CN114921479A - 一种调控抗性淀粉含量的水稻SBEIIb等位基因及其在育种中的应用 - Google Patents
一种调控抗性淀粉含量的水稻SBEIIb等位基因及其在育种中的应用 Download PDFInfo
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- CN114921479A CN114921479A CN202210622297.XA CN202210622297A CN114921479A CN 114921479 A CN114921479 A CN 114921479A CN 202210622297 A CN202210622297 A CN 202210622297A CN 114921479 A CN114921479 A CN 114921479A
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Abstract
本发明属于分子遗传学领域,公开了一种调控抗性淀粉含量的水稻SBEIIb等位基因。该基因在野生型水稻品种日本晴的SBEIIb基因第16外显子上的第56位腺嘌呤(A)被鸟嘌呤(G)替换,其核苷酸序列如SEQ ID No.1所示,编码区序列如SEQ ID No.2所示,所述水稻SBEIIb等位基因编码的蛋白氨基酸序列如SEQ ID No.3所示。上述SBEIIb等位基因对直链淀粉含量无显著影响,但改变水稻支链淀粉的链长;可显著提高米饭的抗性淀粉含量和降低米饭葡萄糖生成速率,且对稻米品质影响较小,在高抗性淀粉水稻育种中具有很强的应用。
Description
技术领域
本发明涉及一种调控抗性淀粉含量的水稻SBEIIb等位基因及其在育种中的应用,属于分子遗传学领域。
背景技术
抗性淀粉(Resistant starch,RS)是指不能被健康人体小肠酶解,但可以在人的结肠中被发酵分解的淀粉及其降解物。摄食富含RS的食物可以维持餐后血糖水平和改善肠道健康。随着人们生活水平提高和生产方式的改变,患超重、肥胖和糖尿病等的人群在逐年增加,而摄食富含RS的食物可以有效预防和延缓上述疾病的发生。淀粉分支酶IIb(Starchbranching enzyme IIb,SBEIIb)负责支链淀粉侧链的形成,调控稻米抗性淀粉含量,因此挖掘水稻SBEIIb等位基因非常重要。目前,国内外育种学家已经挖掘了很多SBEIIb等位基因,它们大多导致SBEIIb缺失或活性显著降低,最终形成粉质籽粒,而粉质籽粒的外观、产量和蒸煮食味品质等较差,限制了其在农业生产和人们生活中的直接应用。
发明内容
本发明旨在提供一种调控抗性淀粉含量的水稻SBEIIb等位基因及其在育种中的应用,尤其是在高抗性淀粉水稻育种中的应用,它显著改变支链淀粉链长分布,提高稻米抗性淀粉含量,但不影响直链淀粉含量和稻米外观品质,显示在高抗性淀粉稻米育种中有很强的应用前景。
本发明提供的技术方案如下:
一种调控抗性淀粉含量的水稻SBEIIb等位基因,在野生型水稻品种日本晴SBEIIb基因基础上,第16外显子上的第56位A被G替换;所述的水稻SBEIIb等位基因其核苷酸序列如SEQ ID No.1所示。
进一步地,其编码区序列如SEQ ID No.2所示。
本发明还提供一种用于检测调控抗性淀粉含量的水稻SBEIIb等位基因的引物,所述引物的上游序列如SEQ ID NO:6所示,所述引物的下游序列如SEQ ID NO:7所示。
本发明还提供上述水稻SBEIIb等位基因所编码的蛋白,在野生型日本晴SBEIIb蛋白基础上,第583位氨基酸精氨酸(Arg)被甘氨酸(Gly)替换;所述的水稻SBEIIb等位基因所编码蛋白的氨基酸序列如SEQ ID No.3所示。
本发明还提供上述水稻SBEIIb等位基因在高抗性淀粉水稻育种中的应用。
进一步地,所述水稻SBEIIb等位基因提高水稻支链淀粉长侧链的比例。
进一步地,水稻SBEIIb等位基因提高米饭的抗性淀粉含量并降低葡萄糖生成速率。
进一步地,水稻SBEIIb等位基因提高稻米的糊化温度。
进一步地,将权利要求1或2所述的基因目标碱基通过基因编辑技术改变目的水稻的目标碱基,得到目标碱基替换的水稻材料;所述目标碱基替换的水稻的稻米尿素糊化抗性高于所述目的水稻。
进一步地,以序列为SEQ ID NO:4和SEQ ID NO:5的引物对T0代植株进行扩增。
本发明提供了一个调控抗性淀粉含量相关的等位基因,其在粳稻日本晴背景的SBEIIb基因(Genebank:AP014958.1,https://www.ncbi.nlm.nih.gov/nucleotide/ AP014958.1)第16外显子上的第56位碱基A被G替换,其基因核苷酸序列如SEQ ID No.1所示,编码区序列如SEQ ID No.2所示。本发明所述的SBEIIb等位基因可以为高抗性淀粉水稻育种提供优异的种质资源。
有益效果
目前尚未有SBEIIb基因该等位变异的报道,是一个新的等位基因突变。本发明从EMS诱变的粳稻日本晴非粉质胚乳突变体库中筛选一个SBEIIb新等位基因,本发明提供的新等位基因编码淀粉分支酶SBEIIb,主要负责支链淀粉短侧链的形成,对淀粉消化特性的调控起关键作用。该SBEIIb新等位基因可以显著改变支链淀粉链长分布但不影响直链淀粉含量,能显著降低米饭的生糖速率和提高米饭的抗性淀粉含量,但对稻米品质影响较小,是高抗性淀粉稻米品质改良育种的优异种质资源,在高抗性淀粉稻米育种中有很强的应用前景。
附图说明
图1为本发明中野生型水稻品种日本晴和携带SBEIIb新等位基因的纯合突变体MY03的糊化特性和籽粒表型。A为籽粒的尿素糊化特性,B为淀粉的热力学特性,C为糙米和精米的表型。
图2为本发明中野生型水稻品种日本晴和携带SBEIIb新等位基因的纯合突变体MY03的淀粉组分。A为淀粉的碘吸收光谱,B为直链淀粉含量,C为支链淀粉链长分布,D为突变体与野生型水稻的支链淀粉链长分布的差异。
图3为本发明中野生型水稻品种日本晴和携带SBEIIb新等位基因的纯合突变体MY03的SBEIIb突变分析。A为SBEIIb基因序列分析,B为SBEIIb氨基酸序列分析,C为突变的目标氨基酸在不同物种淀粉分支酶中的保守性分析。
图4为本发明中SBEIIb新等位基因功能验证。A为T0代转基因苗验证,B为T0代突变株靶位点序列分析,C为T1代纯合突变株的籽粒抗尿素糊化检测。
图5为本发明中野生型水稻品种日本晴和携带SBEIIb新等位基因的纯合突变体MY03的稻米外观品质分析。A为糙米和精米的粒长,B为糙米和精米的粒宽,C为糙米和精米的粒厚,D为糙米和精米的千粒重,E为精米的垩白性状,F为精米的透明度。
具体实施方式
为使本发明的上述目的、特征和优点能够更加明显易懂,下面结合具体实施例对本发明的具体实施方式做详细的说明。
在下面的描述中阐述了很多具体细节以便于充分理解本发明,但是本发明还可以采用其他不同于在此描述的其它方式来实施,本领域技术人员可以在不违背本发明内涵的情况下做类似推广,因此本发明不受下面公开的具体实施例的限制。
1、实验材料
以野生型水稻品种日本晴及其来源的突变体MY03和野生型水稻品种中花11及其来源的突变体ZH11-MY03为实验材料。
MY03是从EMS诱变的日本晴非粉质胚乳突变体库中筛选出来的,筛选方法是用锋利的刀片将成熟糙米从中部横切,取不带胚的半籽粒作为糊化样本,水平放入平底96孔板中,加入200μL 3M的尿素溶液(此浓度能使野生型籽粒糊化导致碘染后染液呈蓝色),置于24℃恒温箱中糊化16h。糊化后在96孔板中加入20μL碘液(0.2%碘,1%碘化钾,0.5%醋酸)染色,并吹打混匀。显色后观察染液颜色,筛选染液颜色呈黄色的材料,将其对应的带胚半粒发苗繁种,单株收种。
ZH11-MY03是利用CRISPR/Cas9单碱基编辑系统(ABE)在粳稻品种中花11(ZH11)背景下创建的与MY03所携带突变基因一致的突变体。具体的转化过程为:挑选饱满透亮无霉点的ZH11成熟糙米,经75%酒精和次氯酸钠溶液清洗消毒,滤纸吸干多余水分后接种于诱导培养基上,光照培养箱中28℃光照培养约28天(每天16h光照/8h黑暗)。将嫩黄愈伤粒转接到继代培养基中相同条件下培养3-7天。将单碱基替换编辑载体转化到EHA105菌株中,挑选阳性菌落摇菌并测定OD600,使其在0.6~0.8。吸取少量菌液后离心,悬浮并转移到15mL侵染液体培养基中。加入继代的愈伤后于22℃暗培养30min。滤纸上吸干愈伤上多余菌液后转移至共培养培养基,22℃暗培养2-3天。用加入羧苄的无菌水充分清洗共培养后的愈伤,滤纸吸干水分后,于恢复培养基中培养3-4天。随后将愈伤依次转移到含有潮霉素的一筛培养基和二筛培养基中筛选15天。将经过筛选培养后新长出的嫩黄小粒愈伤转移到分化培养基中进行分化再生1~2个月。长出的幼苗转移到生根培养基中生长壮大后转移至人工气候箱中,进行炼苗和突变鉴定,将纯合突变植株培育至开花,单株收种。
2、籽粒的糊化特性和表型分析
