CN114634560B - 棉花GhIQD21基因序列及其克隆与应用 - Google Patents
棉花GhIQD21基因序列及其克隆与应用 Download PDFInfo
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Abstract
本发明涉及棉花GhIQD21基因序列及其克隆与应用，属于分子生物学和生物技术领域。GhIQD21基因能重塑拟南芥器官形态，且获得的转基因植株均能正常开花结果，转化基因能稳定遗传至下一代，故该基因可以进一步运用于园艺观赏植物的塑型，培育出姿态各异的新品种，为园艺花卉型态塑造提供新途径；本发明为植物器官形态研究和细胞微管骨架调控提供理论依据；本发明中的转基因拟南芥将成为植物细胞微管骨架研究的有用材料。
Description
技术领域
本发明涉及棉花GhIQD21基因序列及其克隆与应用，属于分子生物学和生物技术领域。
背景技术
微管骨架具有高度的动态特性，其排列方式不断的进行活跃的重组，以响应生长发育及外界生物和非生物刺激信号。微管骨架动态受到精细而复杂的调控，其中微管结合蛋白在微管动态调控上起着关键作用。目前，在植物中已鉴定多种微管结合蛋白如IQD蛋白，都在植物生长发育过程中起重要的作用。IQD蛋白具有保守的67个氨基酸的中心基序，即IQ67保守结构域。IQD蛋白家族目前在多个物种基因组(拟南芥，水稻，番茄，二穗短柄草，大豆，毛果杨，毛竹，玉米和棉花)中进行了全基因组的鉴定和分析，特别是在陆地棉基因组中鉴定到148个 IQD基因，在所有已鉴定的物种中数目最多。
通过对一些IQD蛋白功能的研究，揭示了IQD蛋白在拟南芥叶片细胞形态、水稻籽粒大小和番茄果实形状调控上具有重要的作用。在拟南芥中，AtIQD5主要在营养器官表达，该基因功能缺失后，拟南芥叶片表皮细胞形态改变；IQD11、IQD14和IQD16的异位过表达，改变叶片形态和细胞形状。在水稻中，GRAIN SIZE ONCHROMOSOME 5(GSE5)/IQD21基因功能缺失后，水稻籽粒变宽，相反， GSE5基因过表达后导致水稻籽粒变窄，GSE5基因主要通过影响小穗颖壳细胞的增殖调控籽粒宽度，该基因启动子区的自然变异导致了水稻籽粒大小多样性。在番茄中，SUN/IQD12通过控制细胞分化而调控果实形状，该基因由长末端重复反转座子导致24.7kb的基因复制事件，该事件引起SUN表达量增加，最终形成伸长的果实形状。综上所述，IQD蛋白在调控植物器官发育中起重要作用。
发明内容
本发明首次从棉花中分离出编码含IQ67结构域的GhIQD21蛋白的全长 cDNA，其具有SEQ ID No：1所示的核苷酸序列。
作为本发明基因的改进：上述核苷酸序列还包括在SEQ ID No：1所示的核苷酸序列中添加、取代、插入或缺失一个或多个核苷酸生成的突变体、等位基因或衍生物。
本发明提供一种棉花微管结合蛋白基因GhIQD21编码的蛋白质，其具有SEQ IDNO：2所示的氨基酸序列。
作为本发明蛋白质的改进：上述氨基酸序列还包括在SEQ ID No：2所示的氨基酸序列中添加、取代、插入或缺失一个或多个氨基酸生成的衍生物。
本发明还同时提供了含有上述基因的质粒以及含有上述基因的植物表达载体。
本发明还同时提供了一种宿主细胞，该宿主细胞含有上述基因序列。该宿主细胞为大肠杆菌细胞、农杆菌细胞或植物细胞。
本发明还同时提供了改良拟南芥器官发育的方法：包括用具有SEQ ID No： 1所示的核苷酸序列的基因转化拟南芥。
进一步具体说明：将该基因连接到表达载体上，利用农杆菌浸花法侵染拟南芥，获得的转基因拟南芥本发明的目的是提供一种从棉花中克隆的微管结合蛋白编码新基因GhIQD21，如SEQ ID No：1所示的DNA序列，也包括编码GhIQD21 蛋白的编码区序列(SEQ IDNo：2)。
本发明中的SEQ ID No：2所示的蛋白质属于一类表达蛋白，其中进行一个或几个氨基酸的替换、插入或缺失所获得的功能类似物。另外，也包括在SEQ ID No：1中添加、取代、插入或缺失一个或多个核苷酸而生成的突变体、等位基因或衍生物，具有相同功能的序列也能达到本发明的目的。
本发明的另一个目的是提供一种用GhIQD21基因进行高效的植物转化的方法，具体地说，本发明提供了具有SEQ ID No：1所示的序列基因的载体 pEarleyGate 101-GhIQD21，该载体可以表达有上述核苷酸序列编码的多肽或其同源类似物。
本发明还提供了一种利用植物表达载体转化植物细胞影响拟南芥器官发育的方法。
本发明的有益效果：
