CN112210560A - 一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用 - Google Patents
一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用 Download PDFInfo
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Abstract
本发明公开了一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用。该基因的核苷酸序列如SEQ ID NO.1所示,该基因在编码区第十三个外显子上发生一个单碱基突变,突变位点为G/A;并设计了用于检测与该基因连锁的InDel分子标记的引物对,引物对的具体序列如SEQ ID NO.3和4所示。本发明InDel分子标记与玉米叶片黄化调控基因oy2连锁紧密,可用于鉴定植株中是否含有玉米叶片黄化调控突变基因oy2,检测结果可靠、简单易行。
Description
技术领域
本发明属于分子遗传学领域,具体涉及一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用。
背景技术
玉米是全球种植范围最广、产量最大的谷类作物,居三大粮食作物(玉米、小麦、水稻)之首。我国是玉米生产和消费大国,播种面积、总产量、消费量仅次于美国,均居世界第二位(李少昆等,2017)。但我国玉米单产远不及美国,因此,在有限的土地上提高玉米的产量极为重要,玉米的高产稳产对保障我国粮食安全具有极其重要的意义。
叶片是植物进行光合作用的主要场所,对干物质积累有重要影响,因此提高作物产量的关键是提高叶片的光能利用率,玉米叶色发生变化大多情况下会影响玉米的产量。研究表明,叶绿体是高等植物进行光合作用的主要细胞器。光合色素作为吸收光能的重要色素在光合反应中必不可少,主要包括叶绿素和类胡萝卜素和藻胆素三种类型,其中叶绿素是最重要的色素,导致叶色变化的主要原因是植物体内叶绿素合成、降解或叶绿体发育相关基因的发生突变,影响了叶绿素的生物合成和降解过程,导致叶片中叶绿素的含量发生变化,最终导致了叶色的变化。玉米叶色突变体表型明显,易于观察,已广泛应用于生产实践和科学研究中。除作为性状标记应用于新品种培育外,叶色突变体对于研究植物叶绿体发育、光形态建成等方面具有重要的作用。
基于玉米数据库(MaizeGDB)的结果显示,迄今为止已有大约200余个玉米叶色突变体基因被定位和克隆,大部分主要通过参与叶绿素的生物合成以及影响叶绿体的发育两个途径来调控叶色变异。利用玉米叶色突变体,部分叶绿素生物合成和参与叶绿体发育相关的基因也相继被克隆,包括ygl-1(Guan et al.2016)、yglm1(Wang et al.2014)、ygl3(Du et al.2018)、v-1(Miao et al.2016)、elm1与elm2(Shi et al.2013;Sawers etal.2004)、vyl-Chr.1与vyl-Chr.9(Zhang et al.2006)、tdy1与dty2(Braun et al.2006;Baker and Braun 2008)、zb7(Lu et al.2012)和SN62(Zhong et al.2015)等。其中,玉米中Oy1基因编码叶绿素生物合成过程中镁离子螯合酶亚基,该基因功能失活会导致玉米苗期叶片黄化,从而出现苗期黄化致死表型(Sawers et al.2006);玉米hcf60-ml突变体编码一个叶绿体核糖体蛋白基因,它的突变导致叶绿体核糖体的大小亚基含量显著降低,使植物苗期白化表型(Schulters et al.2000)。尽管已有众多基因被克隆,但植物叶绿体的生物合成和发育是一个及其复杂的过程,受到核基因和自身质体基因组的共同调控,对玉米叶色调控基因的获得及对其调控机理的研究任重而道远。
由于玉米种质资源匮乏和自交系遗传基础狭窄,通过人工诱变方式获得突变体已被视作高效获得功能基因的方式,突变体作为一种遗传学研究的基本材料,是研究基因的生物学功能的理想材料。因此发掘新的玉米叶色突变体,克隆其调控基因对揭示叶绿素生物合成和叶绿体发育的分子机理以及玉米高光效育种实践具有重要的意义。
发明内容
针对现有技术中的上述不足,本发明提供一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用,用本发明分子标记对带有叶片黄化基因oy2的材料进行鉴定,可以提高玉米黄化调控性状的鉴定效率,作为遗传标记应用于玉米分子育种等。
为实现上述目的,本发明解决其技术问题所采用的技术方案是:
一种玉米叶黄化基因oy2,该基因的核苷酸序列如SEQ ID NO.1所示。
进一步地,与野生型相比,该基因在编码区第十三个外显子上发生一个单碱基突变(由G突变成A)。
进一步地,基因编码的蛋白质的氨基酸序列如SEQ ID NO.2所示。
一种与基因oy2连锁的InDel分子标记,运用图位克隆技术,在玉米第5号染色体上鉴定到一个InDel位点(Maize B73 RefGen_v4,Chr5:3207013),该基因编码镁离子螯合酶亚基,其核苷酸序列如SEQ ID NO.5所示。
一种用于扩增上述InDel分子标记的引物对,该引物对的具体序列如下:
InDel-F:GAATTGGTTTGCGCCACAGT(SEQ ID No.3);
InDel-R:TGTTTATTCTTCAAGGCTGTCAGA(SEQ ID No.4)。
