CN114457091A - 一种影响小麦籽粒品质的基因TaXip及其应用 - Google Patents
一种影响小麦籽粒品质的基因TaXip及其应用 Download PDFInfo
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- CN114457091A CN114457091A CN202111577854.2A CN202111577854A CN114457091A CN 114457091 A CN114457091 A CN 114457091A CN 202111577854 A CN202111577854 A CN 202111577854A CN 114457091 A CN114457091 A CN 114457091A
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Abstract
本发明公开了一种影响小麦籽粒品质的基因TaXip及其应用。所述基因TaXip包括基因TaXip‑6A、基因TaXip‑6B和基因TaXip‑6D，其中基因TaXip‑6A包含SNP分子标记，其位于小麦染色体6A上，所述SNP分子标记的多态性为C/G。利用基因TaXip的检测引物能够鉴定出高品质小麦籽粒，方法简单、效率高。且本发明经实验证实TaXip基因敲除具有更高的SDS沉降值和稳定时间，因此能够用来鉴定和筛选高品质的小麦品种，有利于提高小麦育种效率，加速小麦品质遗传改良。
Description
技术领域
本发明属于小麦分子遗传育种技术领域，具体涉及一种影响小麦籽粒品质的基因TaXip及其应用。
背景技术
小麦是世界上广泛种植的三大粮食作物之一，是世界上约30%以上人口的主要营养和能量来源。培育优质小麦新品种是小麦育种的重要目标之一，也是小麦商品粮品质和制作各种符合人们需求食品的基础。我国小麦品质育种还存在较大差距，比如强筋不强、弱筋不弱、专用粉生产较为落后等问题。因此，分离和鉴定小麦品质相关的基因，对小麦品质遗传改良具有重要意义。
小麦中鉴定出三种不同类型的木聚糖酶抑制剂：木聚糖酶抑制剂（the Triticumaestivum. L Xylanase-inhibitor，TAXI）、木聚糖酶抑制蛋白（Xylanse InhibitorProtein，XIP）和类甜蛋白样木聚糖酶抑制剂（Thaumatin-like Xylanase Inhibitor TL-XI）。在小麦中已经鉴定出了XIP-I、XIP-III和XIP-R2三种XIP型抑制蛋白。通过对Xip基因进行CRISPR/Cas9基因编辑验证，可为利用该基因进行优质性状的遗传改良提供理论依据。
发明内容
本发明的目的在于提供了一种影响小麦籽粒品质的基因TaXip及其应用。
为实现上述发明目的，本发明采取的技术方案为：
本发明提供了一种影响小麦籽粒品质的基因TaXip，所述基因TaXip包括基因TaXip-6A、基因TaXip-6B和基因TaXip-6D。
进一步的，所述基因TaXip-6A的核苷酸序列如SEQ ID No.1所示，其编码的氨基酸序列如SEQ ID No.2所示。
进一步的，所述基因TaXip-6B的核苷酸序列如SEQ ID No.3所示，其编码的氨基酸序列如SEQ ID No.4所示。
进一步的，所述基因TaXip-6D的核苷酸序列如SEQ ID No.5所示，其编码的氨基酸序列如SEQ ID No.6所示。
本发明还提供了所述的影响小麦籽粒品质的基因TaXip的检测引物，所述检测引物的序列如SEQ ID No.7~ SEQ ID No.12所示。
本发明还提供了一种高品质小麦籽粒的鉴定方法，所述鉴定方法为：
（1）提取小麦籽粒的DNA；
（2）利用序列如SEQ ID No.7~ SEQ ID No.12所示的检测引物对DNA进行扩增；
（3）扩增结果进行比对，如果序列中出现多态性突变，则样品小麦籽粒高品质存在差异。
进一步的，所述步骤（3）中多态性突变为C/G突变，其发生在位于小麦染色体6A上的基因TaXip-6A上。
进一步的，所述C/G突变中G碱基变异能够提高小麦籽粒的SDS沉降值和稳定时间。
本发明还提供了所述的影响小麦籽粒品质的基因TaXip在用于调控小麦籽粒品质性状中的应用。
进一步的，所述应用方法为：通过敲除基因TaXip来提高小麦籽粒的SDS沉降值和稳定时间。
本发明还提供了所述的影响小麦籽粒品质的基因TaXip的检测引物在用于选育高品质小麦籽粒的应用。
与现有的技术相比，本发明的有益效果和优点是：
1、本发明根据前期QTL的分析结果，从普通小麦中扩增了Xip基因，并利用CRISPR/Cas9基因编辑方法，对3个同源基因TaXip-6A、TaXip-6B和TaXip-6D的靶位点进行编辑，进而通过实验证实了TaXip-6A基因是该QTL位点上影响籽粒品质的主效基因，因此本发明是首次公开了影响小麦籽粒品质的TaXip-6A基因，位于小麦染色体6A上，其多态性SNP为C/G。