利用尿素糊化半籽粒-碘液染色法,从EMS诱变的日本晴非粉质胚乳突变体库中筛选到一个抗尿素糊化突变体MY03。日本晴籽粒在3M尿素中已糊化,而MY03籽粒在6M尿素中才开始糊化(图1A)。进一步利用差示扫描量热仪(DSC)比较日本晴和MY03淀粉的热力学特性发现,MY03(80.8℃)糊化温度显著高于日本晴(70.5℃)(图1B)。分别在反射光和透射光下观察糙米和精米的表型发现,MY03的籽粒表型与日本晴无明显差异,均为饱满的透明籽粒(图1C)。以上结果表明,MY03是一个抗糊化的透明籽粒突变体。
3、淀粉组分分析
利用碘比色法和伴刀豆球蛋白A(Con A)沉淀法(采用直链/支链淀粉检测试剂盒测定)对日本晴和MY03的淀粉组分进行分析。淀粉碘比色的OD620反映与碘分子结合的直链淀粉和支链淀粉长侧链的吸光度,通常用于评估表观直链淀粉含量(AAC)。碘比色测定结果显示MY03的AAC显著比日本晴高(图2A)。Con A可以与支链淀粉特异性结合并形成沉淀,测量的直链淀粉含量(AC)是直链淀粉与直链淀粉和支链淀粉总量的百分比,该方法不受淀粉纯度和支链淀粉长侧链的影响,因此通常将测得的值称为真实直链淀粉含量。本发明使用爱尔兰Megazyme公司的直链/支链淀粉检测试剂盒测得的直链淀粉含量显示,MY03(14.3%)的直链淀粉含量与日本晴(13.5%)无明显差异(图2B)。MY03的AAC显著比日本晴高,而AC与日本晴无明显差异,表明MY03的支链淀粉长侧链的比例升高。进一步,利用荧光辅助毛细管电泳(FACE)比较日本晴和MY03的支链淀粉链长分布发现,MY03的短侧链比例显著降低,而中长侧链比例显著升高(图2C,D)。以上结果表明,MY03与野生型相比,直链淀粉含量未发生明显变化,而支链淀粉链长分布发生显著改变。
4、SBEIIb新等位基因鉴定
SBEIIb被认为主要负责支链淀粉短侧链的形成,通常SBEIIb活性受抑制会导致胚乳淀粉中支链淀粉长侧链比例显著升高,同时短侧链含量显著降低。且水稻SBEIIb基因突变一般会导致淀粉抗糊化,因此本发明首先对MY03的SBEIIb基因进行测序。用CTAB法提取日本晴和MY03的叶片基因组,以叶片DNA为模板进行PCR扩增,扩增产物经琼脂糖凝胶电泳鉴定后委托南京擎科生物公司进行测序。测序结果显示,MY03的SBEIIb基因第16外显子上的第56位碱基A被G替换(图3A),基因序列如SEQ ID No.1所示。该突变导致SBEIIb蛋白第583位氨基酸由带正电荷的碱性Arg替换为不带电荷的极性甘氨酸(图3B),氨基酸序列如SEQ ID No.3所示。本发明进一步将野生型的氨基酸序列粘贴到NCBI(http:// www.ncbi.nlm.nih.gov/)数据库中,进行BlastP对比,下载SBEIIb在常见物种中的同源蛋白,利用DNAMAN软件对突变氨基酸在不同物种中的保守性进行分析。结果显示水稻的SBEIIb第583位氨基酸在很多物种的SBE蛋白家族中高度保守,包括玉米、小麦、大麦和拟南芥的SBEIIb、SBEIIa和SBEI(图3C)。因此,该位点的变异对SBEIIb蛋白的功能具有一定的影响。经查阅文献资料和相关数据库,目前MY03的SBEIIb突变位点未见报道。因此,突变体MY03是一种SBEIIb新等位变异,并且该位点高度保守。
5、SBEIIb新等位基因功能验证
为验证MY03籽粒抗糊化的表型是否由突变的SBEIIb产生,本发明在粳稻ZH11背景下,利用CRISPR/Cas9单碱基替换编辑器将ZH11的SBEIIb目标碱基进行了替换。将构建好的单碱基替换编辑载体通过农杆菌侵染法转化ZH11的愈伤组织,获得10株T0代转基因苗。利用潮霉素(HYG)引物(上游引物:5’-CGAGAGCCTGACCTATTGCAT-3’(SEQ ID NO:4),下游引物:5’-CTGCTCCATACAAGCCAACCAC-3’(SEQ ID NO:5))对T0代植株的潮霉素抗性基因进行扩增,以未进行转化的野生型ZH11的叶片DNA为阴性对照,并用1%琼脂糖凝胶电泳筛选含有目标条带的阳性植株。结果显示,10株苗均为转化成功的植株(图4A)。利用SBEIIb测序引物(上游引物:5’-AGCCTGTCGCCTGTATGGT-3’(SEQ ID NO:6),下游引物:5’-AATGAATGCATTGAGTATATTTGAGT-3’(SEQ ID NO:7))对T0代转基因阳性植株的叶片DNA进行扩增。结果显示有4株转基因苗发生了目标碱基替换,即SBEIIb第16外显子的第56位碱基A替换为G,同时第55位碱基A也被替换为G,这4株转基因植株均为纯合突变(图4B)。进一步分析发现,第55位碱基A突变为G为同义突变(AGA突变为AGG),不改变所编码的氨基酸。因此,我们在ZH11背景下获得了4株SBEIIb目标氨基酸替换的纯合突变株。利用尿素糊化半籽粒-碘液显色法,对上述纯合突变体T1代成熟籽粒进行分析,ZH11在3.0M尿素中已开始发生糊化,而突变体在尿素浓度高达7.0M时才开始糊化。该结果表明,ZH11背景下的SBEIIb目标碱基替换产生了与MY03一致的突变表型,证实了SBEIIb新等位基因即为MY03籽粒抗尿素糊化突变表型产生的调控基因。
6、稻米的外观品质分析
为比较突变体MY03的外观品质与野生型日本晴相比是否发生变化,本发明随机挑选30粒糙米和精米,使用游标卡尺测量其粒长、粒宽和粒厚发现,MY03糙米的粒长与野生型相比略微降低,粒宽与粒厚无明显变化,MY03精米的粒长、粒宽和粒厚与野生型均无明显差异(图5A,B,C)。利用分析天平称量糙米和精米的千粒干重发现,MY03糙米千粒重略微低于日本晴,精米千粒重与日本晴无明显差异(图5D)。利用万深自动考种仪(SC-E)对精米的垩白性状和透明度进行分析发现,MY03的垩白性状与野生型日本晴差异不大,垩白粒率均在10%以下(图5E)。根据中华人民共和国国家标准(大米GB1354-2009),MY03可达到一级粳米垩白粒率的质量指标。中华人民共和国国家标准(大米GB1354-2009)将稻米透明度值划分为1-5五个级别,分析仪器所得出的数值越小,籽粒的透明程度越高。MY03精米透明度与日本晴一致,透明度值均为1,为完全透明的籽粒(图5F)。以上结果表明,突变体MY03的稻米外观品质未发生较大的变化,其突变位点对稻米的外观品质影响较小。
7、米饭的消化特性分析
米饭的抗性淀粉含量以及消化动态反映了稻米的消化特性,本研究利用爱尔兰Megazyme公司抗性淀粉检测试剂盒(K-RSTAR)比较了日本晴和突变体MY03米饭消化16h后的RS含量,操作如下:
(1)称取1g精米于10mL离心管中,加1.5mL去离子水,用锡箔纸封住管口,静置20min;
(2)静置后于电饭锅正常蒸煮模式下蒸煮,煮熟后保温20min;
(3)称取250mg熟米饭于10mL离心管中,玻璃棒研磨充分;
(4)加4mL酶液(10mg/mLα-胰淀粉酶,其中含有3U/mL(AMG)颠倒摇匀至熟米饭不沾管壁;
(5)然后置于摇床中200rpm酶解16h;
(6)酶解结束后立刻以1000g瞬间离心,并加4mL无水乙醇停止反应,然后以6000g离心5min,转移全部上清至50mL容量瓶;
(7)在沉淀中加2mL无水乙醇,于涡旋振荡器上轻轻振荡至管壁无沉淀,再加2mL无水乙醇冲洗管壁,6000g离心5min,转移全部上清与(6)中收集的上清合并,用100mM NaAC(pH 4.5)缓冲液定容,颠倒混匀;
(8)取2mL上述定溶液于2mL离心管中,加20μL AMG原液(3000U)混匀,置于50℃混匀仪中500rpm酶解30min,该步骤酶解的样品即为非抗性淀粉;
(9)在每个样品的沉淀中加一个磁力搅拌转子和2mL 2M KOH,于磁力搅拌器上冰浴碱解30min;
(10)碱解后加8mL 1.2M NaAC(pH 3.8)缓冲液,颠倒混匀;
(11)取2mL上述样品于2mL离心管中,加20μL AMG原液(3000U)混匀,置于50℃混匀仪中500rpm酶解30min,该步骤酶解样品即为抗性淀粉;
(12)分别取步骤(8)和(11)中酶解后的样品35μL于试管中,加1mL GOPOD,用去离子水作空白,用D-葡萄糖标准液作标样,于涡旋振荡器上振荡混匀,并将试管置于50℃水浴锅反应30min,期间每5min振荡一次;
(13)反应结束后,在试管中加1mL去离子水,振荡混匀,静置冷却5min;
(14)用分光光度计测定样品在波长为510nm下的吸光度。