该InDel分子标记引物可用来鉴定玉米叶片黄化调控基因oy2。
一种筛选黄化叶玉米的方法,其特征在于,包括以下步骤:
(1)提取待测玉米DNA;
(2)采用上述引物对待测基因进行PCR扩增;
(3)对扩增产物进行测序分析,如果所得扩增产物在该位点的核苷酸碱基为G/G,待测玉米具有OY2/OY2基因型且叶片正常绿色材料;若该位点的核苷酸碱基为A/A,则待测玉米材料为具有oy2/oy2基因型且叶片黄化材料;若该位点的核苷酸碱基为G/A,则待测玉米材料为具有OY2/oy2杂合基因型且叶片正常绿色材料。
进一步地,PCR程序为:94℃预变性2min;98℃变性10s;60℃退火30s;68℃延伸5s;共35个循环;68℃再延伸10min。
进一步地,PCR反应体系为:Taq Mix 7.5μL,上下游引物分别为0.75μL,DNA模板1μL,用双蒸水定容至15μL。
一种筛选黄化叶玉米的试剂盒,包括上述分子标记和/或引物对。
一种玉米全基因组芯片,包括上述分子标记。
上述分子标记或引物对在玉米分子育种、种质资源改良等中的应用。
本发明的有益效果为:
1、本发明InDel分子标记与玉米叶片黄化调控基因oy2连锁紧密,可用于鉴定植株中是否含有玉米叶片黄化调控突变基因oy2,检测结果可靠、简单易行。
2、用本发明分子标记对带有叶片黄化基因oy2的材料进行鉴定,可以提高玉米黄化调控性状的鉴定效率,作为遗传标记应用于玉米分子育种,具有较高的应用价值。
附图说明
图1为玉米叶片黄化突变体oy2和野生型植株RP125对比图;
图2为玉米叶片斑黄化突变体oy2和野生型RP125显微结构透射电镜图;
图3为玉米叶色调控突变基因oy2的图位克隆示意图;
图4为玉米黄化叶色调控基因oy2的InDel分子标记在野生型和突变体oy2中的测序结果。
具体实施方式
下面对本发明的具体实施方式进行描述,以便于本技术领域的技术人员理解本发明,但应该清楚,本发明不限于具体实施方式的范围,对本技术领域的普通技术人员来讲,只要各种变化在所附的权利要求限定和确定的本发明的精神和范围内,这些变化是显而易见的,一切利用本发明构思的发明创造均在保护之列。
实施例1玉米黄化叶色突变体基因oy2的图位克隆
1、本实验中所用的玉米叶片黄化突变体材料来源于四川农业大学玉米研究所,系玉米研究所玉米自交系RP125经过EMS诱变处理得到(见图1),相比野生型,突变体oy2叶片呈现黄化表型。图1中,(a)为玉米两叶一心时期玉米叶片;(b)为玉米四叶一心时期玉米叶片。标尺为2cm。
突变体植株从幼苗第一片完全展开叶开始就表现出明显的黄化表型,到三叶一心至四叶一心时期,黄化苗逐渐枯萎凋亡。发明人将该突变体命名为oy2。
以上述苗期叶片黄化突变体oy2为母本,以自交系B73为父本杂交,得到F1代;F1代自交得到F2代,即为性状分离群体,经遗传分析表明该突变体黄化表型由单隐性核基因控制。鉴定F2代分离群体中叶片黄化苗的叶片基因组DNA,用于基因定位。玉米叶片基因组DNA采用CTAB法提取。
2、多态性引物筛选及目标基因初定位
前期通过筛选突变体oy2与B73之间具有多态性的SSR标记,将得到多态SSR标记对F2代分离群体中的野生型混合池(20株叶片正常绿色植株的混合DNA)和突变型混合池(20株叶片黄化植株的混合DNA)进行PCR分析,以及对126个F2代分离群体中黄化突变体单株进行PCR,PCR产物经过聚丙烯酰胺凝胶电泳之后,读取条带进行基因的连锁分析,初步将oy2基因定位于第5染色体上,并与SSR标记umc1766紧密连锁(图3)。
3、oy2基因的精细定位
为进一步精细定位oy2基因,首先进一步扩大F2定位群体到304株叶片黄化突变单株。其次根据已公开的B73与Mo17之间的插入(insertion)和删除(deletion)多态标记,设计并筛选到10对多态InDel标记,通过多态性标记对304株叶片黄化突变单株的群体连锁分析发现,其中7个距离目标基因最近的标记分别是InDel7、InDel3、InDel4,InDel10、InDel9、InDel8和InDel5,其交换单株数分别是23、1、0、0、1、3和13(图3)。最终确定oy2基因位于标记InDel3和InDel9之间的约117Kb的区间内,共包含4个候选基因(图3)。
实施例2玉米叶片黄化调控基因oy2的分子标记引物设计
对定位区间内4个候选基因在野生型和突变体植株oy2之间的测序比较分析,发现区间内OY2基因(SEQ ID NO.6)的第十三个外显子上发生一个单碱基突变(由G突变成A)(图3,图4,图4中方框标记的即为点突变碱基,正常为G,黄化为A),即oy2突变基因为OY2基因的等位突变基因。在此突变位点两端设计引物,获得野生型与突变体oy2之间的多态InDel标记。引物序列如下:
上游(F):GAATTGGTTTGCGCCACAGT(SEQ ID No.3);
下游(R):TGTTTATTCTTCAAGGCTGTCAGA(SEQ ID No.4);
用设计的引物进行PCR扩增,其中,PCR反应体系为:Taq Mix 7.5μL,上下游引物分别为0.75μL,DNA模板1μL,用双蒸水定容至15μL。
PCR扩增程序为:94℃预变性2min;98℃变性10s;60℃退火30s;68℃延伸5s;共35个循环;68℃再延伸10min。。
实施例3玉米叶片黄化相关色素变化及叶绿体超微结构观察
1、玉米苗期叶片相关色素变化