2、本发明通过基因编辑试验得到了2种突变体，其基因的表达不同，因此品质性状有差异，aaBBDD突变体的SDS沉降值和稳定时间均极显著高于野生型，AAbbdd突变体的SDS沉降值极显著高于野生型，稳定时间和野生型差异不显著。说明该分子标记TaXIP-6A能够用来筛选和鉴定高品质的小麦品种，有利于提高小麦育种效率，加速小麦品质遗传改良。
附图说明
图1是载体结构图；其中RB/LB表示载体的左右边界；TaU3P是小麦U3基因启动子；sgRNA位点是指引导RNA克隆位点；gRNA SC为gRNA支架；PUbi是泛素基因启动子；zCas9是玉米密码子优化的Cas9；Tnos是Nos终止子；P35S是35S启动子。
图2是CRISPR/ Cas9介导的QSt/Sv-6A-2851定位和TaXIP突变体的开发；其中，(A)利用IciMapping4.1(黑条)、MapQTL5.0(红条)和TASSEL5.0软件对QSt/Sv-6A-2851进行QTL作图；(B) 三个同源基因TaXip-6A、TaXip-6B和TaXip-6D在Fielder中的氨基酸序列比对；(C) TaXip基因sgRNAs靶位点选择及CRISPR/Cas9诱导的T2:3代突变型(aaBBDD和AAbbdd)示意图，条框为外显子，水平线为内含子，PAM序列用红色突出显示；(D) T2:3突变株系与WT(Fielder)的SV和ST差异比较。
图3为小麦和水稻的XIP和chitinase进化树。
图4为TaXip-6A、TaXip-6B和TaXip-6D基因编码区序列比较。
图5为T2:3代突变体氨基酸变化。
具体实施方式
下面结合附图和具体实施方式对本发明的技术方案进一步的详细说明，但本发明要求保护的范围并不局限于实例表述的范围。下述实施例中，如无特殊说明，所使用的实验方法均为常规方法，所用材料、试剂等均可从生物或化学试剂公司购买。
本发明中所用到的小麦品种：QTL分析材料为冬小麦品种“泰农18”（TN18）和“临麦6号”（LM6）及其构建的重组自交系（RIL）群体；基因编辑材料为春性品种Fielder。上述材料均可由山东农业大学处获得。
实施例1：小麦TaXip候选基因的确定
利用IciMapping4.1、MapQTL5.0软件，对TL-RIL群体的品质性稳定时间（stability time，ST）和沉淀值（SDS-sedimentation volume，SV）进行QTL定位（图2A）。利用TASSEL5.0进行GWAS分析。如表1和表2所示，在多个环境下检测到ST和SV的QTLQSt/Sv- 6A-2851，平均贡献率大于5%，峰值区间为2850.3~2860.4。
表1 品质性状QTL定位
表2 品质性状关联分析（TASSEL5.0软件）
对照遗传图谱发现这一区域包含2个在中国春参考基1因组V1.1注释候选基因（表3），TraesCS6A02G076900和TraesCS6A02G077000。TraesCS6A02G076900注释为蛋白激酶超家族（ABC-2 type transporter family protein），含八肽/ Phox/ Bem1p结构域；TraesCS6A02G077000注释为木聚糖酶抑制蛋白（Xylanase inhibitor protein）。
经过序列分析发现，TraesCS6A02G076900具有6个SNP，3个位于5’-UTR区，2个位于内含子区，1个位于3’-UTR区，没有氨基酸的变化。TraesCS6A02G077000在390 bp（从ATG开始）外显子区有1个SNP，可导致氨基酸由半胱氨酸（cysteine）变为色氨酸（tryptophan）。因此，认为TraesCS6A02G077000是该QTL上影响小麦品质的基因（命名为TaXip）。
为了探讨基因TaXip在小麦中的功能，在IWGSC数据库（http://plants.ensembl.org/Triticum_aestivum/Info/Index）上与TaXip的CDS进行BLAST分析，鉴定出三个同源基因，分别位于染色体6A、6B、6D，命名为TaXip-6A（TraesCS6A02G077000）、TaXip-6B（TraesCS6B02G103900）、TaXip-6D（TraesCSU02G026500）。
表3 遗传图谱（部分）
实施例2：TaXip基因的扩增和序列分析
利用Fielder分别扩增了TaXip的3个同源基因。所述引物对的序列如下：
TAXIP6A-F: ccttaggattcactcctgcg（SEQ ID No.7）；
TAXIP6A-R: gttccgagtggtgatcagc（SEQ ID No.8）；
TAXIP6B-F: gcgctagagcagaggatcctaac（SEQ ID No.9）；
TAXIP6B-R: ggcttgtggaagcatagctcc（SEQ ID No.10）；
TAXIP6D-F: gtcggatacgaattggcg（SEQ ID No.11）；
TAXIP6D-R: aactgtgcgaccaatctgttc（SEQ ID No.12）。