同时利用体外模拟人体消化系统NutriScan GI20结合消化酶(α-淀粉酶、胃蛋白酶和胰蛋白酶)模拟米饭在人体内的消化动态(0-300min)。将煮熟的米饭样品放入模拟咀嚼器(Zyliss,Zurich,Switzerland)中,模拟咀嚼样品20次,使米饭形成大小约为2-3mm的细颗粒。将100mg咀嚼过的细颗粒状样品放入体外模拟人体消化系统(Nutri Scan GI20,National Instruments,Australia)的样品杯中,将样品杯置于37℃加热板块中,按仪器的操作,在特定时间点,加入一定量的α-淀粉酶、胃蛋白酶、胰酶和淀粉葡萄糖苷酶,这些酶事先溶解在缓冲溶液(0.02MNaOH和0.2M NaAC)中。消化完成后,用葡萄糖分析仪(GlucoseAnalyser,GM9,Analox,UK)分别测量样品在15min、60min、120min、180min、240min和300min产出的总葡萄糖量。消化动态数据处理使用一级动力学方程y=a*[1-exp(-b*x)],其中x为消化时间,y为在x时间内葡萄糖产生量(g/100g样品),a为300min后最终生成的葡萄糖量,b为消化常数。根据a和b,计算米饭的总葡萄糖生产量(Total glucose production,TGP)、活跃消化时间(Active digestion duration,ADD)和葡萄糖生成速率(Glucose productionrate,GPR),其中TGP=0.95*a、ADD=LN(0.05)/(-b)、GPR=TGP/ADD。结果显示,MY03(2.29%)米饭的抗性淀粉含量显著高于日本晴(1.06%)。MY03(358mg g-1)米饭的总葡萄糖生产量低于日本晴(383mg g-1),而MY03(198min)活跃消化时间显著高于日本晴(112min),从而导致MY03(1.82mg g-1min-1)的葡萄糖生成速率显著低于日本晴(3.45mg g-1min-1)(表1)。上述米饭的消化特性数据表明该SBEIIb等位变异能显著提高米饭的RS含量和降低葡萄糖生成速率,是一个优质的高RS稻米资源。
表1米饭的抗性淀粉含量和消化动态参数
数值为平均值±标准偏差,由Student’s t-test计算WT和MY03之间的差异性(*p<0.05,**p<0.01)。RS:米饭消化16h的抗性淀粉含量;TGP:总葡萄糖生产量;ADD:活跃消化时间;GPR:葡萄糖生成速率。
序列表
<110> 扬州大学
<120> 一种调控抗性淀粉含量的水稻SBEIIb等位基因及其在育种中的应用
<160> 7
<170> SIPOSequenceListing 1.0
<210> 1
<211> 11338
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
gcgcacaccc acacaccgac caccaggcag cgcctcctcg ctttggctct cgcgtgagga 60
gggtttaggt ggaagcagag cgcgggggtt gccgggggat ccgatccggc tgcggtgcgg 120
gcgagatggc ggcgccggcg tctgcggttc ccgggagcgc ggcggggcta cgggcggggg 180
ccgtgcggtt ccccgtgcca gccggggccc ggagctggcg tgcggcggcg gagctcccga 240
cgtcgcggtc gctgctctcc ggccggagat tccccggtaa tgatcccgcg ccaccttgtt 300
gttctcgtcc gccgcgacgc gccacgcgtg cgcctcggct cgaccaggtg gcgtgccgtg 360
tcgcgtcgcg ccgcgggtgc ggtaaatcgt cgtgatcttc attttggttt tgcctgcgtc 420
tcgtgtccag gtgccgttcg cgtggggggt tccggggggc gcgtggccgt gcgcgcggcg 480
ggcgcgtcag gggaggtgat gatccccgag ggcgagagcg acgggatgcc ggtttcagca 540
ggttcagacg atctgcaggt gcgtgatcat gattcatcct gccttgacaa tactattgct 600
acatatctaa cagcattttg catgtcatgt ttgaaaatgg agacatgttc ttgtgcagtg 660
ctaccttacg agatcatata catttcacgc agtccacgtc ttgagacgtg ttgatttctg 720
tttgtgacgg gattatgata tattcagtga ttcagtcctt ttttatgtac ttagtccata 780
ctagttgtct gcgtgcataa tgataatatg atctaggtct agagcaccct atatgaggcc 840
tcaaagggca caccatatgc gtaggattga tctagagtct tgcttgttgt cgctcattcc 900
gttcagttgt cattatgttt tacttctgaa cattttatct cttgtacatt gttgtagttg 960
ccagccttag atgatgaatt aagcacggag gttggagctg aagttgagat tgagtcatct 1020
ggagcaagtg acgttgaagg cgtgaagaga gtggttgaag aattagctgc tgagcagaaa 1080
ccacgagttg tcccaccaac aggagatggg caaaaaatat tccagatgga ctctatgctt 1140
aatggctata agtaccatct tgaatatcgg tatggctttc cgcccttctt tatagatcac 1200
atagtttcat taggttatgt ttttattatt ctccatctgt tcagtctttc acaagatgta 1260
tgcaaataac tacttacgtg tcacagctga tgagactctt gaagttttat ctttagtcct 1320
gcatatatac tctgatctgt tctcaaatgg aagttgtttt agaaatcatt cagtcatgct 1380
ttaaaattta acaagagtcc taaactacta atgtgcgaaa aaaaaataga tttaccatta 1440
taagaatttt taagtcaaca tatagttaga aaaaaaaagt taatggtcaa atgagtatat 1500
tggacataca tagtaaatag tactaataat acatagatag tactcttcac agtggttctg 1560
ggttggggcg gtactgcaaa ccaaacaaaa gagtgatagg atggcgtcag aactaagaaa 1620
tgcaatatta attttgcaaa tatttaatct gaaagttatc tatgatgtaa aatacataat 1680
gggttttaga ccttgctgac gtctgtcaac tggacaacaa ctttatttcc tttcttttat 1740
atcattgtta tactccctct gtcctagaaa aaacaacttt cgtacgtgaa cttggaaagt 1800
tgatttattt tatgatggag ggagcataga tgaagtcaat atgcccatgc taactctact 1860
agaaacctga ttatgcttta attgattgaa tacagatata gcctatatag gagactgcgt 1920
tcagacattg atcagtatga aggaggactg gaaacatttt ctcgcggtta tgagaagttt 1980
ggatttaatc acaggtatag tttcttttac attgtaaccg cacatcactt ctctgggtgt 2040
aacaaaataa ctcagtgata ttcatccttc ttcccatgta gtgctgaagg tgtcacttat 2100
cgagaatggg ctcccggggc acatgtatgt tcttttgata gtctcaccat ttacctaaaa 2160
tgctacatta ttagttccat atttctagta catgtagata taattaccat gccatatatt 2220
ctgagatatt ccagattgta tggaatttat aattctcaga tgaacaactt gtcaatgtac 2280
ttgactagac gttgttaaga actaattttt gcttccaatt gcagtctgca gcattagtag 2340
gtgacttcaa caattggaat ccaaatgcag accgcatgag caaagtaagc ccacaggctc 2400
acaaatatgc ccttcctttc ttcaaaaata ttttgtttac atttctttaa ctaatttatt 2460
tggttatagc attggtttca tctatactct agagttgagc tagtgatata tcatgcgaac 2520
gcatgtatag cgcagtggta aacctgactc gaacctggat tttcgcacaa aaaaaattaa 2580
