CN109666675B - 褐飞虱NlAtg3基因、编码蛋白及其应用 - Google Patents
褐飞虱NlAtg3基因、编码蛋白及其应用 Download PDFInfo
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- CN109666675B CN109666675B CN201811388131.6A CN201811388131A CN109666675B CN 109666675 B CN109666675 B CN 109666675B CN 201811388131 A CN201811388131 A CN 201811388131A CN 109666675 B CN109666675 B CN 109666675B
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Abstract
本发明公开了一种褐飞虱NlAtg3基因、编码蛋白及其应用。NlAtg3基因，其核苷酸序列如SEQ ID NO:1所示，该基因是褐飞虱正常生存所必需，其功能受到抑制会导致褐飞虱存活率下降。NlAtg3基因编码的蛋白，氨基酸序列如SEQ ID NO:2所示。所述的NlAtg3基因或蛋白的应用，用于农药研发和生物防治褐飞虱。本发明对该基因成功实施了RNA干扰，结果表明害虫的存活率下降，有望在实现抑制害虫的同时，充分发挥生态防治的功能。
Description
技术领域
本发明涉及一种褐飞虱NlAtg3基因、编码蛋白及其应用。
背景技术
褐飞虱（Nilaparvata lugens Stål）是一种单食性水稻害虫，主要通过取食水稻韧皮部汁液为害水稻。目前，对褐飞虱的应急防治主要以化学防治为主，但化学防治容易导致害虫的抗药性上升、水稻上有害物质残留、害虫再猖獗及环境污染等问题。在防治褐飞虱的农药中，曾经发挥过较好效果的有效农药，如氟虫腈、吡虫啉、噻嗪酮等，均已淘汰出局或限制使用，究其原因，此类化学农药多以大规模高强度杀灭害虫为目标，但是，由于褐飞虱种群遗传多样性非常复杂等现实原因，其中往往有部分个体能够存活下来，高选择压力的作用结果，最终会导致适应性更强的种群形成。另一方面，化学农药在杀死害虫的同时，也会危及包括天敌在内的非靶标生物，不可避免的会给农田生态***造成负面影响，导致生态防治功能得不到应有的发挥。因此，筛选与发现新的褐飞虱防治靶标，调整目前的褐飞虱防治策略具有重要的现实意义。
CN107988244A公开了一种褐飞虱生存相关的ATPSb基因、编码蛋白及其应用，该基因在维持褐飞虱正常的生存中发挥重要功能，其功能受到抑制会导致褐飞虱存活率下降。
发明内容
为了克服现有技术的不足，本发明提供了一种褐飞虱NlAtg3基因、编码蛋白及其应用。本发明根据NlAtg3基因编码的蛋白质相对保守但核酸序列与其他生物同源性较低的特点，对靶标基因进行RNA干扰，实现了在核酸水平对褐飞虱的抑制，显著地降低了褐飞虱的存活率，并避免了在蛋白质水平采用的喷施农药等可能给非靶标生物造成的杀伤。
本发明的技术方案如下：
一种褐飞虱NlAtg3基因，其核苷酸序列如SEQ ID NO:1 所示，该基因编码ATG3蛋白，是褐飞虱正常生存所必需，其功能受到抑制会导致褐飞虱存活率下降。
一种所述的褐飞虱NlAtg3基因编码的蛋白，氨基酸序列如SEQ ID NO:2所示，该基因功能受到抑制会导致褐飞虱存活率下降。
一种所述的褐飞虱NlAtg3基因的应用，用于农药研发和生物防治褐飞虱。
一种针对所述的褐飞虱NlAtg3基因的RNA干扰技术在控制褐飞虱方面的应用，所述的RNA干扰技术导致褐飞虱的存活率下降。
一种如所述的褐飞虱NlAtg3基因编码的蛋白的应用，用于农药研发和生物防治褐飞虱。
本发明的有益效果：（1）褐飞虱存活率下降，可以减轻害虫取食对水稻作物的直接危害。（2）本发明利用靶标基因的核苷酸序列与天敌核苷酸序列同源性较低的特点，可以在核酸水平上进行RNA干扰，避免了由于蛋白质结构保守对天敌等非靶标生物的伤害，有望在实现抑制害虫的同时，充分发挥生态防治的功能。
附图说明
图1是褐飞虱NlAtg3基因的mRNA表达水平。其中，1-2N: 1-2龄若虫；3-4N: 3-4龄若虫；5N: 5龄若虫；E1-9: 羽化1-9天的雌性褐飞虱。
图2 是RNA干扰对褐飞虱NlAtg3基因表达量的影响；
图中数据为3次重复的平均值±标准偏差，星号表示在统计分析上处理组与对照组之间存在极显著差异（T检验，P<0.01）。dsGFP：对照组；dsNlAtg3：dsNlAtg3饲喂的RNA干扰组。
图3 是NlAtg3基因的RNA干扰对褐飞虱存活率的影响；
图中数据为3次重复的平均值±标准偏差，星号表示在统计分析上处理组与对照组之间存在显著差异（T检验，P<0.05），双星号表示在统计分析上处理组与对照组之间存在显著差异（T检验，P<0.01）。
具体实施方式
ATPSb基因是线粒体中ATP合酶的一个亚基，ATP合酶负责线粒体中ATP的合成过程，这个基因功能被干扰会影响线粒体中ATP的合成（CN107988244A）。