取玉米苗期三叶一心时期的突变体oy2和野生型RP125的叶片,结果显示突变体中叶绿素a、b(Chl a、Chl b)及总叶绿素(Total Chl)含量均比野生型显著降低,而类胡萝卜素(Car)变化不明显(表1)。叶绿素含量的大幅降低造成植物的光合作用产生严重影响。通过测定过氧化氢(H2O2)、过氧化物酶(POD)、超氧化物酶歧化酶(SOD)、过氧化氢酶(CAT)、丙二醛(MDA)等指标,发现突变体与野生型中活性氧相关酶出现了显著性变化(表1)。本研究中色素含量测定均采用常规分光光度计法,测定过氧化氢(H2O2)、过氧化物酶(POD)、超氧化物酶歧化酶(SOD)、过氧化氢酶(CAT)、丙二醛(MDA)试剂盒购置于南京建成生物工程研究所有限公司。
表1突变体oy2和野生型RP125中色素及相关活性酶测定
注:NS表示无差异,**(P<0.01)表示极显著性差异
2、玉米苗期叶绿体超微结构观察
为了研究oy2基因突变对玉米叶绿体发育的影响,我们通过透射电镜观察分析了玉米苗期三叶一心时期的突变体oy2和野生型(RP125)叶绿体超微结构。通过观察发现突变体中叶绿体发育存在明显的障碍,主要表现为叶绿体在叶肉细胞中排列散乱,且叶绿体间缺乏明显的叶绿体边界,缺乏类囊体且类囊体膜不清晰,叶绿体由于缺乏边界而杂乱无章地融合在一起,且没有明显的淀粉粒(图2)。
如图2所示,与野生型相比,突变体叶绿体结构受到损伤,叶绿体和淀粉粒明显减少,类囊体膜结构不完整,嗜锇颗粒增加。图2中(a)(c)为不同放大倍数下,突变体oy2叶绿体及细胞器构造;(b)(d)为不同放大倍数下,野生型叶绿体及细胞器构造。标尺上边为20μm,下边为2μm。CP:叶绿体;SG:淀粉粒;T:类囊体;OG:嗜锇颗粒。
实施例4玉米叶片黄化隐性核基因oy2的分子标记验证试验
取实例1中F2分离群体中叶片正常和黄化单株,提取叶片DNA,利用以上引物进行PCR扩增。PCR反应体系及扩增程序见实例2。PCR产物进行测序(图3,图4),结果叶片正常且无分离的植株中在Chr5:3207013位点碱基为G,即为具有OY2/OY2基因型的材料;叶片正常且表现出分离的植株中,在Chr5:3207013位点碱基为G和A,即为具有OY2/oy2杂合基因型的材料;黄化且无分离的植株中,在Chr5:3207013位点碱基仅为A,即为具有oy2/oy2基因型的材料。
上述结果说明,该InDel分子标记与玉米苗期叶片黄化性状紧密连锁,用本发明分子标记鉴定玉米苗期黄化叶片材料的oy2基因型准确、可靠,可以作为玉米苗期黄化叶片基因oy2的分子标记,提高玉米高光效分子育种水平。
序列表
<110> 四川农业大学
<120> 一种玉米叶黄化基因oy2以及与其连锁的InDel分子标记和应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2268
<212> DNA
<213> oy2(Zea mays oy2)
<400> 1
taccgctgcg ggtggcgcga gaggtggagt gaggggatgg agggcggggc ggcgcagtag 60
agtaagggta ggcggcggcg gcagagggag gggcagtgga gggcggggcg ggccgacgcc 120
ctaagggcgg agcgccggcg ccgttggagc cggaggctcc aggagctcag gtggttgccg 180
cggcaggggt gacgccggtt cctgccgcgc accgcgccca taccctccct catgaagggg 240
gaccgacggc agcaacccgt cctacgatag ttttgacgag acgacgaacc ccgctaacta 300
gcactctagc ctccgtaacg gtagagtccc ttcgcaccct gccgtttctg tcactaccga 360
gcaccaaacg tacgatacga aggtgggtaa cttcaccacc aaccaaggta acgtttacga 420
ctgggattga ggggactgct taccctccta ccaaatcgac tagtttatgt catactgaga 480
ctaccattac agtttaggct ctaacagttt tgtggaaaac acgtctaagg tgaaccacac 540
tgcctcctat ccgagtaacc tagtcaacta caacttcgta gacactctag tccctgatga 600
cataaagttg gaccagaaga acgacttcgt gtatctccac aagaaataca actactttat 660
ttagataacc tactaccgta ttcgttagat gaagacttac agaactgcct ccctcaattg 720
taacaccttt ctctcccgta atcgaaagcg gtagggacgt ttggtgaaga ttaacgatga 780
atgttaggtc tccttcccag acatgcactt gtgaacgaac tagcataacg ttaattaaat 840
tcacgactag aaggttactc aaaactactg gcgcaacttc gtcacctata acgttgtgcc 900
aaagtcctca gatcgtttct tcaaaagttt taccaccttc ttttttgact ttgacgtttt 960