atccctcctt gcaaatgctg gcagaagcgt gctccccatg tgaggttgag caatcccggt 2640
tgtgcgatgc gttcaggggt ggggttctca acttagcttc atgctgcggg ccatcttaac 2700
ccgccgattg agtttttata tatatatatc atgcacactc ttatatgtta ttgactgttg 2760
ccatgagtta actgttagtt atcggttttg atgctactgt agtggtagta cattaatgtt 2820
gtgtatgtta catgtacatt ctttagtacc tagatgtatt ttgcagtttt tgtggacttg 2880
ataggagaag atgagtagct tgaggttgta actaataatg catggaccta gtgttgtaat 2940
tccatatgat tggtaagttg gtagttaccg atgctcatat ctaatgcaac tgtttccaga 3000
atgagtttgg tgtttgggag atttttctgc ctaacaatgc tgatggctca tctcctattc 3060
cacatggctc acgtgtaaag gtaattcctt atcctgcttg gccagtgttt tagctctggg 3120
cctgtgagca tctccctgta tcaagcatgc cttttgtcat ctaggtgcga atggaaactc 3180
catctggtat aaaggattct attcctgcct ggatcaagta ctctgtgcag gccgcaggag 3240
aaatcccata caatggaata tattatgatc ctcctgaaga ggtacctccc cccccccccc 3300
ccaacacgtg cacacgccat ttgaatatgt tcttgttcct tgaaaaaagt gcttcatttg 3360
tacttgtgaa tacttctgta atgtccttgg actcttggaa atttctcttt ttgttggcat 3420
ctttagattg ttctttttat ctagttatta aaatgtttta tatgactatg caggagaagt 3480
acatattcaa gcatcctcaa cctaaaagac caaagtcatt gcggatatac gaaactcatg 3540
ttggaatgag tagcacggta atgcatcttt tcttttaatt actagccata tacagttcaa 3600
gttaacttac attctagaat tatattaaag tgctgttagc cttaattttt ataactgtct 3660
acttctaagc agtgataagt tgttacaagt ttcctctgta ataatatatt aatacaatca 3720
gcacattttt tgcacattcg tcaacaatta gccgtttata taacctttgc tggatggttg 3780
attcaaatat gatgttttga aacaaattac tgatatttac actgatataa tttttgattt 3840
atgttttctg ctaacatagg gtatatttaa ttaatctatg tttgtatttc ttacaaacgg 3900
gtacggttat tattacataa taaaaaatag cgaataacac catgcctacc atttgtatac 3960
attttttttt gggcgtctgt tggtaattta tgattttcct gtactatgct ggctggtatt 4020
aattttaata ttgcaatcag gagccaaaga tcaacacgta tgcaaacttt agggatgagg 4080
tgcttccaag aatcaaaaag cttggataca atgcagtgca aataatggca attcaagagc 4140
atgcatatta tggaagcttt gggtgatcaa attacatatt gctggattta tttcttgtct 4200
ttgcagttta catatatatg cttttagact ttatctttca tcttccttgt tccctttgta 4260
taatgtgata aactgcttca tctttcttat catctttatc ttccttttat actcatgaaa 4320
caatgctgtt actgacccct tttgtttccc cttttgtttt ggtacaattg aactgtaaca 4380
aatgacattc gccacctttt gaatgaatgg ctattgagat aattttcaat tgtcaattaa 4440
gacttttatt atgggtgagc aatgcagatt ttttactttt gtttcaggta ccatgtcacc 4500
aatttctttg caccaagtag tcgtttcggg accccagaag atttaaagtc attgattgat 4560
aaagctcatg agcttggttt agttgtgctc atggatgttg ttcacaggta attaattact 4620
ccctctgctc atttttataa ggcatatttt aattttagaa agtatttatg actgaaattt 4680
gacaattaat ttctcataaa atatatttat catagctata aaaccaatat aatgtgaaaa 4740
tactttgagg tatgaatcta gtggtatatg ttttatactc taaatatact tgtatatcaa 4800
ctaattattg gtcaaaatct ttcgagtttg actttctgaa atcaaaatat gcgttataaa 4860
agtgaacaga atgagtattt gttaactggt ctttcatatc taagttagta aggcgtggcc 4920
acagagataa aagaaagtcc ttataatata gaagaaggct gtggtggtaa gtctgccatt 4980
ataccaccac ctcttgagtt tgtgggatct tctgtggata aatagatcat ttatgcatgt 5040
ggctaggctg tttgtggatt aatgtttggt tccaaattgc ttacaaatta caattttttt 5100
cctgagaagt tcatgcatag ttcatttcat ttttatgata ttttttaaaa aaaatccact 5160
tctactcatg ggtatttagc taacggtatg attctctgtt ctataaaaaa aaagtacatc 5220
tagccctcat gttggtgaac cctaattatg actaatctac aaccgtagat cttctctctt 5280
ttatgcaaca tgtttaactt gaattattgc agtataagag ttttcacaat ttaaattggc 5340
tagcattcga gtttatctta ggatgggaag aaaatgaatt aaacagtttc attcatattt 5400
tctattcaat tattgaattt gagaattcaa attaatattt tattcgtgac atcttttgtt 5460
ggctttctgg tggacaacaa ctagaactaa tttacatgcc aaaatttgtc aatcaccatg 5520
gcccaaagtt tagctcattg gcctctgggt tctaagccct tttggtttca aatattaacc 5580
tttgtgttga tttactaagt ccccttttag ctgcatacct aatagcttta cctttgcccc 5640
ttttttttcc ttttgcagcc atgcgtcaaa taatacccta gatgggttga acggttttga 5700
tggtacagat acgcattact ttcatagtgg ttcacgcggc catcattgga tgtgggattc 5760
tcgccttttc aactatggga attgggaagt aaggaacacg ttaatcgcta cttcccttta 5820
caaagtatca ttattcatat gcattaatct ctattgtgtt tgccaacatt tttatcctta 5880
