CN108624597A - 花生AhGOS1-1基因及其在提高植物抗旱性中的应用 - Google Patents
花生AhGOS1-1基因及其在提高植物抗旱性中的应用 Download PDFInfo
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Abstract
本发明公开了花生AhGOS1‑1及其在提高植物抗旱性中的应用。本发明克隆得到一个花生SNARE蛋白基因AhGOS1‑1,其核苷酸序列如SEQ ID NO:1所示,其所编码的AhGOS1‑1的氨基酸序列如SEQ ID NO:2所示;通过构建AhGOS1‑1基因的过表达载体并转化模式植物拟南芥进行过表达,结果证明转基因拟南芥较野生型植物对ABA的敏感性增加,转基因植物具有更强的耐受干旱胁迫的能力,表明所述花生AhGOS1‑1基因可提高植物的抗旱性能,提高土地利用率,有利于增加作物的产量和农业的可持续发展。
Description
技术领域
本发明涉及生物技术领域,更具体地,涉及花生可溶性N-乙基马来酰亚胺敏感因子附着蛋白受体基因AhGOS1-1及其编码的蛋白在提高植物对干旱和/或渗透胁迫耐受性中的应用。
背景技术
干旱和水分渗透胁迫是限制植物生长、存活、产量以及分布的重要因素之一。植物作为固着生长的生物,面临各种各样的生物和非生物胁迫,高等植物在长期的进化过程中形成了在分子、亚细胞以及生理水平的不同防御体系。在干旱胁迫下,植物激素ABA的含量大量积累进而引起气孔关闭,并且诱导一系列的抗旱相关基因上调表达(Bartels D andSunkar R,Crit Rev Plant Sci,2005,24:23-58; Yamaguchi-Shinozaki K andShinozaki K,Annu Rev Plant Biol,2006,57:781-803)。干旱胁迫下,植物体内存在ABA依赖和ABA不依赖这两条不同的调控通路来调节干旱响应(Yamaguchi-Shinozaki K andShinozaki K,Annu Rev Plant Biol,2006, 57:781-803)。脱落酸(abscisic acid,ABA)作为植物体内一种重要的生长激素,能够调节调节植物生长发育的各个方面,大量的研究报道ABA参与了各种信号途径,尤其在植物对于逆境胁迫抵抗作用的信号通路中发挥重要作用(Yu F,et al., Trends Plant Sci,2015,20:569-575)。植物细胞原生质膜对干旱甚为敏感,质膜透性的大小可以反映细胞受干旱的程度,已知植物体内的ABA含量与植物的生长和抗旱能力相关,因为ABA直接参与维持质膜的结构和功能,并能导致植物体内一系列抵抗干旱的生理反应、形态反应,以提高抗旱性。
膜泡运输作为植物与环境之间物质和信息交流的重要方式,能为改善植物耐受环境胁迫提供物质基础。植物可溶性N-乙基马来酰亚胺敏感因子附着蛋白受体(Soluble N-ethyl-maleimide-sensitive fusion protein attachment protein receptor,缩写为SNAP receptor或SNARE)是一个多功能的蛋白家族,参与了植物细胞中的许多生理过程。SNARE蛋白在介导囊泡的融合过程中作为膜泡运输过程重要的调控因子以及SNARE复合体中的核心蛋白在调控植物响应环境胁迫过程中发挥着重要的作用,而且SNARE蛋白还与植物中多种信号调控途径有着密切的联系(胡重怡和蔡刘体,生物技术通报,2011,7:77-81)。
花生(Arachis hypogaea L.)是一种重要的豆科油料作物,主要栽培在热带和亚热带地区的世界(Sarkar T,et al.,Front Plant Sci,2016,7:935)。在全球范围内,它的种植面积达到20~25万公顷,年产量35~4000万吨。与其它农作物一样,水资源不足也会对花生的生产带来影响,在世界内不同的地方,由于干旱导致的年产量损失大约达到600万吨(Kottapalli K R,et al.,J Proteome Res,2013, 12:5048-5057;Bhatnagar-Mathur Pet al.,Mol Breed,2014,33,327-340)。花生也是旱地作物之一,研究花生的耐旱机制对提高花生的抗旱性,促进花生生产具有重要意义,有研究发现花生ABA转运蛋白基因AhATL1可降低转基因拟南芥对 ABA的敏感性,导致干旱条件下叶片水分散失加快(Ge K,et al.,Front Plant Sci, 2017,8:1150),但目前对花生耐干旱和耐渗透胁迫分子机制的研究仍十分欠缺。尤其是关于花生SNARE蛋白基因在提高植物抗旱性作用方面的研究尚未见报道。
