CN107188941A - 禾谷孢囊线虫的Ha‑62292蛋白、编码基因及其应用 - Google Patents

禾谷孢囊线虫的Ha‑62292蛋白、编码基因及其应用 Download PDF

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CN107188941A
CN107188941A CN201710356195.7A CN201710356195A CN107188941A CN 107188941 A CN107188941 A CN 107188941A CN 201710356195 A CN201710356195 A CN 201710356195A CN 107188941 A CN107188941 A CN 107188941A
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彭德良
罗书介
彭焕
黄文坤
孔令安
崔江宽
王高峰
刘敬
乔芬
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Abstract

本发明公开了一种禾谷孢囊线虫的Ha‑62292蛋白、编码基因及其应用。本发明提供了一种蛋白质，获取自禾谷孢囊线虫，命名为Ha‑62292蛋白，其氨基酸序列如SEQ ID NO：1所示，编码该蛋白的基因序列如SEQ ID NO：2所示。实验表明Ha‑62292基因在线虫侵染寄生过程中发挥重要作用，可作为植物抗线虫工程的靶标基因。本发明对禾谷孢囊线虫致病机理研究以及抗线虫植物开发有重大价值。

Description

禾谷孢囊线虫的Ha-62292蛋白、编码基因及其应用
技术领域
本发明属于生物技术领域，涉及一种来源于禾谷孢囊线虫的Ha-62292蛋白、编码基因及其应用。
背景技术
禾谷孢囊线虫(Heterodera avenae)是一种固着型植物内寄生线虫，主要危害小麦大麦燕麦等禾本科植物的根系，造成严重的产量损失，分布广泛。目前已经在我国的河南、安徽、山东、青海、宁夏、陕西等13个省市地区发现和报道(苏致衡,黄文坤,郑国栋,张宏嘉,刘淑艳,彭德良.北京地区小麦禾谷孢囊线虫病发生动态调查.植物保护,2013,(01):116-120.)(赵洪海,杨远永,彭德良,刘峰.小麦禾谷孢囊线虫在山东省的分布新报道和发生特点浅析.青岛农业大学学报(自然科学版),2011,(04):261-266.)(赵杰,钮绪燕,张管曲,彭德良,康振生.陕西省中南部地区小麦禾谷孢囊线虫的发生与分布.西北农业学报,2011,(06):181-185.)(黄文坤,叶文兴,王高峰,龙海波,欧师琪,彭德良.宁夏地区禾谷孢囊线虫的发生与分布.华中农业大学学报,2011,(01):74-77.)(陈新,周洪友,马玺.内蒙古中西部地区小麦禾谷孢囊线虫的发生分布.植物保护,2009,(05):114-117.)(侯生英,彭德良,王爱玲,黄文坤,耿贵工,张贵.青海省小麦孢囊线虫病调查初报.青海大学学报(自然科学版),2008,(05):84-86.)(杨传广,吴慧平,檀根甲,王月英.安徽省小麦孢囊线虫田间分布及危害调查.植物保护,2008,(02):107-110.)。该种线虫适应低温，通常在冬季之前就可开始侵染，侵染后的幼虫在寄主体内过冬，并在第二年开春后继续侵染和发育，一年只发生一代(向桂林,宋志强,梁旭东,胡小斌,亓竹冉,王暄,李红梅.禾谷孢囊线虫的田间侵染规律及垂直分布研究.麦类作物报,2013,(04):789-794.)。