扩增得到TaXip-6A基因的核苷酸序列如SEQ ID No.1所示，其编码的氨基酸序列如SEQ ID No.2所示。TaXip-6B基因的核苷酸序列如SEQ ID No.3所示，其编码的氨基酸序列如SEQ ID No.4所示。TaXip-6D基因的核苷酸序列如SEQ ID No.5所示，其编码的氨基酸序列如SEQ ID No.6所示。
参见图2B和图4，TaXip-6A基因和TaXip-6D基因只有一个外显子，TaXip-6A和TaXip-6D都有915 bp的开放阅读框，编码305个氨基酸。而TaXip-6B有一个内含子、两个外显子，编码313个氨基酸。基于氨基酸序列和结构域相似性比对结果表明，TaXip-6A和TaXip-6D的同源性（94.75%）高于TaXip-6A和TaXip-6B（91.69%）、TaXip-6B和TaXip-6D（91.37%）。
对亲本泰农18和临麦6号的序列进行扩增。经序列比对发现两亲本TaXip-6A基因在390 bp外显子处有一个SNP位点（泰农18为碱基C，临麦6为碱基G），分别为半胱氨酸和色氨酸。扩增亲本泰农18和临麦6的TaXip-6B基因和TaXip-6D基因，它们外显子序列无差异。
利用"Ensemble.Plants"检索小麦的木聚糖酶抑制蛋白基因及其相似的蛋白序列、水稻中已克隆的木聚糖酶抑制蛋白序列。利用MEGA5.2软件构建系统进化树（图3）表明，所有基因可被优先化分为2个进化枝，一枝是几丁质酶，另一枝是木聚糖酶抑制蛋白。这些基因都具有保守的糖基水解酶家族18（GH18）结构域，参与碳水化合物的代谢。但是，木聚糖酶抑制蛋白不具有几丁质酶活性，不能降解几丁质。它们在进化过程中产生了新的功能。本发明中的TaXip-6A、TaXip-6B和TaXip-6D可归为XIP分支，并与XIP-I、XIP-III关系密切，说明TaXip具有木聚糖酶抑制剂的功能。·
实施例3：TaXip基因的CRISPR/Cas9基因编辑
从Fielder获得的TaXip基因的序列，用于在CRISPR-direct（http://crispr.dbcls.jp/）和CRISPOR （http://crispor.tefor.net/）中设计引导RNA（sgRNA）靶序列。
sgRNA1为：5' -ACAACATCCGCGGCGGCCCG-3 '（SEQ ID No.13），PAM序列为GGG；sgRNA2为：5' -GTCCAACCGCTCCGCGCTCG-3'（SEQ ID No.14），PAM序列为CCC。
通过对目标位点测序，将TaU3启动子调控的二元载体（图1）经农杆菌介导转化至Fielder。2019年8月，共获得54株T0植物，共获得29株编辑植株。T1、T2代在温室种植不同编辑类型。将T2代突变体植株的种子播种在花盆中（T2：3代），每个花盆4株，每个编辑类型30盆。野生型Fielder株系的籽粒也同样收获，并作为对照。
在本发明构建的CRISPR/Cas9编辑系统中，sgRNA2起主要作用，sgRNA1起次要作用。在T0代得到了29株基因编辑材料，经鉴定编辑效率为53.7%。在T2代，得到2个基因型的突变体：编辑率大于80%的A基因型和编辑率小于20%的B、D基因型归为aaBBDD基因型；编辑率小于20%的A基因型，编辑率大于80%的B、D基因型，归为AAbbdd基因型。
实施例4：TaXip-6A，TaXip-6B和TaXip-6D基因型鉴定
基因编辑植株的突变类型鉴定采用第二代测序技术进行测序鉴定。
（1）设计第一轮特异PCR引物：根据常规PCR引物设计原则设计第一轮引物（18-23nt）。引物序列为
g6aF：ggagtgagtacggtgtgcGTTGGCGGCTACGGCACC（SEQ ID No.15）；
g6aR：gagttggatgctggatggCACCGGACCGTCGCCGT（SEQ ID No.16）；
g6bF：ggagtgagtacggtgtgcATCGGCGGCTACGGCACC（SEQ ID No.17）；
g6bR：gagttggatgctggatggCACCGGACCGTCGCCGTT（SEQ ID No.18）；
g6dF：ggagtgagtacggtgtgcCATCGGCGGCTACGGCG（SEQ ID No.19）；
g6dR：gagttggatgctggatggCGGACCGTCGCCGTCAGGT（SEQ ID No.20）。
靶位点需在离正向引物或反向引物的10-100bp范围内，扩增长度在150-300bp左右。正向引物5’端加搭桥序列5’-ggagtgagtacggtgtgc-3’（SEQ ID No.21）；反向引物5’端加搭桥序列5’-gagttggatgctggatgg-3’（SEQ ID No.22）。
（2）扩增第一轮PCR目的片段：利用设计的特异引物进行第一轮常规PCR扩增条带，取3-5µL琼脂糖凝胶电泳检测 PCR 产物，确保目标产物存在且特异性良好。
（3）用Hi-TOM试剂盒进行第二轮PCR。
（4）混合样本切胶回收，网上样品登记：在网站http://www.hi-tom.net/hi-tom/注册账号，登录并录入样本信息。