发明内容
本发明所要解决的技术问题是克服上述现有技术的缺陷和不足,提供一种花生可溶性N-乙基马来酰亚胺敏感因子附着蛋白受体基因AhGOS1-1。
本发明的第二个目的是提供由所述AhGOS1-1基因编码的AhGOS1-1蛋白。
本发明的第三个目的是提供所述AhGOS1-1基因或AhGOS1-1蛋白在提高植物对干旱和/或渗透胁迫耐受性中的应用。
本发明的上述目的是通过以下技术方案给予实现的:
花生AhGOS1-1基因,其核苷酸序列如SEQ ID NO:1所示。
花生AhGOS1-1基因编码的AhGOS1-1,其氨基酸序列如SEQ ID NO:2所示。
发明人在已构建花生cDNA文库的基础上,通过EST测序得到一个花生 SNARE蛋白基因序列,再克隆得了到编码该蛋白的完整cDNA序列。其核苷酸序列如SEQ ID NO:1所示,该序列含有一个大小为675bp的开放阅读框(ORF)。开放阅读框编码224个氨基酸残基的蛋白,所述蛋白的氨基酸序列如SEQ ID NO: 2所示。在NCBI数据库中搜索比对发现,该基因与GenBank中来源于花生祖先种野生蔓花生的SNARE蛋白基因(XM_016078250)、狭叶羽扁豆SNARE蛋白基因(XM_019601639)、鹰嘴豆SNARE蛋白基因(XM_004486564)和大豆 SNARE蛋白基因(XM_003531683)的同源性分别达到97%、85%、85%和82%,这类SNARE蛋白也称为高尔基体SNAP受体复合物1-1(Golgi SNAP receptor complex member 1-1),因此将克隆得到的花生SNARE蛋白基因命名为AhGOS1-1。
本发明还请求保护用于扩增AhGOS1-1基因的引物对,包括上游引物F和下游引物R,其核苷酸序列依次如SEQ ID NO:3~4所示。
正向引物(SEQ ID NO:3):
5’-F-TCCCCCGGGAGTCAAATCGGAGCGAGTCATCACC-3’
反向引物(SEQ ID NO:4):
5’-CGAGCTCGCAACCTCAATAAATCCAAGTGTAG-3’
通过转化植物验证AhGOS1-1基因的功能,与对照组比较,发现转基因过表达AhGOS1-1的拟南芥植株对ABA更加敏感,对干旱的抵抗能力明显增强。由此验证了AhGOS1-1基因可以提高植物的抗旱性,并证明该基因参与了ABA依赖的非生物胁迫抗逆调控途径。将花生AhGOS1-1基因应用于植物的遗传转化和农作物基因工程改良,可提高植物对干旱和渗透胁迫的抗性,增强植物抵抗不良环境胁迫的能力。
因此,SEQ ID NO:1所示基因或SEQ ID NO:2所示蛋白的下列应用均在本发明保护范围内。
SEQ ID NO:1所示的花生SNARE蛋白基因AhGOS1-1在提高植物在干旱胁迫和渗透胁迫下的生存和恢复生长的能力,以及该基因在提高植物抗旱性方面的应用。
SEQ ID NO:2所示的花生SNARE蛋白AhGOS1-1在提高植物在干旱胁迫和渗透胁迫下的生存和恢复生长的能力,以及该基因在提高植物抗旱性方面的应用。
具体地,所述应用为构建花生AhGOS1-1基因的过表达载体并转化到植物中。
优选地,所述过表达载体为pBI121-AhGOS1-1。
优选地,所述植物为花生、水稻、玉米、小麦、大豆、油菜、油茶、芝麻、向日葵等重要农作物。
与现有技术相比,本发明具有以下有益效果:
(1)本发明从花生中分离到AhGOS1-1基因,通过转化模式植物拟南芥证实了AhGOS1-1能提高植物抗旱性。这将为植物基因工程育种提供一种具有抗旱性能的候选基因。