tgagtctatt aaaaacgttc tctcatagac ttcctacaat gataatcgtg tctcgtcgag 1020
tttatagaac agtaccttcg atatgctcca ccgacagtcc ccgtagcacg actcaacata 1080
cgacgagctc aacgttttac agatcgacga taccttcctg cactttttca taaacaccta 1140
ctggagttct ttcgacatct cgatcagtaa gatggagcga ggtaggatag actattaggt 1200
gtcctagtcg tcgttctcgt tggtgggggt gggggcggtg gtggtggagg tcttttagtt 1260
ctaagaagtc ttctggttct actccttctg cttctggttc tcctactact tcttctttta 1320
cttgttgttc tggttgtcta tggactcctc aagtataaac tacgacttcc accaaatcat 1380
ctactgtttg aagaaaagaa acgggtcgtt cgtgtctctg ctgcaccttt tcgacccgct 1440
cgtttcttac agtagaagag tcttctatcc ccggcaatgt atttcggata cgaaggattc 1500
ccaggtcatt cctccaatcg acaactacgg tgcgaatctc gtcgacgtgg tatggttttt 1560
gacgcgtctc tctttcttgc actgttttgt tctttccaaa aacaactttt ctgactgtac 1620
tctcggtttt cttaccgagc ttttcgtcca cgagatcagt ataaacaaca cctgcgatca 1680
ccatcgtacc gagacttagc atacgtctta cgatttccac gccgcaactt caacgaacgt 1740
ctttcgatgt ggtcgtctct agtccaaagt taataaggaa aagcacctct aatacgactc 1800
caaaacgaag gtggtagttc tagatatcgt taccgggcct ttgcagaact cttcgatggt 1860
acaccaccac caagaggaaa tcgagtaccg gattcatgtc gacagtctca cccagactta 1920
cgacttttct caccgctaca acccgcatag tactaacaac gttagtagct accttctcga 1980
ttacatagtg acttctctag gtgactgggt cttcgacgac gacgaagtct acgtggttct 2040
ggaagaagag ttcttgactt cctgctctat gaactccacc gaccgtttta tatgttccgt 2100
ccttacaggg aagaacagta gctgtgactc ttgttcaaac ataggtgccc taaacggttc 2160
ctttaacgtt cccaacgggt cccctttata ataatggagg gattacgaag cctacgacat 2220
taaagacgac ggtggttctg gcgggactgt ctgaacttct cgagtact 2268
<210> 2
<211> 729
<212> PRT
<213> oy2(Zea mays oy2)
<400> 2
Met Ala Thr Pro Thr Ala Leu Ser Thr Ser Leu Pro Tyr Leu Pro Pro
1 5 10 15
Arg Arg Val Ile Ser Phe Pro Ser Ala Ala Ala Val Ser Leu Pro Val
20 25 30
Thr Ser Arg Pro Ala Arg Leu Arg Asp Ser Arg Leu Ala Ala Ala Ala
35 40 45
Thr Ser Ala Ser Glu Val Leu Glu Ser Thr Asn Gly Ala Val Pro Thr
50 55 60
Ala Ala Lys Asp Gly Ala Trp Arg Gly Tyr Gly Arg Glu Tyr Phe Pro
65 70 75 80
Leu Ala Ala Val Val Gly Gln Asp Ala Ile Lys Thr Ala Leu Leu Leu
85 90 95
Gly Ala Ile Asp Arg Glu Ile Gly Gly Ile Ala Ile Ser Gly Lys Arg
100 105 110
Gly Thr Ala Lys Thr Val Met Ala Arg Gly Leu His Ala Met Leu Pro
115 120 125
Pro Ile Glu Val Val Val Gly Ser Ile Ala Asn Ala Asp Pro Asn Ser
130 135 140
Pro Asp Glu Trp Glu Asp Gly Leu Ala Asp Gln Ile Gln Tyr Asp Ser
145 150 155 160