1、GhIQD21基因能重塑拟南芥器官形态，且获得的转基因植株均能正常开花结果，转化基因能稳定遗传至下一代，故该基因可以进一步运用于园艺观赏植物的塑型，培育出姿态各异的新品种，为园艺花卉型态塑造提供新途径；
2、本发明为植物器官形态研究和细胞微管骨架调控提供理论依据；
3、本发明中的转基因拟南芥将成为植物细胞微管骨架研究的有用材料。
附图说明
图1为本发明棉花GhIQD21基因序列及其克隆与应用的蛋白特征示意图。
图2为本发明棉花GhIQD21基因序列及其克隆与应用的蛋白亚细胞定位示意图。
图3为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥植株鉴定和子叶表型示意图。
图4为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥植株莲座叶和细胞表型示意图。
图5为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥植株花和细胞表型示意图。
图6为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥植株茎秆和细胞表型示意图。
图7为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥植株果荚和细胞表型示意图。
图8为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥下胚轴oryzalin处理表型示意图。
图9为本发明棉花GhIQD21基因序列及其克隆与应用的基因拟南芥下胚轴和子叶微管变化示意图。
具体实施方式
下面将结合附图，对本发明的优选实施例进行详细的描述。
实施例1
实施例1：棉花GhIQD21基因的克隆和表达载体构建
(1)引物设计
根据cottonFGD已报道的棉花基因组序列，通过生物信息学比对，确定GhIQD21 基因cDNA序列，根据该序列设计编码框(ORF)全长序列的引物，引物序列如下：
IQD-F:5'GGGGACAAGTTTGTACAAAAAAGCAGGCTTAATGGGAAAGAAAG GGAGTGG3'
IQD-R:5'GGGGACCACTTTGTACAAGAAAGCTGGGTACCAGCCATGACTGCC AATAG3'
(2)陆地棉TM-1棉花叶片总RNA提取
取幼嫩叶片加液氮研磨至粉末状，然后使用试剂盒RNAprep Pure Plant Kit (天根，中国)提取叶片总RNA。经分光光度计检测RNA浓度及质量，浓度约为600ng/ul，OD260/OD280＝2.0，OD260/OD230＝2.0，提取的总RNA立即用于下一步实验。
(3)RNA反转录
取1μg RNA至0.2ml无RNAase的离心管中，使用试剂盒(Takala，日本) 将RNA反转录为cDNA，经分光光度计检测DNA浓度及质量，浓度约为 1000ng/μl，OD260/OD280＝1.8，OD260/OD230＝2.0，得到的cDNA下一步实验。
(4)PCR扩增和纯化
使用高保真酶进行PCR扩增，具体如下：PCR反应体系为：1000ng/μl棉花 cDNA 1μl,10uM引物F和R向各2μl,2×Phanta Master Mix：25μl，ddH2O 20μl, 总体系为50μl。PCR扩增条件具体为：95℃预变性2分钟；95℃变性15秒，55℃退火15秒，72℃延伸40秒，35个循环；1％琼脂糖凝胶电泳分离和Gelred染色检测扩增结果。使用Omega PCR纯化回收试剂盒，进行PCR纯化回收，浓度达到100ng/μl，进行连接反应。
(5)PCR片段连接表达载体
采用Gateway系统进行表达载体构建，首先将PCR产物与入门载体 pDONAR-Zeo进行BP反应，获得pDONAR-Zeo-GhIQD21入门载体。将入门载体送至上海生工生物工程有限公司进行测序，载体中包含SEQ ID No：1所示的核苷酸序列，其蛋白质序列中含有典型的IQ67结构域。然后将入门载体与目的载体pEarleyGate 101进行LR反应，得到过表达载体pEarleyGate 101-GhIQD21。将pEarleyGate 101-GhIQD21载体转化至GV3101农杆菌中，利用PCR鉴定阳性菌落。
(6)烟草瞬时表达
将本氏烟烟草在16h光照，8h黑暗，24℃的温室中培养4-5周后，将含 pEarleyGate101-GhIQD21载体和YFP空载对照的GV3101农杆菌摇菌后，用 20mM MES，150nMAS，10mMMgCl2的溶液重悬，OD600值调至1.0左右，黑暗放置3小时后，将农杆菌注射到烟草叶片中。培养36小时左右，用奥林巴斯 FV1200(奥林巴斯，日本东京)采集叶片的EYFP信号，同时用20μM微管解聚剂oryzalin进行处理2小时，采集EYFP信号。可以观察到GhIQD21-YFP荧光信号呈现丝状分布，并且可以被oryzalin降解，说明GhIQD21定位于微管骨架。
