKX5-CAS4d和CKX5-CAS10d)幼穗中iPR和tZR比亲本科农199分别增加18.7％和32.6％。双突变的增加效果明显高于单突变，说明敲除TaCKX5对幼穗中细胞分裂素含量的影响存在基因剂量效应。敲除突变体中iPR和tZR含量的增加解释了穗粒数增加的表型。敲除TaCKX5-3A和TaCKX5-3D并不影响幼穗中IAA含量。
三、TaCKX5突变系在正常氮磷和缺磷条件下的产量
在河北省石家庄市农林科学研究院赵县实验基地，在正常氮磷和低磷条件下比较了突变系CKX5-CAS4d(敲除TaCKX5-3D的单突变)、CKX5-CAS10ad(敲除TaCKX5-3A和TaCKX5-3D的双突变)、CKX5-CAS10d(敲除TaCKX5-3D的单突变)和野生型亲本科农199的产量性状。
试验设正常施用氮磷肥(纯氮12kg/亩、纯磷6kg/亩)和低磷(纯氮12kg/亩、不施磷肥)两个处理，每个处理4次生物学重复。每个重复每个材料2m行长，株距5cm，行距23cm。在成熟期取每个重复每个材料取20株的主穗考查穗粒数和千粒重，考察2行的穗数和籽粒产量。结果见表4-表7：
表4.敲除TaCKX5对小麦籽粒产量的影响
表5.敲除TaCKX5对小麦穗数的影响
表6.敲除TaCKX5对小麦主穗粒数的影响
表7.敲除TaCKX5对小麦主穗千粒重的影响
表4-表7中，数据为4个生物学重复的Mean±SE。CKX5-CAS4d为株系17CAS184-01的数据，CKX5-CAS10ad为株系17CAS168-02的数据，CKX5-CAS10d为株系17CAS162-07的数据。*和**表示转基因系与受体亲本之间的差异显著性分别达到P<0.05和P<0.01。
结果表明，在正常氮磷条件下，CKX5-CAS4d(敲除TaCKX5-3D的单突变)、CKX5-CAS10ad(敲除TaCKX5-3A和TaCKX5-3D的双突变)和CKX5-CAS10d(敲除TaCKX5-3D的单突变)比受体亲本增产19.9％-24.8％；在低磷条件下，比受体亲本增产23.5％-28.7％(表4)。对产量三要素的分析结果显示，在低磷条件下三个突变系的穗粒数均显著高于野生型亲本科农199；在正常氮磷条件下有两个突变系的主穗粒数显著高于野生型亲本科农199(表6)，穗粒数增加是突变系在正常氮磷和低磷条件下增产的主要因素。突变系穗粒数的增加可能是因为突变系幼穗中iPR和tZR含量增加所致。
以上对本发明进行了详述。对于本领域技术人员来说，在不脱离本发明的宗旨和范围，以及无需进行不必要的实验情况下，可在等同参数、浓度和条件下，在较宽范围内实施本发明。虽然本发明给出了特殊的实施例，应该理解为，可以对本发明作进一步的改进。总之，按本发明的原理，本申请欲包括任何变更、用途或对本发明的改进，包括脱离了本申请中已公开范围，而用本领域已知的常规技术进行的改变。按以下附带的权利要求的范围，可以进行一些基本特征的应用。
参考文献
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序列表
<110> 中国科学院遗传与发育生物学研究所
<120> 调控细胞分裂素氧化酶基因TaCKX5的物质在提高小麦产量中的应用
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<213> 小麦(Triticum aestivum)
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tggcaatggc gcggtgcttg gtcttcatgg tgttccagat atactgcctc atctcgaccg 60
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cgctgctggg cgacctgaac ttcctcccgg gcacggtgtt cacgacggac ctggcgtacg 1080
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aggtgccgca cccgtggctg aacctgttcg tgccggcgtc gcgcatcgcc gacttcgacc 1200
gcggcgtctt ccgcggcatc ctcggcagcc gcacctccgg cgggcccatc ctcatctacc 1260
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<211> 531
<212> PRT
<213> 小麦(Triticum aestivum)
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Met Ala Arg Cys Leu Val Phe Met Val Phe Gln Ile Tyr Cys Leu Ile
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Ser Thr Val Gly Leu Pro Leu Glu Pro Ala Glu Leu Leu Gln Leu Gly
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Glu Ala Ser Arg Asp Phe Gly Gly Phe Thr Arg Gly Glu Pro Leu Ala
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Val Tyr His Pro Ser Gly Ala Gly Asp Val Ala Ala Leu Val Arg Ala
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Ala Tyr Gly Ser Ala Arg Asp Ile Arg Val Ser Ala Arg Gly His Gly
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His Ser Ile Arg Gly Gln Ala Gln Val Ala Gly Gly Val Val Val Asp
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Met Ser Arg Gly Gly Lys Ala Pro Ser Arg Ala Leu Pro Val Tyr Ser
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Ser Trp Thr Asp Tyr Leu Tyr Leu Ser Val Gly Gly Thr Leu Ser Asn