Asp Gly Asn Val Lys Ser Glu Ile Val Lys Thr Pro Phe Val Gln Ile
165 170 175
Pro Leu Gly Val Thr Glu Asp Arg Leu Ile Gly Ser Val Asp Val Glu
180 185 190
Ala Ser Val Arg Ser Gly Thr Thr Val Phe Gln Pro Gly Leu Leu Ala
195 200 205
Glu Ala His Arg Gly Val Leu Tyr Val Asp Glu Ile Asn Leu Leu Asp
210 215 220
Asp Gly Ile Ser Asn Leu Leu Leu Asn Val Leu Thr Glu Gly Val Asn
225 230 235 240
Ile Val Glu Arg Glu Gly Ile Ser Phe Arg His Pro Cys Lys Pro Leu
245 250 255
Leu Ile Ala Thr Tyr Asn Pro Glu Glu Gly Ser Val Arg Glu His Leu
260 265 270
Leu Asp Arg Ile Ala Ile Asn Leu Ser Ala Asp Leu Pro Met Ser Phe
275 280 285
Asp Asp Arg Val Glu Ala Val Asp Ile Ala Thr Arg Phe Gln Glu Ser
290 295 300
Ser Lys Glu Val Phe Lys Met Val Glu Glu Lys Thr Glu Thr Ala Lys
305 310 315 320
Thr Gln Ile Ile Phe Ala Arg Glu Tyr Leu Lys Asp Val Thr Ile Ser
325 330 335
Thr Glu Gln Leu Lys Tyr Leu Val Met Glu Ala Ile Arg Gly Gly Cys
340 345 350
Gln Gly His Arg Ala Glu Leu Tyr Ala Ala Arg Val Ala Lys Cys Leu
355 360 365
Ala Ala Met Glu Gly Arg Glu Lys Val Glu Leu Val Ile Leu Pro Arg
370 375 380
Ser Ile Leu Ser Asp Asn Pro Gln Asp Gln Gln Gln Glu Gln Pro Pro
385 390 395 400
Pro Pro Pro Pro Pro Pro Pro Pro Glu Asn Gln Asp Ser Ser Glu Asp
405 410 415
Gln Asp Glu Glu Asp Glu Asp Gln Glu Asp Asp Glu Glu Glu Asn Glu
420 425 430
Gln Gln Asp Gln Gln Ile Pro Glu Glu Phe Ile Phe Asp Ala Glu Gly
435 440 445
Gly Leu Val Asp Asp Lys Leu Leu Phe Phe Ala Gln Gln Ala Gln Arg
450 455 460
Arg Arg Gly Lys Ala Gly Arg Ala Lys Asn Gly Pro Val Arg Arg Leu
465 470 475 480
Ala Val Asp Ala Thr Leu Arg Ala Ala Ala Pro Tyr Gln Lys Leu Arg
485 490 495
Arg Glu Lys Glu Arg Asp Lys Thr Arg Lys Val Phe Val Glu Lys Thr
500 505 510
Asp Met Arg Ala Lys Arg Met Ala Arg Lys Ala Gly Ala Leu Val Ile
515 520 525
Phe Val Val Asp Ala Ser Gly Ser Met Ala Leu Asn Arg Met Gln Asn
530 535 540
Ala Lys Gly Ala Ala Leu Lys Leu Leu Ala Glu Ser Tyr Thr Ser Arg
545 550 555 560
Asp Gln Val Ser Ile Ile Pro Phe Arg Gly Asp Tyr Ala Glu Val Leu
565 570 575
Leu Pro Pro Ser Arg Ser Ile Ala Met Ala Arg Lys Arg Leu Glu Lys
580 585 590
Leu Pro Cys Gly Gly Gly Ser Pro Leu Ala His Gly Leu Ser Thr Ala
595 600 605
Val Arg Val Gly Leu Asn Ala Glu Lys Ser Gly Asp Val Gly Arg Ile
610 615 620
Met Ile Val Ala Ile Thr Asp Gly Arg Ala Asn Val Ser Leu Lys Arg
625 630 635 640
Ser Thr Asp Pro Glu Ala Ala Ala Ala Ser Asp Ala Pro Arg Pro Ser