实施例2：转基因拟南芥的获得
(1)拟南芥转化
将野生型拟南芥Col0放入4℃冰箱中进行春化处理4天，营养土和蛭石按 3:1搅拌均匀后，加水润湿后分装至小盆中，将Col0种子直接用枪头吸至小盆中，每盆9棵。拟南芥放置于16h光照，8h黑暗，24℃的温室中进行培养。培养45 天左右，花苞较多时进行转化。取pEarleyGate 101-GhIQD21阳性农杆菌200μl 于2ml LB+rif+kan培养基中，28℃，200rpm振荡培养过夜，取1ml菌液于100ml 相应的培养基中扩大培养，28℃，200rpm振荡培养12小时，4000rpm，10min 收集菌液，使用侵染液(1/2ms+5％蔗糖+0.05％的silwetl-77)重新悬浮菌液，将 OD600调至1.5左右。将配置好的侵染液倒置培养皿中，使拟南芥花絮浸泡于侵染液中1min，转化完成的拟南芥植株横放，保鲜膜覆盖进行保湿，暗处理24小时后放置温室培养，拟南芥结实后，收取种子，进行抗性筛选转基因阳性植株。
(2)阳性植株的筛选和鉴定
将获得的转基因拟南芥T0代种子均匀的撒播在营养土基质中，待种子发芽后，开始喷施Basta除草剂，每隔5天左右喷施一次，共喷施3次。总共得到8 株抗性植株，利用CTAB法提取每一株抗性拟南芥基因组DNA，并进行目的基因PCR鉴定，经鉴定均为阳性植株，分别命名为OE-1～8。由于我们将GhIQD21 与YFP进行了融合表达，所以进一步用anti-GFP抗体对OE-1～3进行western blot 检测，结果发现三个株系的蛋白均有表达。同时，以空载转基因株系为对照，观察了拟南芥幼苗的表型，可以看出过表达株系比对照材料的子叶更小，叶柄短粗。
实施例3：转基因拟南芥的器官形态变化
转基因T3代纯合株系，在16h光照，8h黑暗，24℃的温室中培养27天后，观察转基因株系莲座叶表型，可以观察到三个过表达转基因株系OE-1～3的叶片均比对照小，叶柄更加短粗。进一步对第7片莲座叶进行了细胞学观察，发现 OE-1～3叶片表皮的扁平细胞变小。植株生长40天，观察拟南芥花的表型，可以看到OE-1～3的花瓣比对照长度变短，宽度变宽，但是花瓣的大小变化不明显。植株生长45天，观察拟南芥茎秆的表型，可以看出OE-1～3的茎秆比对照更粗，而且对照茎秆上表皮毛大多数是二叉的，而OE-1～3的表皮毛是单叉的。(通过石蜡切片，我们看出OE-1～3茎秆的细胞数目明显比对照增多。植株生长50天，观察拟南芥果荚的表型，可以看出OE-1～3的果荚比对照更加的短粗，通过扫描电镜，观察了细胞形态变化，发现OE-1～3的细胞变短，而且细胞形状更加的不规则。
实施例4：转基因拟南芥的微管变化
为了分析转基因拟南芥的微管变化，首先分析了OE-1～3转基因株系和对照对微管解聚剂oryzalin的反应。转基因纯合系和对照拟南芥种子经消毒，4℃春化，播种至含0和300nM oryzalin 1/2MS固体培养基，黑暗培养4天后，记录下胚轴长度和宽度。可以看出，OE-1～3转基因株系的下胚轴比对照更加的短粗，对微管解聚剂更加敏感。
以TUB6-mCherry作为对照，利用激光共聚焦观察了OE-1下胚轴和子叶中微管的变化。在下胚轴中，对照中微管是很长的细丝，大部分微管都具有相同的方向和近似平行的排列，微管束较粗，而且光滑。过表达OE-1的微管没有长而完整的丝状，并且没有明确的排列方向和平行特征，微管束不光滑。在子叶中，观察到相似的微管表型变化。
以上对本发明及其实施方式进行了描述，这种描述没有限制性，附图中所示的也只是本发明的实施方式之一，实际的结构并不局限于此。总而言之如果本领域的普通技术人员受其启示，在不脱离本发明创造宗旨的情况下，不经创造性的设计出与该技术方案相似的结构方式及实施例，均应属于本发明的保护范围。
序列表
<110> 中国农业科学院棉花研究所
<120> 棉花GhIQD21基因序列及其克隆与应用
<130> 2022.3.8
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1353
<212> DNA
<213> 人工序列(Manual sequence)
<400> 1
atgggaaaga aagggagtgg atggttctcg acggtgaaga aagtgttcaa atcatctcac 60