NlAtg3基因是褐飞虱中编码ATG3蛋白的基因，ATG3蛋白是细胞自噬(Autophagy)发生过程中的一种类泛素化E2蛋白，它催化ATG8蛋白磷脂酰乙醇胺化(Phosphatidylethanolamine, PE)。细胞自噬是普遍存在于生物体的一种细胞自我吞噬行为，是真核生物中进化保守的对细胞内物质进行周转的重要过程。该过程中一些损坏的蛋白或细胞器被双层膜结构的自噬小泡(自噬小体)包裹后，送入溶酶体中进行降解并得以循环利用。细胞自噬小体的形成有许多细胞自噬蛋白(Autophagy-related gene, ATG)参与并最终与溶酶体融合完成目标蛋白的降解。昆虫发育过程中细胞自噬发挥了重要作用，NlAtg3基因受到干扰，会影响昆虫细胞的自噬过程，进而影响昆虫的正常生长发育。
所以，ATPSb基因参与的是细胞线粒体中ATP的合成，NlAtg3基因参与的是细胞的自噬过程，两个基因分属不同的细胞生理过程，作为褐飞虱防治的不同靶标，二者有本质的不同。
以下结合附图和具体实施方式对本发明做进一步的说明。
实施例1
1 材料与方法
1.1 供试褐飞虱
供试褐飞虱种群为感虫水稻品种TN1上饲养的Tn种群，由本实验室连续在TN1上饲养60代以上，饲养温度为26±2℃，相对湿度为80%±5%，光周期为12L:12D。
主要试剂
TaKaRa MiniBEST Universal RNA Extraction Kit，TaKaRa MiniBEST AgaroseGel DNA Extraction Kit，PrimeScript RT reagent Kit With gDNA Eraser，DNA 2000Marker，Premix Taq™（TaKaRa Taq™ Version 2.0 plus dye），SYBR Premix Ex Taq 均购自大连 TaKaRa公司，SMARTer® RACE 5’/3’ Kit User Manual购自美国Clontech公司，MEGAscript® T7 High Yield Transcription Kit购自美国Ambion公司，测序以及引物合成有由上海桑尼生物科技有限公司完成。
褐飞虱NlAtg3基因全长cDNA的克隆
收集TN1水稻上不同龄期的褐飞虱若虫和成虫混合材料，立即放入液氮中冻存。取样时若虫取30-50只，成虫取5只。使用TaKaRa MiniBEST Universal RNA Extraction Kit提取总RNA，具体步骤参照其说明书进行。使用琼脂糖凝胶电泳和Nanodrop 2000（Thermo）对RNA进行完整性及纯度检测。以1μg总RNA为模板，使用PrimeScript RT reagent KitWith gDNA Eraser试剂盒反转录合成cDNA，贮存于-20℃备用。
根据本实验室的转录组测序序列信息，获得褐飞虱NlAtg3基因的部分核心序列，并经NCBI网站序列比对鉴定。应用Primer Premier 5.0软件设计引物NlAtg3-F和NlAtg3-R（表1），对核心序列进行验证。以TN1上不同龄期的混合褐飞虱cDNA为模板，验证其核心序列。NlAtg3全长cDNA的克隆参照Hao et al（2015）的方法，PCR扩增反应体系为50 μL，其中包括PCR Mix 25 μL，10 μmol/L正反引物各2 μL，cDNA模板2 μL，ddH2O 19 μL。PCR反应程序为：94℃ 4 min；94℃ 30 s，55℃ 30 s，72℃ 3 min, 30个循环；72℃ 10 min；4℃保存。PCR扩增产物经1%琼脂糖凝胶电泳检测，用TaKaRa MiniBEST Agarose Gel DNAExtraction Kit回收目的片段，连接到载体pMD-18T上4℃过夜，转化到JM109感受态细胞中加1 mL LB液体培养基37℃摇床培养2 h，取200 μL菌液涂布在含有1% Amp的LB固体培养基37℃倒置培养9 h，随机挑取5个单菌落于含1% Amp的 LB 液体培养基1 mL的1.5 mL离心管中37℃摇床培养12 h，取1 μL菌液进行菌液PCR鉴定阳性克隆菌株并送到上海桑尼生物科技有限公司测序。测序结果用DNAMAN软件和原序列比对验证。
选取符合要求的RNA样品，用SMARTer® RACE 5’/3’ Kit User Manual 合成5'-RACE和3'-RACE的模板。分别设计外引物NlAtg3-5O和NlAtg3-3O（表1），以及内引物NlAtg3-5I和NlAtg3-3I（表1）。根据RACE试剂盒说明书，采用巢式PCR对目的基因的2端进行扩增，电泳，胶回收，连接，转化，测序，测序结果应用DNAMAN软件进行比对拼接获得NlAtg3的全长cDNA。根据获得的全长cDNA序列，设计全长验证引物NlAtg3-FL-F和NlAtg3-FL-R（表1）对拼接的全长序列进行验证。
表1 基因克隆、荧光定量PCR及合成dsRNA的引物
1.4 褐飞虱NlAtg3基因的序列分析