由于禾谷孢囊线虫寄主范围窄、侵染周期长、研究背景不明，其致病机制研究相对较落后。随着效应蛋白作为病原物致病机制的关键因素的持续研究开发，禾谷孢囊线虫相关研究也得到进一步增长(彭德良.重要植物线虫致病相关基因研究进展.中国植物病理学会.中国植物病理学会2006年学术年会论文集.中国植物病理学会:,2006:10.)。随着基因表达和功能研究相关技术的成熟，RNA表达定量、定位分析、RNAi等均可应用到禾谷孢囊线虫中(黄文坤,彭德良,彭焕.RNA干扰技术在植物寄生线虫中的应用.农业生物技术学报,2009,(01):170-175.)。借鉴于其它植物寄生线虫的研究方法，禾谷孢囊线虫效应蛋白的研究已取得长足的进步，例如内切葡聚糖酶、细胞壁扩展蛋白、膜联蛋白等一些具有明显促进寄生作用的效应蛋白在禾谷孢囊线虫中被发现(龙海波.小麦禾谷孢囊线虫β-1，4-内切葡聚糖酶和细胞壁扩展蛋白基因的克隆及功能分析.中国农业科学院,2012.)，并且这些被发现的基因可应用于寄主介导的RNA干扰或异位表达以抑制病原线虫的侵入危害。
发明内容
本发明的目的是提供一种来源于禾谷孢囊线虫的Ha-62292蛋白及其编码基因，从表达时间上来看，主要集中在侵染前二龄、侵染后二龄第1天。而从表达部位来看，其主要集中在食道腺细胞中。实验表明该基因在禾谷孢囊线虫寄生致病过程中发挥重要作用，可作为植物抗线虫工程的靶标基因。
一种来源于禾谷孢囊线虫的Ha-62292蛋白，其氨基酸序列如SEQ ID NO：1所示。
编码禾谷孢囊线虫的Ha-62292蛋白的基因。
编码禾谷孢囊线虫的Ha-62292蛋白的基因，其核苷酸序列如SEQ ID NO：2所示。
禾谷孢囊线虫的Ha-62292蛋白在禾谷孢囊线虫防控中的应用。
所述应用为将Ha-62292蛋白作为禾谷孢囊线虫的寄主防御反应的诱导剂，用于激活寄主免疫反应增强对线虫抗性，或者获得在寄主中的Ha-62292蛋白的受体蛋白，转移到不含有该种受体蛋白的寄主中以获得抗性。
本发明的序列表Ha-62292蛋白，长度为218个氨基酸；如SEQ ID NO：1所述；编码该蛋白的基因长度为654个核苷酸，其序列如SEQ ID NO：2所示。上述蛋白来源于禾谷孢囊线虫，但来源于其它孢囊线虫甚至根结线虫的此蛋白也属于本发明保护范围。此蛋白可通过生物技术直接人工合成，先合成其编码基因，连接到表达载体后再导入生物体内表达得到。
上述编码基因的杂交探针、dsRNA、重组表达载体、干扰载体、重组病毒、转基因细胞系、转基因植物或组织、重组菌也属于本发明的保护范围。杂交探针和dsRNA列于序列表中序列3和序列4、5、6、7，其中dsRNA片段有四个，表达载体和干扰载体的构建建立在相应的启动子区基础上，具体可为单独或者多个组合使用的诱导型、增强型、组成型、组织特异型，并且伴随有合适的筛选标记，例如抗性、化学发光或染色、荧光。转化目标生物可以是细菌、真菌、植物或动物细胞系。
Ha-62292蛋白在禾谷孢囊线虫病害防控中的应用。
蛋白质的功能应用研究常用RNA干扰、体内异位表达等基于编码基因操作的方法，本发明使用RNA干扰有效影响了基因的表达水平，并且在线虫的寄生致病能力方面表现出差异。此差异建立在对禾谷孢囊线虫敏感的寄主上，例如大麦Golden Promise。
Ha-62292基因的应用主要包括
(d1)在线虫不同发育阶段表达水平变化
(d2)在线虫不同组织部位表达水平
孢囊线虫不同阶段其表达的蛋白质有差异，并且反映在编码基因的转录水平上。在涉及到对寄主的寄生致病和抗逆方面，往往在寄生过程的前期表达，而与自生发育相关的蛋白质，其编码基因很可能在后期表达。此外编码基因在线虫的转录表达部位也与其应用有关，例如食道腺细胞中表达的蛋白很可能与寄生植物有关，而在头感器或者表皮中表达则可能与寻找寄主或者抑制寄主免疫反应有关。本发明提供的Ha-62292基因，从表达时间上来看，主要集中在侵染前二龄、侵染后二龄第1天。而从表达部位来看，其主要集中在食道腺细胞中。
本发明涉及到的寄主品种包括但不限于大麦Golden Promise，也可为其它感病麦类作物，例如小麦和燕麦。本发明通过基因沉默，将针对Ha-62292基因设计的dsRNA片段(如SEQ ID NO：4、5、6)分别导入禾谷孢囊线虫体内，结果显示，4个dsRNA片段RNA干扰后线虫侵入根内的数量相比对照明显上升，推测此基因可能与激活寄主免疫反应有关。