(2)将本发明中的AhGOS1-1基因,通过分子生物学等技术构建过表达载体,使之在其它植物中过表达,定向提高某些物种的抗旱性能,为植物的遗传育种和利用提供新的途径。
(3)将AhGOS1-1基因转入到水稻、玉米、小麦、大豆、油菜、油茶、芝麻、向日葵等重要农作物中,则有可能提高这些作物的抗旱性能,提高了土地利用率,有利于增加作物的产量和农业的可持续发展。
附图说明
图1为AhGOS1-1基因的克隆条带电泳图。
图2为植物表达载体pBI121-AhGOS1-1构建示意图。RB:右边界;LB:左边界。
图3为转基因过表达拟南芥AhGOS1-1基因的qRT-PCR检测结果。WT表示野生型,OE-1、OE-2表示不同的转基因过表达株系。
图4为对野生型植物和AhGOS1-1过表达植物进行干旱耐受性检测的结果。
图5为野生型植物和AhGOS1-1过表达植物离体叶片失水速率检测。WT表示野生型,OE-1、OE-2表示不同的转基因株系。
图6为ABA处理下AhGOS1-1过表达植物气孔变化检测结果。A为ABA 处理下对照与AhGOS1-1过表达植物的气孔变化;WT表示野生型,OE-1、OE-2 表示不同的转基因株系。B为ABA处理下对照与AhGOS1-1过表达植物气孔开度的测量结果;WT表示野生型,OE-1、OE-2表示不同的转基因株系。
具体实施方式
以下结合说明书附图和具体实施例来进一步说明本发明,但实施例并不对本发明做任何形式的限定。除非特别说明,本发明采用的试剂、方法和设备为本技术领域常规试剂、方法和设备。
除非特别说明,以下实施例所用试剂和材料均为市购。
实施例1花生AhGOS1-1基因的克隆
(1)花生种子材料的准备:收获发育中期(授粉后果针入土发育30-50天) 的花生种子,剥取种仁。
(2)总RNA的提取:采用Trizol Reagent试剂(Invitrogen公司)提取种子总RNA。
(3)cDNA文库构建:采用Clontech公司的SMART cDNA library construction kit构建文库。
(4)AhGOS1-1基因的克隆:根据含有花生AhGOS1-1基因cDNA全长的 EST序列(GenBank登录号EG029455)设计引物,并在5’端增加Sma I酶切位点和保护碱基,在3’端增加Sac I酶切位点和保护碱基,以花生种子cDNA文库为模板进行PCR扩增;
正向引物:SEQ ID NO:3:
5’-F-TCCCCCGGGCACTCCCTTCCTTCTTCATTCTCCA-3’
反向引物:SEQ ID NO:4:
5’-CGAGCTCCCATGAGACCTTATTTGGTTAGCC-3’
PCR反应体系:1μL cDNA文库模板,1μL正向引物,1μL反向引物,1μL dNTP(10mM),5μL 10×PCR Buffer,0.5μl EX-Taq酶(TaKaRa公司产品),最后补充去离子水,使总体积为50μl。PCR程序:94℃4分钟;然后进入如下循环:94℃45秒,53℃30秒,72℃45秒,共30个循环;最后72℃延伸5 分钟。扩增得到符合预期大小的目的片段,结果如图1所示。
(5)测序载体的构建:扩增完成进行琼脂糖电泳检测,回收合适大小的条带。进行回收。取5.3μL回收产物与0.7μL pMD20-T载体(TaKaRa公司产品) 连接,按产品说明书操作。
(6)大肠杆菌转化:连接产物转化大肠杆菌DH5α感受态细胞,感受态细胞购自天根生化科技(北京)有限公司,转化后的细胞在含氨苄青素(100mg/L) 的LB培养基上涂板,倒置于37℃过夜培养,挑取单菌落在含氨苄青素(100mg/L) 的LB液体培养基中培养,取少量菌液进行PCR鉴定;
(7)细菌质粒DNA的制备:选取PCR检测阳性的菌落与上述LB液体培养基中培养,收集菌体,采用天根公司的质粒小提试剂盒制备细菌质粒DNA。测序由上海英潍捷基(Invitrogen)公司完成。
实施例2植物过表达载体的构建
(1)将实施例1中测序正确的测序载体质粒与pBI121载体质粒分别用Sma I和SacI进行双酶切,酶切完成后采用天根生化科技(北京)有限公司的DNA 回收试剂盒回收酶切片段;