根据DNAMAN 拼接获得的NlAtg3全长cDNA 序列信息，利用开放阅读框分析软件(ORF finder https://www.ncbi.nlm.nih.gov/orffinder/) 预测开放阅读框及蛋白质翻译情况，使用NCBI Blastx 进行氨基酸序列同源性比对，使用在线工具ExPASy ( http://web.expasy.org/protparam/) 对蛋白质的分子量、理论等电点等进行预测，采用SignalP4.1 server (http://www.cbs.dtu.dk/services/SignalP/) 对信号肽进行预测，采用在线工具InterProScan (http://www.ebi.ac.uk/interpro/search/sequence-search) 对蛋白质功能域进行预测。
褐飞虱NlAtg3基因的表达规律分析
利用荧光定量PCR技术检测NlAtg3基因在TN1水稻上不同龄期的褐飞虱种群的相对表达量，包括1-2龄、3-4龄、5龄若虫和羽化1、3、5、7、9天的雌成虫以及雄虫。荧光定量PCR特异性引物为QNlAtg3-F和QNlAtg3-R（表1），以RPS11基因为内参（Yuan et al, 2014），检测褐飞虱NlAtg3基因的相对表达量。荧光定量PCR参考马艳等（2013）的反应体系及方法，其中退火温度改为53℃。
饲喂法进行RNAi
根据cDNA全长序列设计用于合成dsRNA干扰片段的引物dsNlAtg3-F和dsNlAtg3-R（表1），并在特异性引物5' 端加保护碱基( GGATCC ) 及T7 启动子( TAATACGACTCACTATA )。为了不影响RNAi后的转录水平检测，干扰片段不包括荧光定量PCR检测的片段。按照MEGAscript® T7 High Yield Transcription Kit（Ambion）试剂盒说明书合成dsRNA。合成后的dsRNA采用LiCl沉淀法进行纯化：向dsRNA反应体系中加入30ul的ddH2O和30ul的LiCl Precipitation Solution，-20℃放置1h左右，11000rpm，4℃离心15min，去上清后加入用DEPC水配制的70%的乙醇1ml进行洗涤，离心后去除乙醇，加入20ulddH2O溶解，放置-20℃等待使用。
取饲养在TN1水稻品种上2龄褐飞虱若虫进行人工饲养，饲养条件参照Fu 等（2001）的营养液配方，饲喂装置为两端覆盖Parafilm膜（内含取食液）的双通玻璃管（2.5cm × 15 cm），待试虫饲喂取食液5天适应后，进行RNA干扰处理，纯化的dsRNA用取食液配制到终浓度为0.5 μg/μL。处理时，以取食液加入相应体积的dsRNA稀释液为对照组（CK），每饲喂装置接入20只经过上述预处理且发育基本一致的褐飞虱若虫，设置3个重复，每天更换取食液，清理和统计死亡的褐飞虱，计算存活率。
此外，设置平行的RNA干扰处理，每饲喂装置放入25只发育一致的褐飞虱若虫，且两管作为一个重复，每2天对各组分别进行取样，每组取6只若虫，并进行RNA的抽提及后续NlAtg3基因表达量的荧光定量PCR检测。
数据统计与分析
采用WPS Excel软件进行数据整理，运用SPSS16.0独立样本T检验进行显著性差异分析。
结果与分析
2.1 褐飞虱NlAtg3基因的cDNA全长克隆及序列分析
用生物学软件分析获得的褐飞虱自噬相关基因NlAtg3基因的cDNA全长序列。使用ORF Finder分析发现，NlAtg3基因的ORF共990 bp（SEQ ID NO：1），编码329个氨基酸（SEQID NO：2）。ExPASy软件分析发现NlAtg3基因编码的蛋白分子量为37.78 kDa，等电点（pI）为4.46。蛋白结构域预测发现NlAtg3基因有一个典型的N-末端(位于8-158位氨基酸)和C-末端结构(位于300-323位氨基酸)。
褐飞虱NlAtg3基因的表达规律分析
利用荧光定量PCR检测NlAtg3基因在褐飞虱不同发育阶段的表达规律，结果发现，NlAtg3基因在褐飞虱不同发育阶段均有表达，1-2龄若虫相对较高，3-4龄居中，5龄最低（图1，左）；雌虫各发育阶段变化不明显（图1，右）。
干扰对褐飞虱NlAtg3基因表达水平的影响
荧光定量PCR检测注射5 μg/μl的dsRNA 4d后的褐飞虱，结果显示，NlAtg3基因注射dsRNA后的表达水平均出现大幅下降，与dsGFP对照组相比，达到了极差异显著（P<0.01）（图2）。
2.4 NlAtg3基因的RNA干扰对褐飞虱存活率的影响
RNA干扰结果发现，注射ds NlAtg3的致死作用显著，在RNA干扰的第 4天，注射ds NlAtg3的处理组的存活率与对照组达到了显著差异（P<0.05），从第5天开始就与对照组达到了极显著差异（P<0.01），到第7天时存活率仅有6.67%（图3）。
序列表
<110> 中国计量大学
<120> 褐飞虱NlAtg3基因、编码蛋白及其应用
<141> 2018-11-21
<160> 2
<170> SIPOSequenceListing 1.0