根据基因沉默结果推测，该基因编码的蛋白可能在被寄主识别后激活相应的免疫反应，而该基因也可能属于无毒基因。无毒基因表达产物在寄主中往往有固定的受体，该受体行使抗病作用，可通过酵母双杂交、免疫共沉淀等蛋白互作研究技术挖掘受体蛋白和其编码基因，再将编码基因转化至不含该种受体的感病寄主中获得对相应线虫的抗性。本发明对于孢囊线虫致病机理研究以及抗线虫植物制备具有重大价值。
附图说明
图1为发育表达定量分析，结果显示，Ha-62292基因表达主要集中在侵染前二龄和侵染后二龄第1天，数据收集自三次独立重复。
图2为侵染前二龄幼虫组织表达定位，结果显示，Ha-62292基因表达主要集中在食道腺细胞中。
图3为RNA干扰显示Ha-62292基因表达被干扰后二龄幼虫侵入根内的数量统计图，结果表明，被干扰后二龄幼虫侵入根内的数量明显上升。
具体实施方式
以下实施例中所用到的试剂如无特别说明均为国产分析纯，所用方法如无特别说明均按照试剂盒使用说明书进行，涉及到RNA的实验所用试剂耗材均经过DEPC处理。
实施例1：Ha-62292蛋白及其编码基因的分离
使用TRIzol(Ambion)试剂盒提取禾谷孢囊线虫总RNA，DNase Treatment andRemoval Reagents(Ambion)处理后使用III First-Strand SynthesisSystem for RT-PCR反转录试剂盒(invitrogen)合成cDNA。并以此cDNA为模板，PCR扩增基因序列，所用试剂为Max DNA Polymerase高保真酶(Takara)，引物序列为：5'-GATTTGTTGTCCGCAGTGCCTTTG-3'，5'-GCCAAAGGCACTGCGGACAACAAA-3'。将扩增得到的片段纯化后连接测序。测序结果显示其包含有序列表中SEQ ID NO：2所示的开放阅读框，编码SEQID NO：1的蛋白质序列，并分别将该蛋白质序列和基因序列命名为Ha-62292蛋白和Ha-62292基因。
实施例2：Ha-62292基因的发育表达变化
酶解法收集不同发育阶段的禾谷孢囊线虫，使用TRIzol(Ambion)试剂盒提取总RNA，DNase Treatment and Removal Reagents(Ambion)处理后使用IIIFirst-Strand Synthesis System for RT-PCR反转录试剂盒(invitrogen)合成cDNA。并以此cDNA为模板，real-time PCR检测不同阶段表达水平变化，所用PCR试剂为SelectMaster Mix(Applied Biosystems)，引物为5'-CTACAAAGCGGCGGGAAAT-3'，5'-ATGCACCAGAATAATCTGAACAGC-3'，内参基因为GAPDH，内参基因引物为5'-AGCGGCACAGAACATCATCC-3'，5'-GGTCCTCCGTGTAGCCCAAA-3'。结果表明此基因表达水平主要集中在侵染前二龄、侵染后二龄第1天。
实施例3：Ha-62292基因的组织表达定位
以禾谷孢囊线虫线虫cDNA为模板，Max DNA Polymerase高保真酶(Takara)和引物5'-CTTCTTGCTCATTGCCCTAA-3'、5'-AAATTCTCGCCCATCCCT-3'PCR扩增，回收纯化得到的扩增片段，并以此为模板，PCR DIG Probe Synthesis Kit(Roche)和反义引物作单引物扩增，以扩增产物为杂交探针。杂交方法参考文献de J.M.Boer YY,Geert Smant,Eric L.Davis,Thomas J.Baum(1998)In-situ Hybridization to Messenger RNA inHeterodera glycines.Journal of Nematology 30:309-312.杂交结果显示，此基因表达定位在禾谷孢囊线虫的食道腺细胞中。
实施例4：Ha-62292基因RNA干扰