(2)植物表达载体构建:将上述酶切回收片段进行连接,构建植物表达载体pBI121-AhGOS1-1(图2)。取2.5μL基因片段与15μL pBI121载体片段,按T4Ligase连接酶(TaKaRa公司产品)说明书操作;
(3)大肠杆菌转化:连接产物转化大肠杆菌DH5α感受态细胞,转化后的细胞在卡那霉素(50mg/L)的LB培养基上涂板,倒置于37℃过夜培养,挑取单菌落在含卡那霉素(50mg/L)的LB液体培养基中培养,取少量菌液进行PCR 鉴定;收集阳性菌落培养的菌体,提取质粒进行测序鉴定。
实施例3拟南芥的遗传转化
一、拟南芥转化前处理
拟南芥转化前处理,开花期拟南芥,转化前一天浇足水分。主苔或侧苔长至 6-8cm,去除已有果荚和已开花蕾。
二、农杆菌菌液的准备
(1)接种带有表达质粒的农杆菌EHA105到5ml LB液体培养基(卡那霉素50mg/L、利福平30mg/L)进行菌种活化,200rpm,28℃震荡培养24-36小时;
(2)按1:100的体积比例加入菌液到新鲜LB液体培养基(卡那霉素50mg/L、利福平30mg/L)中,28℃,200rpm震荡培养16-20小时,OD600为1.0-1.8;
(3)室温3000rpm离心15分钟沉淀收集菌体;
(4)用新鲜配制的浸染培养基(5%蔗糖,0.02-0.05%表面活性剂Silwet L-77)重悬菌体至OD600在0.6-0.8。
三、拟南芥转化
(1)将重悬过后的菌液倒入干净烧杯,将待转化拟南芥的花序浸入到烧杯内60秒,吸水纸吸干残留浸染液,黑暗条件下放置16小时后,放于正常培养条件下继续培养至结实。
(2)正常管理植物,收取成熟的种子。收获后的种子在含有50mg/L卡那霉素和150mg/L羧苄青霉素的1/2MS固体培养基上筛选转基因植株。
实施例4转基因植物的qRT-PCR分析
(1)RNA的提取:RNA的提取采用北京康为世纪生物科技有限公司的超纯RNA提取试剂盒提取。
(2)第一链cDNA的合成:RNA首先用DNase I消化,去除基因组DNA 污染。根据Promega公司的MMLV逆转录酶使用方法进行cDNA第一链的合成。在冰上依次加入以下成分:2μg总RNA,1μL Olig dT(10mM),纯水补至总体积10μL,75℃水浴中变性5分钟。将上述产物置于冰上,再依次加入5μL 5×buffer,1.5μL dNTPs(10mM),1μL MMLV Reversetranscriptase(200U/μL), 1μL Ribonuclease inhibitor 0.7μL,用纯水补齐至总体积25μL,42℃反应1小时,反应完成后将cDNA存储于-20℃备用。
(3)qRT-PCR检测:
AhGOS1-1基因检测特异引物:
正向引物(SEQ ID NO:5):
5’-CACTCCCTTCCTTCTTCATTCTCCA-3’
反向引物(SEQ ID NO:6):
5’-GCCGCTCAATCCAAGACTCCAAATC-3’
内参基因Actin2引物:
正向引物(SEQ ID NO:7):5’-CACTTGCACCAAGCAGCATGAAGA-3’;
反向引物(SEQ ID NO:8):5’-AATGGAACCACCGATCCAGACACT-3’。
将cDNA原液稀释10倍后作为qRT-PCR的模板,反应体系按照BIO-RAD 公司的SsoFastTMEvaGreen Supermix说明书配制,具体如下:5μL Mix,0.5μL 正向引物,0.5μL反向引物,4μL cDNA。以拟南芥检测中稳定表达的基因Actin2 作为内参。应用荧光定量PCR仪(RoChe)LightCycler480按照以下反应程序进行qRT-PCR反应:98℃30秒;进入以下循环98℃5秒,55℃5秒,72℃ 10秒,共40个循环。应用2—ΔΔCT相对定量分析方法进行结果分析。
Q-PCR结果如图3所示,野生型植物中没有检测到AhGOS1-1的表达,而在 2个转基因拟南芥株系中均检测到较高的AhGOS1-1表达。
实施例5转基因植物的干旱耐受性检测
转基因过表达拟南芥和野生型拟南芥种子消毒后播种于1/2MS培养基平板上,4℃层积2天后转入正常萌发培养1周。幼苗移栽至土中正常培养,植物生长 1周后停止浇水,予以持续干旱处理2周,观察并记录干旱表型。将干旱后植物进行复水处理,48小时后观察并统计植物成活率。