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Ala Gly Ile Ser Gly Gln Ala Phe His His Gly Pro Gln Ile Ser Asn
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Val Tyr Glu Leu Asp Val Val Thr Gly Lys Gly Glu Ala Val Thr Cys
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Lys Leu Ser Ser Leu Lys His His Thr Gly Val Leu Tyr Cys Leu Glu
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Ile Tyr Pro Met Asn Lys His Lys Trp Asp Pro Arg Ser Ser Val Val
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Thr Pro Asp Glu Glu Val Phe Tyr Leu Val Ala Phe Leu Arg Ser Ala
435 440 445
Leu Pro Gly Ala Pro Gln Ser Leu Glu Ala Leu Ala Arg Gln Asn Arg
450 455 460
Glu Ile Leu Asp Phe Cys Ala Glu Ala Gly Ile Gly Ala Arg Gln Tyr
465 470 475 480
Leu Pro Asn His Lys Ser Gln Pro Glu Trp Glu Ala His Phe Gly Glu
485 490 495
Glu Arg Trp Ala Arg Phe Ala Gly Leu Lys Ala Gln Phe Asp Pro Arg
500 505 510
Ala Met Leu Ala Thr Gly Gln Gly Ile Phe Pro Pro Pro Ala Leu Leu
515 520 525
Ser Asp Phe
530
<210> 7
<211> 198
<212> PRT
<213> 小麦(Triticum aestivum)
<400> 7
Met Ala Arg Cys Leu Val Phe Met Val Phe Gln Ile Tyr Cys Leu Ile
1 5 10 15
Ser Thr Val Gly Leu Pro Leu Glu Pro Ala Glu Leu Leu Gln Leu Gly
20 25 30
Gly Asp Val Gly Gly Gly Arg Leu Ser Thr Asp Pro Ala Asp Val Leu
35 40 45
Glu Ala Ser Arg Asp Phe Gly Gly Phe Thr Arg Gly Glu Pro Leu Ala
50 55 60
Val Tyr His Pro Ser Gly Ala Asp Asp Val Ala Ala Leu Val Arg Ala
65 70 75 80
Ala Tyr Gly Ser Ala Arg Asp Ile Arg Val Ser Ala Arg Gly His Gly
85 90 95
His Ser Ile Ser Gly Gln Ala Gln Val Pro Gly Gly Val Val Val Asp
100 105 110
Met Ser Leu Gly Gly Lys Ala Pro Ser Arg Ala Leu Pro Val Tyr Ser
115 120 125
Pro Glu Leu Gly Gly His Tyr Val Asp Val Trp Gly Gly Glu Leu Trp
130 135 140
Ile Asp Val Leu Asn Trp Thr Leu Ser His Gly Gly Leu Ala Pro Arg
145 150 155 160
Ser Trp Thr Asp Tyr Leu Tyr Leu Ser Val Gly Gly Thr Leu Ser Asn
165 170 175
Ala Gly Ile Ser Gly Gln Ala Phe His His Gly Pro Arg Ser Ala Met
180 185 190
Ser Thr Ser Ser Thr Ser
195
<210> 8
<211> 198
<212> PRT
<213> 小麦(Triticum aestivum)
<400> 8
Met Ala Arg Cys Leu Val Phe Met Val Phe Gln Ile Tyr Cys Leu Ile
1 5 10 15
Ser Thr Val Gly Leu Pro Leu Glu Pro Ala Glu Leu Leu Gln Leu Gly
20 25 30
Gly Asp Val Gly Gly Gly Arg Leu Ser Thr Asp Pro Ala Asp Val Leu
35 40 45
Glu Ala Ser Arg Asp Phe Gly Gly Phe Thr Arg Gly Glu Pro Leu Ala
50 55 60
Val Tyr His Pro Ser Gly Ala Gly Asp Val Ala Ala Leu Val Arg Ala
65 70 75 80
Ala Tyr Gly Ser Ala Arg Asp Ile Arg Val Ser Ala Arg Gly His Gly
85 90 95
His Ser Ile Arg Gly Gln Ala Gln Val Ala Gly Gly Val Val Val Asp
100 105 110
Met Ser Arg Gly Gly Lys Ala Pro Ser Arg Ala Leu Pro Val Tyr Ser
115 120 125
Pro Glu Leu Gly Gly His Tyr Val Asp Val Trp Gly Gly Glu Leu Trp
130 135 140
Ile Asp Val Leu Asn Trp Thr Leu Ser His Gly Gly Leu Ala Pro Arg
145 150 155 160
Ser Trp Thr Asp Tyr Leu Tyr Leu Ser Val Gly Gly Thr Leu Ser Asn
165 170 175