645 650 655
Ser Gln Glu Leu Lys Asp Glu Ile Leu Glu Val Ala Gly Lys Ile Tyr
660 665 670
Lys Ala Gly Met Ser Leu Leu Val Ile Asp Thr Glu Asn Lys Phe Val
675 680 685
Ser Thr Gly Phe Ala Lys Glu Ile Ala Arg Val Ala Gln Gly Lys Tyr
690 695 700
Tyr Tyr Leu Pro Asn Ala Ser Asp Ala Val Ile Ser Ala Ala Thr Lys
705 710 715 720
Thr Ala Leu Thr Asp Leu Lys Ser Ser
725
<210> 3
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
gaattggttt gcgccacagt 20
<210> 4
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
tgtttattct tcaaggctgt caga 24
<210> 5
<211> 2190
<212> DNA
<213> 玉米(Zea mays oy2)
<400> 5
taccgctgcg ggtggcgcga gaggtggagt gaggggatgg agggcggggc ggcgcagtag 60
agtaagggta ggcggcggcg gcagagggag gggcagtgga gggcggggcg ggccgacgcc 120
ctaagggcgg agcgccggcg ccgttggagc cggaggctcc aggagctcag gtggttgccg 180
cggcaggggt gacgccggtt cctgccgcgc accgcgccca taccctccct catgaagggg 240
gaccgacggc agcaacccgt cctacgatag ttttgacgag acgacgaacc ccgctaacta 300
gcactctagc ctccgtaacg gtagagtccc ttcgcaccct gccgtttctg tcactaccga 360
gcaccaaacg tacgatacga aggtgggtaa cttcaccacc aaccaaggta acgtttacga 420
ctgggattga ggggactgct taccctccta ccaaatcgac tagtttatgt catactgaga 480
ctaccattac agtttaggct ctaacagttt tgtggaaaac acgtctaagg tgaaccacac 540
tgcctcctat ccgagtaacc tagtcaacta caacttcgta gacactctag tccctgatga 600
cataaagttg gaccagaaga acgacttcgt gtatctccac aagaaataca actactttat 660
ttagataacc tactaccgta ttcgttagat gaagacttac agaactgcct ccctcaattg 720
taacaccttt ctctcccgta atcgaaagcg gtagggacgt ttggtgaaga ttaacgatga 780
atgttaggtc tccttcccag acatgcactt gtgaacgaac tagcataacg ttaattaaat 840
tcacgactag aaggttactc aaaactactg gcgcaacttc gtcacctata acgttgtgcc 900
aaagtcctca gatcgtttct tcaaaagttt taccaccttc ttttttgact ttgacgtttt 960
tgagtctatt aaaaacgttc tctcatagac ttcctacaat gataatcgtg tctcgtcgag 1020
tttatagaac agtaccttcg atatgctcca ccgacagtcc ccgtagcacg actcaacata 1080
cgacgagctc aacgttttac agatcgacga taccttcctg cactttttca tctcgatcag 1140
taagatggag cgaggtagga tagactatta ggtgtcctag tcgtcgttct cgttggtggg 1200
ggtgggggcg gtggtggtgg aggtctttta gttctaagaa gtcttctggt tctactcctt 1260
ctgcttctgg ttctcctact acttcttctt ttacttgttg ttctggttgt ctatggactc 1320
ctcaagtata aactacgact tccaccaaat catctactgt ttgaagaaaa gaaacgggtc 1380
gttcgtgtct ctgctgcacc ttttcgaccc gctcgtttct taccaggtca ttcctccaat 1440
cgacaactac ggtgcgaatc tcgtcgacgt ggtatggttt ttgacgcgtc tctctttctt 1500