检测结果见表4和图5：aaBBDD基因型6A缺失或插入了一个碱基；AAbbdd基因型6B缺失22个碱基，6D插入1个碱基。利用网站（http://www.detaibio.com/sms2/translate.html），进行蛋白质预测。我们发现以上突变均可引起氨基酸移码突变，终止密码子提前出现进而导致蛋白质失活。
表4 不同突变型的编辑类型
实施例5：TaXip基因对品质的影响
一、品质性状SV、ST的测定
面粉由Brabende Quadrumat Junior的小型实验磨研磨，然后过80目筛。
1、SDS沉降试验方法参考Pena等（1990）。
（1）将5克全麦粉加到带塞子的100 ml量筒中，加入50 ml溴酚蓝水溶液。
（2）塞好塞子后将量筒水平握在手中，左右交替摆动12次，使面粉和溶液充分混合。
（3）将具塞量筒置于机械摇床上摇混4 min。
（4）4 min末，从机械要床上取下量筒，立即加入50 ml SDS-乳酸储备溶液，置于机械摇床摇混6 min。
（5）6 min末，从机械摇床上取下量筒，立即将其竖直放置40 min。
（6）40 min末，读取沉淀值毫升数，精确至0.1 ml。
溶液配制：通过在1000毫升水中添加20克SDS，然后添加20毫升原液稀释的乳酸溶液，1份乳酸和8份体积的水来制备。
用公式
，计算修正后的面粉含水率14%的SV。
2、粉质仪参数检测参照AACC54-21标准方法，采用Brabender粉质仪测定面团稳定时间等指标。使用近红外分析仪测定不同基因型面粉的水分含量，以计算确定所需的面粉重量（田纪春，2006)。
采用SPSS17.0进行ST和SV的统计分析。采用单因素方差分析（One-way ANOVA）确定不同基因型之间的显著差异。使用LSD进行多重比较，以确定差异显著性。
经测定，aaBBDD、AAbbdd 2种突变类型和野生型Fielder的SV分别为31.77、27.30和20.08 ml。基因编辑型的SV均高于野生型对照，aaBBDD的SV显著高于AAbbdd和野生型（图2D）。结果表明，TaXip-6A、TaXip-6B和TaXip-6D对SV都有影响，但TaXip-6A对沉淀值的影响最大。
aaBBDD、AAbbdd 2种突变类型和野生型Fielder的ST分别为2.60、2.24 和2.25。突变基因型aaBBDD的ST显著高于野生型和AAbbdd，而野生型和AAbbdd之间没有显著差异（图2D）。结果说明，TaXip-6A对稳定时间影响显著，而TaXip-6B和TaXip-6D对稳定时间影响不大。
表5 突变型和野生型的面团流变学特性
以上述的结果可知，TaXip-6A基因是影响小麦籽粒品质的主效基因，所以利用该基因可以对优质小麦品种进行筛选，或者用来调控小麦的籽粒品质。
以上实施例仅用以说明本发明的技术方案，而非对其进行限制；尽管参照前述实施例对本发明进行了详细的说明，对于本领域的普通技术人员来说，依然可以对前述实施例所记载的技术方案进行修改，或者对其中部分技术特征进行等同替换；而这些修改或替换，并不使相应技术方案的本质脱离本发明所要求保护的技术方案的精神和范围。
序列表
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225 230 235 240
Glu Glu Ala Trp Asp Arg Trp Thr Ala Ala Tyr Pro Ala Thr Arg Phe
245 250 255
Phe Ile Gly Leu Thr Ala Asp Glu Lys Ser Tyr Gln Trp Ile His Pro
260 265 270
Lys Asn Val Tyr Tyr Gly Ile Thr Pro Val Val Gln Lys Lys Asp Asn
275 280 285
Tyr Gly Gly Val Met Leu Trp Asp Arg Tyr Phe Asp Lys Gln Ser Asp
290 295 300
Tyr Ser Ser Tyr Ile Lys Tyr Tyr Ala Tyr
305 310
<210> 5
<211> 919
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
atggcgccgc tcgcacccgg gaggccacca gcctgcctcc taacccttct ctccgtcgtc 60
gcggccctat ccctggccgc gccgggcctg gcggcgggga agaccggcca ggtgacggtg 120
ttctggggac ggaacaaggc cgaggggtcc ctgcgcgagg cctgcgactc cggcatgtac 180
accatggtca ccatctcttt cctcgacgtc ttcggcgcca acggaaagta ccaccttgac 240
ctctccggcc acgacctctc cgccgtcggc gccgacatca agcactgcca gtccaagggc 300