aaagacttgc ctgacaaaaa aagagagaat aatgtggaga aatggccaaa tgaggccccg 120
gatgcagtgt cgtttgaaca ttttccagcg gagagttcac cggacatcac gaataatgag 180
agtgcaatgt cgactcctct gaatgaagat aggaaccatg ccattgctgt cgccatggca 240
actgctgctg ctgcagaagc agctgttgcg gcagctcaag ctgccgctaa agtcgtccgg 300
ttggcgggtt atggacgaca gtctaaggaa gaaagagctg caacgcctat acagtcgtat 360
tatagaggct atctggctcg gagagctttg cgtgctttga agggactggt gaggttgcag 420
gcactagtga gaggttataa cgtgcgcaaa caagcgcaaa tgacaatgcg gtgcatgcaa 480
gcattagtcc gggtgcaggc aagagttcaa gctcgtaggc tgcagctaac tgatcatgat 540
aagatcccca agagatcaga cgaggactat gaagaaaaaa aaggaacggg agaagtagaa 600
cgaaagccaa ggagcctgtt gaagagatat gacagttggg atggtggaca gcaaagctcg 660
gagaaggtta aggaaagtgt ttcgaagaaa catgatgctg cgatgagacg agaacgagcc 720
cttgcttatg cctatagcta tcagcaacag ccacagcaac aactcatgca accgcacccc 780
aatggcaaag atgttggagt ccaccttaat gatcgcgaga aggcacaatg gggttggaac 840
tggctcgagc attggatgtc atcgcaaccg tacgacgctc gtcaactagg ccatcaagaa 900
ggttcttaca tgacactgct gaccaccacc gcaaccacta ccgttaccga taacatgtct 960
gagaaaactg ttgaaatgga tgttgtgaca gcaatggatt caggctcata ctcaaccctg 1020
cagcaggctc aatcagggtc aagtaatgta cctagctaca tggccccaac ccaatctgcc 1080
aaagccaagg ttcgaagcca aggacctatg attatgaagc agcaacgtgt accacatgca 1140
ccccaatgga acccatcaac aaaaaaatcc ccaggttgtg attcatcaag ttcaggtgga 1200
ggaacaacag tctaccaggc accaaggagc cctggccaaa agagtaacgg tgctcgcata 1260
ccgtctaggc gtctaggtgg atgtagcccg gataccgctg gtggccacgg tggagaagat 1320
tggaggttgc ctattggcag tcatggctgg tga 1353
<210> 2
<211> 450
<212> PRT
<213> 人工序列(Manual sequence)
<400> 2
Met Gly Lys Lys Gly Ser Gly Trp Phe Ser Thr Val Lys Lys Val Phe
1 5 10 15
Lys Ser Ser His Lys Asp Leu Pro Asp Lys Lys Arg Glu Asn Asn Val
20 25 30
Glu Lys Trp Pro Asn Glu Ala Pro Asp Ala Val Ser Phe Glu His Phe
35 40 45
Pro Ala Glu Ser Ser Pro Asp Ile Thr Asn Asn Glu Ser Ala Met Ser
50 55 60
Thr Pro Leu Asn Glu Asp Arg Asn His Ala Ile Ala Val Ala Met Ala
65 70 75 80