gcactgtttt gttctttcca aaaacaactt ttctgactgt actctcggtt ttcttaccga 1560
gcttttcgtc cacgagatca gtataaacaa cacctgcgat caccatcgta ccgagactta 1620
gcatacgtct tacgatttcc acgccgcaac ttcaacgaac gtctttcgat gtggtcgtct 1680
ctagtccaaa gttaataagg aaaagcacct ctaatacgac tccaaaacga aggtggtagt 1740
tctagatatc gttaccgggc ctttgcagaa ctcttcgatg gtacaccacc accaagagga 1800
aatcgagtac cggattcatg tcgacagtct cacccagact tacgactttt ctcaccgcta 1860
caacccgcat agtactaaca acgttagtgg ctaccttctc gattacatag tgacttctct 1920
aggtgactgg gtcttcgacg acgacgaagt ctacgtggtt ctggaagaag agttcttgac 1980
ttcctgctct atgaactcca ccgaccgttt tatatgttcc gtccttacag ggaagaacag 2040
tagctgtgac tcttgttcaa acataggtgc cctaaacggt tcctttaacg ttcccaacgg 2100
gtccccttta taataatgga gggattacga agcctacgac attaaagacg acggtggttc 2160
tggcgggact gtctgaactt ctcgagtact 2190
<210> 6
<211> 2268
<212> DNA
<213> 玉米(Zea mays oy2)
<400> 6
taccgctgcg ggtggcgcga gaggtggagt gaggggatgg agggcggggc ggcgcagtag 60
agtaagggta ggcggcggcg gcagagggag gggcagtgga gggcggggcg ggccgacgcc 120
ctaagggcgg agcgccggcg ccgttggagc cggaggctcc aggagctcag gtggttgccg 180
cggcaggggt gacgccggtt cctgccgcgc accgcgccca taccctccct catgaagggg 240
gaccgacggc agcaacccgt cctacgatag ttttgacgag acgacgaacc ccgctaacta 300
gcactctagc ctccgtaacg gtagagtccc ttcgcaccct gccgtttctg tcactaccga 360
gcaccaaacg tacgatacga aggtgggtaa cttcaccacc aaccaaggta acgtttacga 420
ctgggattga ggggactgct taccctccta ccaaatcgac tagtttatgt catactgaga 480
ctaccattac agtttaggct ctaacagttt tgtggaaaac acgtctaagg tgaaccacac 540
tgcctcctat ccgagtaacc tagtcaacta caacttcgta gacactctag tccctgatga 600
cataaagttg gaccagaaga acgacttcgt gtatctccac aagaaataca actactttat 660
ttagataacc tactaccgta ttcgttagat gaagacttac agaactgcct ccctcaattg 720
taacaccttt ctctcccgta atcgaaagcg gtagggacgt ttggtgaaga ttaacgatga 780
atgttaggtc tccttcccag acatgcactt gtgaacgaac tagcataacg ttaattaaat 840
tcacgactag aaggttactc aaaactactg gcgcaacttc gtcacctata acgttgtgcc 900
aaagtcctca gatcgtttct tcaaaagttt taccaccttc ttttttgact ttgacgtttt 960
tgagtctatt aaaaacgttc tctcatagac ttcctacaat gataatcgtg tctcgtcgag 1020
tttatagaac agtaccttcg atatgctcca ccgacagtcc ccgtagcacg actcaacata 1080
cgacgagctc aacgttttac agatcgacga taccttcctg cactttttca taaacaccta 1140
ctggagttct ttcgacatct cgatcagtaa gatggagcga ggtaggatag actattaggt 1200
gtcctagtcg tcgttctcgt tggtgggggt gggggcggtg gtggtggagg tcttttagtt 1260