干旱耐受性实验结果表明,随着干旱处理时间的延长,野生型植物叶片失水颜色变深并逐渐萎蔫,此时过表达植物叶片还能保持相对较绿的状态。干旱处理之后的植物复水48小时后,转基因植物成活的植株数量明显多于野生型对照,说明AhGOS1-1过表达植物的干旱耐受性强于野生型(图4)。
实施例6转基因AhGOS1-1过表达植物离体叶片失水实验
选取在土中生长2周的健壮、未抽薹的转基因过表达拟南芥和野生型拟南芥幼苗,将整个植株根以上莲座叶簇生部分剪下,放入铺有称量纸的培养皿中,培养皿置于培养箱中,70%相对湿度,22℃恒温。预先称量并记录植物起始重量,此后每隔30分钟准确称量一次重量,直到完成5小时的检测,统计并计算离体叶片失水率。
测定结果显示,AhGOS1-1过表达植物离体叶片的失水速率显著低于对照(图 5)。
实施例7ABA处理下AhGOS1-1过表达植物气孔变化检测
1、正常状态叶片气孔观察:将正常生长4周左右的野生型和转基因型拟南芥浇足水生长3天,在当天光照开始1.5h后取叶片,用透明胶带粘取叶片背面表皮,放入ZEISSImager.AI显微镜下观察气孔,用Axio软件拍照并统计气孔长宽比。
2、ABA处理叶片气孔观察
(1)取与上述正常状态观察相同大小的拟南芥,取其平展的莲座叶浸泡在气孔缓冲液(10mM KCl,50μM CaCl2,10mM MES-Tris,0.05%Triton X-100, pH 6.15)中,置于强光下照射3小时;
(2)将叶片换至添加50μM ABA的气孔缓冲液中处理3~5小时;
(3)将叶片取出用滤纸迅速吸干,用透明胶带粘取叶片下表皮,置于载玻片上,盖上盖玻片,在ZEISS Imager.AI显微镜下观察气孔,用Axio软件拍照并统计气孔长宽比。
观察结果显示,ABA处理前,转基因和野生型植物叶片的气孔在强光下完全打开,50μM ABA处理3h后,气孔关闭。但转基因植物气孔的开度明显小于野生型对照植物(图6),说明AhGOS1-1过表达植物对ABA更敏感,导致更明显的气孔关闭现象,因而在干旱状态下气孔能够迅速反应避免大量失水。
序列表
<110> 中山大学
<120> 花生AhGOS1-1基因及其在提高植物抗旱性中的应用
<130> YG18103955AA042
<141> 2018-05-23
<160> 8
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<211> 675
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atggaggtca ccgcttcgtg ggattccctc cgccaacagg caagaaagct ccaagctcag 60
ctggatgagc aaatgagttt gtaccgcaaa ttggtgtcag caaacatttc tacgaaatct 120
gatgctgctg agagtgattt ggagtcttgg attgagcggc tactcaaaca actccaacaa 180
gtaaactctc agatgaaagc ctgggtgtca tccggtgggt cagaaatggt ttctcatact 240
ctaactagac accaagaaat tcttcaagat ctcacccagg agttttatcg cctccagtct 300
agtcttagag ctaagcaaga gcatgcttca ctactagatg attttagaga gttcgatcgg 360
acaagactag acctggaaga gggtgcaggt tctgaacagc aaaccctcct aaaagagtat 420
gcatctataa gccgaagcac cggacagatg gacactgtta tttctcaagc acaagcaacg 480
atgggttcac tagtccacca gcgctcaact tttggtggaa tcaattcaaa gctcagcaat 540