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<211> 1002
<212> DNA
<213> 褐飞虱(Nilaparvata lugens Stal)
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cctgttttaa aggaatcgaa atttcgtgag acgggagtga tcactcctga agagtttgtg 120
gcggccggcg atcacctggt gcatcactgc cccacctggc agtgggcgtc cggggacgaa 180
gccagggcca agacctactt gcccaagacc aaacagtttc tcatcaccaa gaatgtgccc 240
tgctcacgaa ggtgcaaaga gattgagtat tgtgacgagc aagagaaaat tctagagcct 300
gatgatcctg acggtggctg ggtggacact caccgcgtgg accccactag tggcctcgac 360
gaaaatgtct ccggaatgac attagactca aacacaactg ttcagagggt caatgattca 420
cctccatctg gagactttca gacgaacacc acaaacgcca ataataatgt cgatgatgat 480
gatgatgacg atgatgataa cgaagaagca gccgacatgg aaatgtttga ggaaagtggt 540
cttctagacg aggaggatga ggcgactgct gaggatgtca aaatcgaaaa agatgaaaga 600
aatgctgcag gtgacggcga gatagtcaaa accagaacgt atgacttaca tatcacatac 660
gacaaatatt accagactcc gcgattatgg ctttttggtt atgatgagaa ccataagcca 720
ctgaatgttg aacagatgta tgaagatgtg aatcaagatt atgcgaagaa aactgtgaca 780
atggaaacac atccgcacgt tccagggcct cccatggctt ctgtccatcc ttgcaggcac 840
gctgaagtga tgcagaagat aattcagacc gtattggaag gaggaggcga gctaggtgtt 900
catatgtatt tgataatttt tctgaaattt gttcaatcag tcattccaac tatagagtat 960
gattacacac aaaattttac aatgtggtct aaaacgctgt ga 1002
<210> 2
<211> 333
<212> PRT
<213> 褐飞虱(Nilaparvata lugens Stal)
<400> 2
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His Cys Pro Thr Trp Gln Trp Ala Ser Gly Asp Glu Ala Arg Ala Lys
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Val Lys Ile Glu Lys Asp Glu Arg Asn Ala Ala Gly Asp Gly Glu Ile
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Leu Asn Val Glu Gln Met Tyr Glu Asp Val Asn Gln Asp Tyr Ala Lys
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Lys Thr Val Thr Met Glu Thr His Pro His Val Pro Gly Pro Pro Met
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Asp Tyr Thr Gln Asn Phe Thr Met Trp Ser Lys Thr Leu
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Claims (2)
1.一种褐飞虱NlAtg3基因的应用，其特征在于，利用RNA干扰技术抑制褐飞虱NlAtg3基因的功能从而降低褐飞虱的存活率，所述的褐飞虱NlAtg3基因的核苷酸序列如SEQ ID NO:1 所示。
2.如权利要求1所述的应用，其特征在于，用于生物防治褐飞虱。
Priority Applications (1)
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