ctaagaagtc ttctggttct actccttctg cttctggttc tcctactact tcttctttta 1320
cttgttgttc tggttgtcta tggactcctc aagtataaac tacgacttcc accaaatcat 1380
ctactgtttg aagaaaagaa acgggtcgtt cgtgtctctg ctgcaccttt tcgacccgct 1440
cgtttcttac agtagaagag tcttctatcc ccggcaatgt atttcggata cgaaggattc 1500
ccaggtcatt cctccaatcg acaactacgg tgcgaatctc gtcgacgtgg tatggttttt 1560
gacgcgtctc tctttcttgc actgttttgt tctttccaaa aacaactttt ctgactgtac 1620
tctcggtttt cttaccgagc ttttcgtcca cgagatcagt ataaacaaca cctgcgatca 1680
ccatcgtacc gagacttagc atacgtctta cgatttccac gccgcaactt caacgaacgt 1740
ctttcgatgt ggtcgtctct agtccaaagt taataaggaa aagcacctct aatacgactc 1800
caaaacgaag gtggtagttc tagatatcgt taccgggcct ttgcagaact cttcgatggt 1860
acaccaccac caagaggaaa tcgagtaccg gattcatgtc gacagtctca cccagactta 1920
cgacttttct caccgctaca acccgcatag tactaacaac gttagtggct accttctcga 1980
ttacatagtg acttctctag gtgactgggt cttcgacgac gacgaagtct acgtggttct 2040
ggaagaagag ttcttgactt cctgctctat gaactccacc gaccgtttta tatgttccgt 2100
ccttacaggg aagaacagta gctgtgactc ttgttcaaac ataggtgccc taaacggttc 2160
ctttaacgtt cccaacgggt cccctttata ataatggagg gattacgaag cctacgacat 2220
taaagacgac ggtggttctg gcgggactgt ctgaacttct cgagtact 2268
Claims (10)
1.一种玉米叶黄化基因oy2,其特征在于,所述基因的核苷酸序列如SEQ ID NO.1所示。
2.根据权利要求1所述的玉米叶黄化基因oy2,其特征在于,所述基因在编码区第十三个外显子上发生一个单碱基突变,突变位点为G/A。
3.根据权利要求1所述的玉米叶黄化基因oy2,其特征在于,所述基因编码的蛋白质的氨基酸序列如SEQ ID NO.2所示。
4.一种与基因oy2连锁的InDel分子标记,其特征在于,所述InDel分子标记定位于玉米5号染色体上,其核苷酸序列如SEQ ID NO.5所示。
5.一种用于扩增权利要求3所述的InDel分子标记的引物对,其特征在于,所述引物对的具体序列如下:
InDel-F:GAATTGGTTTGCGCCACAGT;
InDel-R:TGTTTATTCTTCAAGGCTGTCAGA。
6.一种筛选黄化叶玉米的方法,其特征在于,包括以下步骤:
(1)提取待测玉米DNA;
(2)采用权利要求4所述的引物对待测基因进行PCR扩增;
(3)对扩增产物进行测序,如若扩增产物在突变位点的核苷酸碱基为A/A,则其为黄化叶玉米。
7.根据权利要求6所述的方法,其特征在于,所述PCR程序为:94℃预变性2min;98℃变性10s;60℃退火30s;68℃延伸5s;共35个循环;68℃再延伸10min。
8.根据权利要求6所述的方法,其特征在于,所述PCR反应体系为:Taq Mix7.5μL,上下游引物分别为0.75μL,DNA模板1μL,用双蒸水定容至15μL。
9.一种筛选黄化叶玉米的试剂盒,其特征在于,包括权利要求4所述分子标记,和/或权利要求5所述的引物对。
10.一种玉米全基因组芯片,其特征在于,包括权利要求4所述的分子标记。
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CN114480717A (zh) * | 2022-03-04 | 2022-05-13 | 广东省科学院南繁种业研究所 | 玉米籽粒颜色相关的InDel标记及其应用 |
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CN113528539B (zh) * | 2021-08-18 | 2022-06-07 | 四川农业大学 | 一种玉米苗期斑马叶及白粒基因zb10及其连锁的分子标记和应用 |
CN114480717A (zh) * | 2022-03-04 | 2022-05-13 | 广东省科学院南繁种业研究所 | 玉米籽粒颜色相关的InDel标记及其应用 |
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