gtccccgtct ccctctccat cggcggctac ggcgcccgct actcgctccc gtccaaccgc 360
tccgcgctcg acctcttcga ccacctctgg gactcctact tcggcgggtc caagccgggc 420
gtgccccgcc ccttgggcga cgcgtggctc gacggcgtcg acctcttcct ggagcacggc 480
acgccggcgg accgctacga cgtgctggcg ctggagctgg cgaagcacaa catccgcggc 540
ggcccgggga agccgctgca cctgacggcg acggtccggt gcgggtaccc gccggcggcg 600
cacgtggggc gagcgctggc gacggggatc ttcgagcgcg tccacgtgag gatctacgag 660
gagagcgaca aggcgtgcaa ccagtacggg gcgtgggagg aggcgtggga caggtggacg 720
gcggcgtacc cggccacccg gttcttcatc gggctcacgg cggacgacaa gtcgtaccag 780
tggatacacc ccaagaacgt ctactacggc atcacgccgg tggtgcagaa gaaggagaac 840
tatggcgggg tcatgctctg ggaccgatac ttcgacaagc agagcgacta cagtagctac 900
atcaagtact acgcctgaa 919
<210> 6
<211> 306
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 6
Met Ala Pro Leu Ala Pro Gly Arg Pro Pro Ala Cys Leu Leu Thr Leu
1 5 10 15
Leu Ser Val Val Ala Ala Leu Ser Leu Ala Ala Pro Gly Leu Ala Ala
20 25 30
Gly Lys Thr Gly Gln Val Thr Val Phe Trp Gly Arg Asn Lys Ala Glu
35 40 45
Gly Ser Leu Arg Glu Ala Cys Asp Ser Gly Met Tyr Thr Met Val Thr
50 55 60
Ile Ser Phe Leu Asp Val Phe Gly Ala Asn Gly Lys Tyr His Leu Asp
65 70 75 80
Leu Ser Gly His Asp Leu Ser Ala Val Gly Ala Asp Ile Lys His Cys
85 90 95
Gln Ser Lys Gly Val Pro Val Ser Leu Ser Ile Gly Gly Tyr Gly Ala
100 105 110
Arg Tyr Ser Leu Pro Ser Asn Arg Ser Ala Leu Asp Leu Phe Asp His
115 120 125
Leu Trp Asp Ser Tyr Phe Gly Gly Ser Lys Pro Gly Val Pro Arg Pro
130 135 140
Leu Gly Asp Ala Trp Leu Asp Gly Val Asp Leu Phe Leu Glu His Gly
145 150 155 160
Thr Pro Ala Asp Arg Tyr Asp Val Leu Ala Leu Glu Leu Ala Lys His
165 170 175
Asn Ile Arg Gly Gly Pro Gly Lys Pro Leu His Leu Thr Ala Thr Val
180 185 190
Arg Cys Gly Tyr Pro Pro Ala Ala His Val Gly Arg Ala Leu Ala Thr
195 200 205
Gly Ile Phe Glu Arg Val His Val Arg Ile Tyr Glu Glu Ser Asp Lys
210 215 220
Ala Cys Asn Gln Tyr Gly Ala Trp Glu Glu Ala Trp Asp Arg Trp Thr
225 230 235 240
Ala Ala Tyr Pro Ala Thr Arg Phe Phe Ile Gly Leu Thr Ala Asp Asp
245 250 255
Lys Ser Tyr Gln Trp Ile His Pro Lys Asn Val Tyr Tyr Gly Ile Thr
260 265 270
Pro Val Val Gln Lys Lys Glu Asn Tyr Gly Gly Val Met Leu Trp Asp