atgagcagtc gcctccccac ggtaaataac atcctatcag caataaggag aaaaaaatca 600
agggatacaa taattctttc gcttgtcata tccgtgtgca cgttcttgat tttaatatac 660
tggctaacca aataa 675
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<213> 花生(Arachis hypogaea L.)
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1 5 10 15
Leu Gln Ala Gln Leu Asp Glu Gln Met Ser Leu Tyr Arg Lys Leu Val
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Ser Trp Ile Glu Arg Leu Leu Lys Gln Leu Gln Gln Val Asn Ser Gln
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Met Lys Ala Trp Val Ser Ser Gly Gly Ser Glu Met Val Ser His Thr
65 70 75 80
Leu Thr Arg His Gln Glu Ile Leu Gln Asp Leu Thr Gln Glu Phe Tyr
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Arg Leu Gln Ser Ser Leu Arg Ala Lys Gln Glu His Ala Ser Leu Leu
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Asp Asp Phe Arg Glu Phe Asp Arg Thr Arg Leu Asp Leu Glu Glu Gly
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Ala Gly Ser Glu Gln Gln Thr Leu Leu Lys Glu Tyr Ala Ser Ile Ser
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Arg Ser Thr Gly Gln Met Asp Thr Val Ile Ser Gln Ala Gln Ala Thr
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Met Gly Ser Leu Val His Gln Arg Ser Thr Phe Gly Gly Ile Asn Ser
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Claims (7)
1.花生AhGOS1-1基因,其特征在于,所述基因的核苷酸序列如SEQ ID NO:1所示。
2. 由权利要求1所述花生AhGOS1-1基因编码的AhGOS1-1,其特征在于,所述AhGOS1-1的氨基酸序列如SEQ ID NO:2所示。
3. 一种用于扩增权利要求1所述花生AhGOS1-1基因的扩增引物,其特征在于,包括上游引物F和下游引物R,其核苷酸序列依次如SEQ ID NO:3~4所示。
4.权利要求1所述花生AhGOS1-1基因或权利要求2所述花生AhGOS1-1在提高植物对干旱和/或渗透胁迫耐受性中的应用。
5.根据权利要求4所述的应用,其特征在于,所述应用为构建花生AhGOS1-1基因的过表达载体并转化到植物中。
6.根据权利要求4所述的应用,其特征在于,所述过表达载体为pBI121-AhGOS1-1。
7.根据权利要求4所述的应用,其特征在于,所述植物为拟南芥。
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