tacaggttct aagatttcta ctatccaatg caagatggtg gctcgaggag tataagtttg 5940
atggtttcag atttgacggt gtaacctcaa tgatgtacac tcatcatgga ttacaagtat 6000
attgcttcgt tttcattact ttatcttcac taccttctta attatgatgg ggtcaaacct 6060
agcagttgct atctgttaca atttcatatt tatttgcgtc ataaccttta tcttatgatt 6120
ccaccctcta ttttaggtag catttacggg gaactacagt gaatactttg gatttgccac 6180
tgatgctgat gcagtagttt acttgatgct ggtaaatgat ttaattcatg gactttatcc 6240
tgaggccata accatcggtg aagatgtaag tgctgactat atttgtcttt atgtactcat 6300
ttctttagca tgcatttata gcaatatttt ggcatctaga catgcttaag taagaagttg 6360
aaagcttgca ctgcatatta cgctaatgtc agttttgcct tctctagttc ataatgttgt 6420
atacccaaag cttagctttc caatcttagt gttacatatt tactattaac cccatcttga 6480
tagttgatca aattatgtcg ctggttcaag aaattgtttt ccttgggctt agaaaacagc 6540
aaggagtaat gatagttacc aaagattagt gaccacctgg tctctagtac agaagttgac 6600
catcaatgtt atacgagtat gttagtaaag aattatacaa atgtattgtt atgaaaaaaa 6660
ttcgtgacga atataatggc accatttgca tgtgaataac ctgattgttt ttatatatta 6720
ttctgagtta gagaagtttg aaatatgaac gcgtgcattc taaaaattta catatttctg 6780
aacaaaaaga gtacattttt tctgtaaaaa aagagagtat aattgtgtcc tcaaaaacag 6840
ctgtgcatgt tccatggtgc catactgtca taaccttttc ttctcttggg aaatccagcc 6900
tccctggaag tttgtgaagt acaaaaagca agttatacag tagctctgta tggtctacat 6960
ttttttttgg gaatacgcaa aatgcgtatc ttttgtatta agatcgggaa aagttgttag 7020
aacaggcaca gcaactatgt acagggagaa aggaagaaaa atgggtgtgg gaaacatata 7080
gtaatatctc gcggaacaat ttgatgacca aatgctccac ctagatgcgc gctcctgcct 7140
ccctccaggg cgcggcctcc gatgccacta gctccgctat ctcctgcatg gtcttgcatt 7200
gaccattgaa aacccggttt ttccactcct tttgtatcgt ccaggccacc aggatatgat 7260
gatggtgtca aaccctcatc tttgggccgc tatgattttg tctcggctgt cgcaccacca 7320
tacgaggaag ggctgctcat ttgatgggaa gcaatgggat tggccaatgc cgcatagaac 7380
tgtccaccat agctgtagtg tttctggaca ttggattaga aggtggtcaa ttgactccac 7440
actttgatca cagaagacac acttctgcag accacttgct cagcctgtcg cctgtatggt 7500
ctagagttgc aatagtagta acaatactct aaatcccata cgtaaaccta actctattgc 7560
aaaagataac ttaatttcaa ctgttctgtg gttaaatggc atataactac ttcaggtcag 7620
tggaatgcct acatttgccc ttcctgttca agatggtggg gttggttttg attatcgcct 7680
tcatatggct gttcctgaca aatggattga actcctcaag taagtgtttc aaattttggt 7740
taatagaaat actaattaca tatctaatga acataaactg ttttcttgtg ttaaattctt 7800
ttttgtatac attggtttcc ttgaccatga tatctctatt tctcttctga agttaattgg 7860
cataactcct gcccttgatt tttttttaaa aaaaaaaaag atcactcata tatagggtta 7920
taggcattta ccaatgtgat gtgctggatt tggtcatatg aacagtggtc gtgtgagttc 7980
cattattagt ataaaataag atgcaactac caacttatag agtttgactt ggcaacgtta 8040
catttttaat gctcctttat tatgtggtat aaaaatgccc atgctcatcg tatctcactt 8100
ttttgtcagc tgtttagctt ttatttagaa actgggaaag attgttattt tgtatccttt 8160
aattgaaaga tgtggtgcca gagtttcaat ttttatttta cacaggcaaa gtgatgaatc 8220
ttggaagatg ggtgatattg tgcacacact gactaacaga gggtggtcag agaagtgtgt 8280
tacttatgct gaaagtcatg atcaagcact agttggtgac aaaactattg cattctggtt 8340
gatggacaag gttaccacac atcaatctct aggttttagt gttttactga ttaggactgt 8400
aactaatggc tctttaatat atccttgata attatttgta ctgattttac tcaaatatac 8460
tcaatgcatt cattgtggat agtaagaacg gtatgaaaac cacagctaga tcagtgtttt 8520
aagttggata gctttaggct ttctgcagtc tacacccagt ttttcttgta ttagttgcaa 8580
catgtattta tgaattttct attctttact agttctttgg aaaatgttat ttccttgttt 8640
taatacttaa aacatatttg tgtatcagtg tatcaccgta ctattcttga gataatagag 8700
ttgttctttg agttttaatt gttgagaaaa tgcatataca tgcaggatat gtatgaattt 8760
taatgaatac tggcatgaga acctcacata ctgtacttat gcaggatatg tatgatttta 8820
tggctctgga cagaccggca acacctagca ttgatcgtgg aatagcattg cataaaatga 8880
ttagacttat cacaatgggg ttaggaggag aaggctatct taactttatg ggaaatgagt 8940
tcggacatcc tggtgagatt taataacaat cacacaccat gtttgtttct tattttatat 9000
caacttctca tttgatgaca atggttcttg tcacctagta ttttctggga aacaattggc 9060
ttaaaggcca ttgatctacc ttttattttg cagaatggat tgattttcca agagctccac 9120
aagtacttcc aaatggtaaa ttcatcccag ggaataacaa cagttatgat aaatgccgtc 9180