275 280 285
Arg Tyr Phe Asp Lys Gln Ser Asp Tyr Ser Ser Tyr Ile Lys Tyr Tyr
290 295 300
Ala Tyr
305
<210> 7
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
ccttaggatt cactcctgcg 20
<210> 8
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
gttccgagtg gtgatcagc 19
<210> 9
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
gcgctagagc agaggatcct aac 23
<210> 10
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
ggcttgtgga agcatagctc c 21
<210> 11
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
gtcggatacg aattggcg 18
<210> 12
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
aactgtgcga ccaatctgtt c 21
<210> 13
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 13
acaacatccg cggcggcccg 20
<210> 14
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 14
gtccaaccgc tccgcgctcg 20
<210> 15
<211> 36
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 15
ggagtgagta cggtgtgcgt tggcggctac ggcacc 36
<210> 16
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 16
gagttggatg ctggatggca ccggaccgtc gccgt 35
<210> 17
<211> 36
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 17
ggagtgagta cggtgtgcat cggcggctac ggcacc 36
<210> 18
<211> 36
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 18
gagttggatg ctggatggca ccggaccgtc gccgtt 36
<210> 19
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 19
ggagtgagta cggtgtgcca tcggcggcta cggcg 35
<210> 20
<211> 37
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 20
gagttggatg ctggatggcg gaccgtcgcc gtcaggt 37
<210> 21
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 21
ggagtgagta cggtgtgc 18
<210> 22
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 22
gagttggatg ctggatgg 18
Claims (10)
1.一种影响小麦籽粒品质的基因TaXip，其特征在于，所述基因TaXip包括基因TaXip- 6A、基因TaXip-6B和基因TaXip-6D。
2.根据权利要求1所述的影响小麦籽粒品质的基因TaXip，其特征在于，所述基因TaXip-6A的核苷酸序列如SEQ ID No.1所示，其编码的氨基酸序列如SEQ ID No.2所示。
3.根据权利要求1所述的影响小麦籽粒品质的基因TaXip，其特征在于，所述基因TaXip-6B的核苷酸序列如SEQ ID No.3所示，其编码的氨基酸序列如SEQ ID No.4所示。
4.根据权利要求1所述的影响小麦籽粒品质的基因TaXip，其特征在于，所述基因TaXip-6D的核苷酸序列如SEQ ID No.5所示，其编码的氨基酸序列如SEQ ID No.6所示。
5.权利要求1-4任一项所述的影响小麦籽粒品质的基因TaXip的检测引物，其特征在于，所述检测引物的序列如SEQ ID No.7~ SEQ ID No.12所示。
6.一种小麦籽粒品质的鉴定方法，其特征在于，所述鉴定方法为：
（1）提取小麦籽粒的DNA；