Ala Gly Ile Ser Gly Gln Ala Phe His Gln Gly Pro Arg Ser Ala Met
180 185 190
Ser Thr Ser Ser Thr Ser
195
Claims (4)
1.抑制或降低TaCKX5基因的表达的sgRNA在提高小麦低磷条件下产量中的应用，所述TaCKX5编码TaCKX5蛋白，所述TaCKX5蛋白是如下A1或A2的蛋白质：
A1、氨基酸序列是序列表中序列2、序列4和序列6中任一种所示的氨基酸序列的蛋白质；
A2、在A1)的N末端或/和C末端连接蛋白标签得到的融合蛋白质；
所述TaCKX5基因具体为如下D1或D2所示的基因：
D1、编码链的编码序列是序列表中序列1、序列3、序列5中任一种的DNA分子；
D2、核苷酸序列是序列表中序列1、序列3、序列5中任一种的DNA分子。
2.一种提高小麦低磷条件下产量的方法，其特征在于：包括如下步骤：抑制受体小麦中权利要求1中所述TaCKX5蛋白的表达、降低权利要求1中所述TaCKX5蛋白的丰度、和/或敲除编码权利要求1中所述TaCKX5蛋白的基因，得到低磷条件下产量高于所述受体小麦的目的小麦。
3.根据权利要求2所述的方法，其特征在于：所述抑制受体小麦中权利要求1中所述TaCKX5蛋白的表达、降低权利要求1中所述TaCKX5蛋白的丰度、和/或敲除编码权利要求1中所述TaCKX5蛋白的基因通过CRISPR/Cas9系统实现，所述CRISPR/Cas9系统包括表达含有sgRNA和Cas9的质粒，所述sgRNA的靶序列为序列表中序列1的第552-第574位所示的核苷酸序列。
4.权利要求2或3所述的方法，其特征在于：所述目的小麦为满足如下条件的小麦：A基因组、B基因组和D基因组其中一个或者两个或者三个在靶点区域发生突变。
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Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN101235381A (zh) * | 2007-10-30 | 2008-08-06 | 山东农业大学 | 小麦细胞分裂素氧化酶基因TaCKX1序列及其克隆与应用 |
| CN107723311A (zh) * | 2017-11-28 | 2018-02-23 | 中国科学院遗传与发育生物学研究所 | 一种与植物种子粒重相关的TaCKX2‑3D蛋白及其编码基因与应用 |
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Patent Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN101235381A (zh) * | 2007-10-30 | 2008-08-06 | 山东农业大学 | 小麦细胞分裂素氧化酶基因TaCKX1序列及其克隆与应用 |
| CN107723311A (zh) * | 2017-11-28 | 2018-02-23 | 中国科学院遗传与发育生物学研究所 | 一种与植物种子粒重相关的TaCKX2‑3D蛋白及其编码基因与应用 |
Non-Patent Citations (8)
| Title |
|---|
| Different sets of TaCKX genes affect yield-related traits in wheat plants grown in a controlled environment and in field conditions;szala k等;bmc plant biology;第第20卷卷(第第1期期);第496篇，第1-13页 * |
| NCBI genbank.PREDICTED: Aegilops tauschii subsp. strangulata cytokinin dehydrogenase 5 (LOC109757480), transcript variant X2, mRNA，NCBI Reference Sequence: XM_020316300.2，1937bp mRNA linear.NCBI genbank.2021,第1-2页. * |
| NCBI genbank.PREDICTED_ Triticum dicoccoides cytokinin dehydrogenase 5-like (LOC119277171), mRNA，NCBI Reference Sequence: XM_037558417.1，1956bp mRNA linear.NCBI genbank.2020,第1-2页. * |
| NCBI genbank.REDICTED: Triticum dicoccoides cytokinin dehydrogenase 5-like (LOC119269181), mRNA，NCBI Reference Sequence: XM_037550962.1，1980bp mRNA linear.NCBI genbank.2020,第1-2页. * |
| PREDICTED: Aegilops tauschii subsp. strangulata cytokinin dehydrogenase 5 (LOC109757480), transcript variant X2, mRNA，NCBI Reference Sequence: XM_020316300.2，1937bp mRNA linear;NCBI genbank;NCBI genbank;第1-2页 * |
| PREDICTED_ Triticum dicoccoides cytokinin dehydrogenase 5-like (LOC119277171), mRNA，NCBI Reference Sequence: XM_037558417.1，1956bp mRNA linear;NCBI genbank;NCBI genbank;第1-2页 * |
| REDICTED: Triticum dicoccoides cytokinin dehydrogenase 5-like (LOC119269181), mRNA，NCBI Reference Sequence: XM_037550962.1，1980bp mRNA linear;NCBI genbank;NCBI genbank;第1-2页 * |
| 小麦细胞分裂素氧化/脱氢酶基因(TaCKX5)的克隆及其染色体定位;张磊等;中国农业科学;第第41卷卷(第第3期期);第636-642页 * |
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