gaagatttga cctggtaact ttctttgctc tgtcacctga ttattgatga aaataagttc 9240
tgcattccct ggattcaatt tagatatctt tttaccccct tctgcagtgt tttaatcatg 9300
aaggtcccat tatgacaatt catctaattc atcaattcct ttttattcac attaacattt 9360
gtgcgaggcc agcaacattg aaccatgatt ttgttatgga tcagtaagtt ttttcattgc 9420
tttgtcaggg tgatgcggac tatcttaggt atcgtggcat gctagagttt gaccgcgcga 9480
tgcagtctct cgaggaaaaa tatggggtat gttgtattta tcacttatct ttttctgggg 9540
gttgcacaaa tatgagccat tatgcttatg tcctgttact gatacttgct atgtcaacta 9600
tttatccaaa accacttgaa atatatggtg aaatgacgag ctgcacttgt gcattatgca 9660
attatgtcca atactcttag gaatagctcc cattgtcatt atctaggtgc atcattatgt 9720
tttctcatgt cttccaatca ttgcaatagt aatcaatact gttaaataat ggacagcttt 9780
gttcgatgcc aattaaatta ttttgtcttg tttgcatggg ctatgtggac agtaatttta 9840
tatttttatt gatgaataaa tgcattatca ataatataat gcatatattt cccttcatag 9900
aatttctttt ttgtatttat aggaaaaact ttgcaagtaa aatgatggtt ttgttttaac 9960
attgctctga cactgatggg ggttttctta tggctatgtt atttcagttc atgacatcag 10020
accaccagta catatctcga aagcatgaag aggataagat gattatattt gagaagggag 10080
atctggtatt tgtgttcaac ttccattgga gtaacagcta ttttgactac cgtgttggtt 10140
gtttaaagcc agggaaatat aaggtatgca gctatttgtg cattgcccgt gatcaattat 10200
agtttttgtt cactaggtaa ggctttaccg accttactgt taaccaagta gaaaatggca 10260
aaatgccaaa aaaatcggat ttgtatgatt tctgcccaag aataaaatag attgacaact 10320
atgcttgcat atgcaccccg gtggtcaagg agttgccttt gttagcgata gtcctaggtc 10380
tggatcctca aattgcaacc attcattttt ttttcctatt cagattaaaa aatattgaat 10440
gtggactaaa aaaatttcat gttatgtacg tctccaattg tactgtaact aatatttctc 10500
cctgggtctc tgggatatgt caagtgccat tatttattga ttggtgttaa atcaggaatt 10560
acgagcaaag aagttcttgg taatcatcaa cacttcatat ttacctacac tagtcgtgcc 10620
cattatgttg accctgccat agtttactcc ttccatatat ttgaaaataa ccagtaaaaa 10680
ttactcaaaa gtcctctagt cagttttgca gcgcgttcat gcttcttaca gacgaactgg 10740
atgtgttggt tgctgaactg caaaataatt tatgttgctt tctatcaggt ggtcttggac 10800
tcagatgctg gactctttgg tggatttggc aggatccatc acactgcaga gcacttcact 10860
gccgtaagtc ttgctcagat gaaattgcgt accgtatatt gtgtgctctt tattaacctc 10920
tgttgtgctc attccttgca ggattgttca catgacaaca ggccctactc gttctcagtt 10980
tattctccta gcagaacctg cgttgtctat gctccagcgg aatgagaaca ccaagaggca 11040
gcatgcaagt gtgtgcggct gctagtgcga aggagcaaga aaaactagtt gccagcaatc 11100
tgtgaacggc tttcctaggt tctgcttcga tgaatgccgg atagactaga cagcttgctt 11160
ttgtgctttg cgctcccaat ttgtagtttt agtttgtgag ggaaagaaac gtttatttgt 11220
aattatctat ggctgtcgaa cggcgacgaa accatgaacc ccgtatattt gttggtaccg 11280
ttcgaactgc cagttataca tagttctgca cttctgtaca tcttgtgatg cttgaatc 11338
<210> 2
<211> 2478
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 2
atggcggcgc cggcgtctgc ggttcccggg agcgcggcgg ggctacgggc gggggccgtg 60
cggttccccg tgccagccgg ggcccggagc tggcgtgcgg cggcggagct cccgacgtcg 120
cggtcgctgc tctccggccg gagattcccc ggtgccgttc gcgtgggggg ttccgggggg 180
cgcgtggccg tgcgcgcggc gggcgcgtca ggggaggtga tgatccccga gggcgagagc 240
gacgggatgc cggtttcagc aggttcagac gatctgcagg tgccagcctt agatgatgaa 300
ttaagcacgg aggttggagc tgaagttgag attgagtcat ctggagcaag tgacgttgaa 360
ggcgtgaaga gagtggttga agaattagct gctgagcaga aaccacgagt tgtcccacca 420
acaggagatg ggcaaaaaat attccagatg gactctatgc ttaatggcta taagtaccat 480
cttgaatatc gatatagcct atataggaga ctgcgttcag acattgatca gtatgaagga 540
ggactggaaa cattttctcg cggttatgag aagtttggat ttaatcacag tgctgaaggt 600
gtcacttatc gagaatgggc tcccggggca cattctgcag cattagtagg tgacttcaac 660
aattggaatc caaatgcaga ccgcatgagc aaaaatgagt ttggtgtttg ggagattttt 720
ctgcctaaca atgctgatgg ctcatctcct attccacatg gctcacgtgt aaaggtgcga 780
atggaaactc catctggtat aaaggattct attcctgcct ggatcaagta ctctgtgcag 840
gccgcaggag aaatcccata caatggaata tattatgatc ctcctgaaga ggagaagtac 900
atattcaagc atcctcaacc taaaagacca aagtcattgc ggatatacga aactcatgtt 960
ggaatgagta gcacggagcc aaagatcaac acgtatgcaa actttaggga tgaggtgctt 1020
ccaagaatca aaaagcttgg atacaatgca gtgcaaataa tggcaattca agagcatgca 1080