（2）利用序列如SEQ ID No.7~ SEQ ID No.12所示的检测引物对DNA进行扩增；
（3）扩增结果进行比对，如果序列中出现多态性突变，则样品小麦籽粒品质存在差异。
7.根据权利要求6所述的小麦籽粒品质的鉴定方法，其特征在于，所述步骤（3）中多态性突变为C/G突变，其发生在位于小麦染色体6A上的基因TaXip-6A上；所述C/G突变中G碱基变异能够提高小麦籽粒的SDS沉降值和稳定时间。
8.权利要求1-4任一项所述的影响小麦籽粒品质的基因TaXip在用于调控小麦籽粒品质性状中的应用。
9.根据权利要求8所述的应用，其特征在于，所述应用方法为：通过敲除基因TaXip来提高小麦籽粒的SDS沉降值和稳定时间。
10.权利要求5所述的影响小麦籽粒品质的基因TaXip的检测引物在用于选育高品质小麦籽粒的应用。
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Cited By (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN115011617A (zh) * | 2022-05-16 | 2022-09-06 | 山东农业大学 | 一种控制小麦株高的主效qtl及其候选基因和应用 |
Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| US8173866B1 (en) * | 2008-01-11 | 2012-05-08 | Pioneer Hi-Bred International, Inc. | Modulation of plant xylan synthases |
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2021
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Patent Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| US8173866B1 (en) * | 2008-01-11 | 2012-05-08 | Pioneer Hi-Bred International, Inc. | Modulation of plant xylan synthases |
Non-Patent Citations (7)
| Title |
|---|
| W. DEBYSER ET AL.: "Triticum aestivum Xylanase Inhibitor (TAXI), a New Class of Enzyme Inhibitor Affecting Breadmaking Performance", 《JOURNAL OF CEREAL SCIENCE》, pages 39 * |
| 无: "hypothetical protein CFC21_086812 [Triticum aestivum] Accession NO. KAF7082986.1", 《GENBANK DATABASE》 * |
| 无: "PREDICTED: Triticum aestivum xylanase inhibitor protein 1-like (LOC123141024), mRNA", 《GENBANK DATABASE》 * |
| 无: "PREDICTED: Triticum dicoccoides xylanase inhibitor protein 1-like (LOC119314698), mRNA", 《GENBANK DATABASE》 * |
| 无: "Triticum aestivum xylanase inhibitor protein 1-like (LOC123133608), mRNA", 《GENBANK DATABASE》 * |
| 无: "xylanase inhibitor protein 1-like [Triticum aestivum]，Accession NO. XP_044416208.1", 《GENBANK DATABASE》 * |
| 马名章;侯春晓;王谦;张传亮;刘建新;翁晓燕;: "谷物中蛋白类木聚糖酶抑制剂研究进展", 中国生物工程杂志, no. 04, pages 129 - 133 * |
Cited By (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN115011617A (zh) * | 2022-05-16 | 2022-09-06 | 山东农业大学 | 一种控制小麦株高的主效qtl及其候选基因和应用 |
| CN115011617B (zh) * | 2022-05-16 | 2023-11-14 | 山东农业大学 | 一种控制小麦株高的主效qtl及其候选基因和应用 |
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