Thr Ala Ala Ala Ala Glu Ala Ala Val Ala Ala Ala Gln Ala Ala Ala
85 90 95
Lys Val Val Arg Leu Ala Gly Tyr Gly Arg Gln Ser Lys Glu Glu Arg
100 105 110
Ala Ala Thr Pro Ile Gln Ser Tyr Tyr Arg Gly Tyr Leu Ala Arg Arg
115 120 125
Ala Leu Arg Ala Leu Lys Gly Leu Val Arg Leu Gln Ala Leu Val Arg
130 135 140
Gly Tyr Asn Val Arg Lys Gln Ala Gln Met Thr Met Arg Cys Met Gln
145 150 155 160
Ala Leu Val Arg Val Gln Ala Arg Val Gln Ala Arg Arg Leu Gln Leu
165 170 175
Thr Asp His Asp Lys Ile Pro Lys Arg Ser Asp Glu Asp Tyr Glu Glu
180 185 190
Lys Lys Gly Thr Gly Glu Val Glu Arg Lys Pro Arg Ser Leu Leu Lys
195 200 205
Arg Tyr Asp Ser Trp Asp Gly Gly Gln Gln Ser Ser Glu Lys Val Lys
210 215 220
Glu Ser Val Ser Lys Lys His Asp Ala Ala Met Arg Arg Glu Arg Ala
225 230 235 240
Leu Ala Tyr Ala Tyr Ser Tyr Gln Gln Gln Pro Gln Gln Gln Leu Met
245 250 255
Gln Pro His Pro Asn Gly Lys Asp Val Gly Val His Leu Asn Asp Arg
260 265 270
Glu Lys Ala Gln Trp Gly Trp Asn Trp Leu Glu His Trp Met Ser Ser
275 280 285
Gln Pro Tyr Asp Ala Arg Gln Leu Gly His Gln Glu Gly Ser Tyr Met
290 295 300
Thr Leu Leu Thr Thr Thr Ala Thr Thr Thr Val Thr Asp Asn Met Ser
305 310 315 320
Glu Lys Thr Val Glu Met Asp Val Val Thr Ala Met Asp Ser Gly Ser
325 330 335
Tyr Ser Thr Leu Gln Gln Ala Gln Ser Gly Ser Ser Asn Val Pro Ser
340 345 350
Tyr Met Ala Pro Thr Gln Ser Ala Lys Ala Lys Val Arg Ser Gln Gly
355 360 365
Pro Met Ile Met Lys Gln Gln Arg Val Pro His Ala Pro Gln Trp Asn
370 375 380
Pro Ser Thr Lys Lys Ser Pro Gly Cys Asp Ser Ser Ser Ser Gly Gly
385 390 395 400
Gly Thr Thr Val Tyr Gln Ala Pro Arg Ser Pro Gly Gln Lys Ser Asn
405 410 415
Gly Ala Arg Ile Pro Ser Arg Arg Leu Gly Gly Cys Ser Pro Asp Thr
420 425 430
Ala Gly Gly His Gly Gly Glu Asp Trp Arg Leu Pro Ile Gly Ser His
435 440 445
Gly Trp
450
Claims (2)
1.棉花GhIQD21基因序列的用途，其特征在于：基因具有Seq ID No：1所示的核苷酸序列，用于构建转基因植物，所述转基因植物的器官形态得以改良。
2.一种改良植物器官形态的方法，其特征在于：包括用具有Seq ID No：1所示的核苷酸序列的基因转化植物细胞，再将转化后的植物细胞培育成植株。
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