tattatggaa gctttgggta ccatgtcacc aatttctttg caccaagtag tcgtttcggg 1140
accccagaag atttaaagtc attgattgat aaagctcatg agcttggttt agttgtgctc 1200
atggatgttg ttcacagcca tgcgtcaaat aataccctag atgggttgaa cggttttgat 1260
ggtacagata cgcattactt tcatagtggt tcacgcggcc atcattggat gtgggattct 1320
cgccttttca actatgggaa ttgggaagtt ctaagatttc tactatccaa tgcaagatgg 1380
tggctcgagg agtataagtt tgatggtttc agatttgacg gtgtaacctc aatgatgtac 1440
actcatcatg gattacaagt agcatttacg gggaactaca gtgaatactt tggatttgcc 1500
actgatgctg atgcagtagt ttacttgatg ctggtaaatg atttaattca tggactttat 1560
cctgaggcca taaccatcgg tgaagatgtc agtggaatgc ctacatttgc ccttcctgtt 1620
caagatggtg gggttggttt tgattatcgc cttcatatgg ctgttcctga caaatggatt 1680
gaactcctca agcaaagtga tgaatcttgg aagatgggtg atattgtgca cacactgact 1740
aacagagggt ggtcagagaa gtgtgttact tatgctgaaa gtcatgatca agcactagtt 1800
ggtgacaaaa ctattgcatt ctggttgatg gacaaggata tgtatgattt tatggctctg 1860
gacagaccgg caacacctag cattgatcgt ggaatagcat tgcataaaat gattagactt 1920
atcacaatgg ggttaggagg agaaggctat cttaacttta tgggaaatga gttcggacat 1980
cctgaatgga ttgattttcc aagagctcca caagtacttc caaatggtaa attcatccca 2040
gggaataaca acagttatga taaatgccgt cgaagatttg acctgggtga tgcggactat 2100
cttaggtatc gtggcatgct agagtttgac cgcgcgatgc agtctctcga ggaaaaatat 2160
gggttcatga catcagacca ccagtacata tctcgaaagc atgaagagga taagatgatt 2220
atatttgaga agggagatct ggtatttgtg ttcaacttcc attggagtaa cagctatttt 2280
gactaccgtg ttggttgttt aaagccaggg aaatataagg tggtcttgga ctcagatgct 2340
ggactctttg gtggatttgg caggatccat cacactgcag agcacttcac tgccgattgt 2400
tcacatgaca acaggcccta ctcgttctca gtttattctc ctagcagaac ctgcgttgtc 2460
tatgctccag cggaatga 2478
<210> 3
<211> 825
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 3
Met Ala Ala Pro Ala Ser Ala Val Pro Gly Ser Ala Ala Gly Leu Arg
1 5 10 15
Ala Gly Ala Val Arg Phe Pro Val Pro Ala Gly Ala Arg Ser Trp Arg
20 25 30
Ala Ala Ala Glu Leu Pro Thr Ser Arg Ser Leu Leu Ser Gly Arg Arg
35 40 45
Phe Pro Gly Ala Val Arg Val Gly Gly Ser Gly Gly Arg Val Ala Val
50 55 60
Arg Ala Ala Gly Ala Ser Gly Glu Val Met Ile Pro Glu Gly Glu Ser
65 70 75 80
Asp Gly Met Pro Val Ser Ala Gly Ser Asp Asp Leu Gln Val Pro Ala
85 90 95
Leu Asp Asp Glu Leu Ser Thr Glu Val Gly Ala Glu Val Glu Ile Glu
100 105 110
Ser Ser Gly Ala Ser Asp Val Glu Gly Val Lys Arg Val Val Glu Glu
115 120 125
Leu Ala Ala Glu Gln Lys Pro Arg Val Val Pro Pro Thr Gly Asp Gly
130 135 140
Gln Lys Ile Phe Gln Met Asp Ser Met Leu Asn Gly Tyr Lys Tyr His
145 150 155 160
Leu Glu Tyr Arg Tyr Ser Leu Tyr Arg Arg Leu Arg Ser Asp Ile Asp
165 170 175
Gln Tyr Glu Gly Gly Leu Glu Thr Phe Ser Arg Gly Tyr Glu Lys Phe
180 185 190
Gly Phe Asn His Ser Ala Glu Gly Val Thr Tyr Arg Glu Trp Ala Pro
195 200 205
Gly Ala His Ser Ala Ala Leu Val Gly Asp Phe Asn Asn Trp Asn Pro
210 215 220
Asn Ala Asp Arg Met Ser Lys Asn Glu Phe Gly Val Trp Glu Ile Phe
225 230 235 240
Leu Pro Asn Asn Ala Asp Gly Ser Ser Pro Ile Pro His Gly Ser Arg
245 250 255
Val Lys Val Arg Met Glu Thr Pro Ser Gly Ile Lys Asp Ser Ile Pro
260 265 270
Ala Trp Ile Lys Tyr Ser Val Gln Ala Ala Gly Glu Ile Pro Tyr Asn
275 280 285
Gly Ile Tyr Tyr Asp Pro Pro Glu Glu Glu Lys Tyr Ile Phe Lys His
290 295 300
Pro Gln Pro Lys Arg Pro Lys Ser Leu Arg Ile Tyr Glu Thr His Val
305 310 315 320
Gly Met Ser Ser Thr Glu Pro Lys Ile Asn Thr Tyr Ala Asn Phe Arg
325 330 335
Asp Glu Val Leu Pro Arg Ile Lys Lys Leu Gly Tyr Asn Ala Val Gln
340 345 350
Ile Met Ala Ile Gln Glu His Ala Tyr Tyr Gly Ser Phe Gly Tyr His
355 360 365
Val Thr Asn Phe Phe Ala Pro Ser Ser Arg Phe Gly Thr Pro Glu Asp
370 375 380
Leu Lys Ser Leu Ile Asp Lys Ala His Glu Leu Gly Leu Val Val Leu
385 390 395 400
Met Asp Val Val His Ser His Ala Ser Asn Asn Thr Leu Asp Gly Leu
405 410 415
Asn Gly Phe Asp Gly Thr Asp Thr His Tyr Phe His Ser Gly Ser Arg
420 425 430
Gly His His Trp Met Trp Asp Ser Arg Leu Phe Asn Tyr Gly Asn Trp
435 440 445
Glu Val Leu Arg Phe Leu Leu Ser Asn Ala Arg Trp Trp Leu Glu Glu
450 455 460
Tyr Lys Phe Asp Gly Phe Arg Phe Asp Gly Val Thr Ser Met Met Tyr
465 470 475 480
Thr His His Gly Leu Gln Val Ala Phe Thr Gly Asn Tyr Ser Glu Tyr
485 490 495
Phe Gly Phe Ala Thr Asp Ala Asp Ala Val Val Tyr Leu Met Leu Val
500 505 510
Asn Asp Leu Ile His Gly Leu Tyr Pro Glu Ala Ile Thr Ile Gly Glu
515 520 525
Asp Val Ser Gly Met Pro Thr Phe Ala Leu Pro Val Gln Asp Gly Gly
530 535 540
Val Gly Phe Asp Tyr Arg Leu His Met Ala Val Pro Asp Lys Trp Ile
545 550 555 560
Glu Leu Leu Lys Gln Ser Asp Glu Ser Trp Lys Met Gly Asp Ile Val
565 570 575
His Thr Leu Thr Asn Arg Gly Trp Ser Glu Lys Cys Val Thr Tyr Ala
580 585 590
Glu Ser His Asp Gln Ala Leu Val Gly Asp Lys Thr Ile Ala Phe Trp
595 600 605
Leu Met Asp Lys Asp Met Tyr Asp Phe Met Ala Leu Asp Arg Pro Ala
610 615 620
Thr Pro Ser Ile Asp Arg Gly Ile Ala Leu His Lys Met Ile Arg Leu
625 630 635 640
Ile Thr Met Gly Leu Gly Gly Glu Gly Tyr Leu Asn Phe Met Gly Asn
645 650 655
Glu Phe Gly His Pro Glu Trp Ile Asp Phe Pro Arg Ala Pro Gln Val
660 665 670
Leu Pro Asn Gly Lys Phe Ile Pro Gly Asn Asn Asn Ser Tyr Asp Lys
675 680 685
Cys Arg Arg Arg Phe Asp Leu Gly Asp Ala Asp Tyr Leu Arg Tyr Arg
690 695 700
Gly Met Leu Glu Phe Asp Arg Ala Met Gln Ser Leu Glu Glu Lys Tyr
705 710 715 720
Gly Phe Met Thr Ser Asp His Gln Tyr Ile Ser Arg Lys His Glu Glu
725 730 735
Asp Lys Met Ile Ile Phe Glu Lys Gly Asp Leu Val Phe Val Phe Asn
740 745 750
Phe His Trp Ser Asn Ser Tyr Phe Asp Tyr Arg Val Gly Cys Leu Lys
755 760 765
Pro Gly Lys Tyr Lys Val Val Leu Asp Ser Asp Ala Gly Leu Phe Gly
770 775 780
Gly Phe Gly Arg Ile His His Thr Ala Glu His Phe Thr Ala Asp Cys
785 790 795 800
Ser His Asp Asn Arg Pro Tyr Ser Phe Ser Val Tyr Ser Pro Ser Arg
805 810 815
Thr Cys Val Val Tyr Ala Pro Ala Glu
820 825
<210> 4
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
cgagagcctg acctattgca t 21
<210> 5
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
ctgctccata caagccaacc ac 22
<210> 6
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
agcctgtcgc ctgtatggt 19
<210> 7
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
aatgaatgca ttgagtatat ttgagt 26
Claims (10)
1.一种调控抗性淀粉含量的水稻SBEIIb等位基因,其特征在于,在野生型水稻品种日本晴SBEIIb基因基础上,第16外显子上的第56位A被G替换;所述的水稻SBEIIb等位基因其核苷酸序列如SEQ ID No.1所示。
2.根据权利要求1所述的调控抗性淀粉含量的水稻SBEIIb等位基因,其特征在于,其编码区序列如SEQ ID No.2所示。
3.一种用于检测调控抗性淀粉含量的水稻SBEIIb等位基因的引物,其特征在于,所述引物的上游序列如SEQ ID NO:6所示,所述引物的下游序列如SEQ ID NO:7所示。
4.根据权利要求1或2所述的水稻SBEIIb等位基因所编码的蛋白,其特征在于,在野生型日本晴SBEIIb蛋白基础上,第583位氨基酸精氨酸(Arg)被甘氨酸(Gly)替换;所述的水稻SBEIIb等位基因所编码蛋白的氨基酸序列如SEQ ID No.3所示。
5.根据权利要求1所述的水稻SBEIIb等位基因在高抗性淀粉水稻育种中的应用。
6.根据权利要求5所述的应用,其特征在于,所述水稻SBEIIb等位基因提高水稻支链淀粉长侧链的比例。
7.根据权利要求5所述的应用,其特征在于,水稻SBEIIb等位基因提高米饭的抗性淀粉含量并降低葡萄糖生成速率。
8.根据权利要求5所述的应用,其特征在于,水稻SBEIIb等位基因提高稻米的糊化温度。
9.根据权利要求5所述的应用,其特征在于,将权利要求1或2所述的基因目标碱基通过基因编辑技术改变目的水稻的目标碱基,得到目标碱基替换的水稻材料;所述目标碱基替换的水稻稻米尿素糊化抗性高于所述目的水稻。
10.根据权利要求9所述的应用,其特征在于,以序列为SEQ ID NO:4和SEQ ID NO:5的引物对T0代植株进行扩增。
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Citations (2)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
CN102816778A (zh) * | 2012-07-30 | 2012-12-12 | 上海市农业科学院 | 一种水稻淀粉分支酶sbe3基因的突变基因及其用途 |
CN107384946A (zh) * | 2017-07-21 | 2017-11-24 | 上海市农业科学院 | 水稻淀粉分支酶sbe3基因的人工定点突变体及其应用 |
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CN102816778A (zh) * | 2012-07-30 | 2012-12-12 | 上海市农业科学院 | 一种水稻淀粉分支酶sbe3基因的突变基因及其用途 |
CN107384946A (zh) * | 2017-07-21 | 2017-11-24 | 上海市农业科学院 | 水稻淀粉分支酶sbe3基因的人工定点突变体及其应用 |
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Title |
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RUIFANG YANG等: "A single amino acid mutation of OsSBEIIb contributes to resistant starch accumulation in rice", 《BREEDING SCIENCE》, vol. 66, no. 4 * |
SHENGNAN ZHAO等: "Screening and identification of rice non-floury endosperm mutants with different starch components", 《JOURNAL OF CEREAL SCIENCE》, vol. 103 * |
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