以禾谷孢囊线虫cDNA为模板，Max DNA Polymerase高保真酶(Takara)和下表引物扩增，将扩增产物纯化并以此为模板，HiScribe T7 Quick HighYield RNA Synthesis Kit(NEB)试剂盒做体外转录，MEGAclearTM Kit Purification forLarge Scale Transcription Reactions(Ambion)纯化转录得到的dsRNA后-80℃保存备用。
按照下表配制浸泡体系，无dsRNA体系为阴性对照
浸泡条件为黑暗16℃缓慢摇动36h。浸泡后接种大麦，每个50ml离心管中3棵大麦，接种线虫400，实验重复至少3次，接种后的大麦培养在16度光照培养箱中。接种后第8天酸性品红染色统计根中线虫数量。结果显示，4个dsRNA片段RNA干扰后线虫侵入根内的数量相比对照明显上升，推测此基因可能与激活寄主免疫反应有关。
SEQUENCE LISTING
<110> 中国农业科学院植物保护研究所
<120> 禾谷孢囊线虫的Ha-62292蛋白、编码基因及其应用
<130> PP17066-ZWB
<160> 7
<170> PatentIn version 3.3
<210> 1
<211> 218
<212> PRT
<213> Ha-62292蛋白序列
<400> 1
Met Glu Leu His Ser Lys Val Val Cys Phe Phe Leu Leu Ile Ala Leu
1 5 10 15
Ile Ala Ile Pro Tyr Asn Val Trp Ala Leu Ser Ala Gly Gly Arg Val
20 25 30
Ser Val Leu Asn Cys His Asn Asn Tyr Arg Ser Gln Leu Ala Lys Gly
35 40 45
Thr Ala Asp Asn Lys Ser Gly Lys Met Pro Ala Gly Ser Asn Leu Ile
50 55 60
Glu Leu Lys Tyr Leu Asn Glu His Glu Lys Gly Ala Gln Ser Trp Ala
65 70 75 80
Asp Gly Cys Ser Met Ser His Ser Ser Ser Ser Gln Arg Gln Gly Met
85 90 95
Gly Glu Asn Leu Tyr Met Ser Ser Ser Ser Thr Ile Ser Glu Ala Glu
100 105 110
Ala Leu Lys Gln Ala Cys Asp Met Trp Trp Ala Glu Leu Lys Glu Phe
115 120 125
Gly Phe Asp Gln Ser Leu Val Leu Asn Met Asn Glu Phe Asn Lys Gly
130 135 140
Ile Gly His Trp Ser Gln Gln Ala Trp Ala Lys Thr Ala Gln Ile Gly
145 150 155 160
Cys Ala Leu Ala Arg Cys Pro Ser Ser Gln Trp Gln Thr Trp Val Val
165 170 175
Cys Arg Tyr Lys Ala Ala Gly Asn Met Leu Asn Glu Met Val Tyr Lys
180 185 190
Lys Gly Thr Ala Cys Ser Gly Cys Ser Asp Tyr Ser Gly Ala Ser Cys
195 200 205
Asn Asn Ala Asn Gly Leu Cys Val Val Pro
210 215
<210> 2
<211> 654
<212> DNA
<213> Ha-62292基因序列
<400> 2
atggaactcc actcaaaagt tgtctgcttc ttcttgctca ttgccctaat tgctattcct 60
tacaatgttt gggcactttc tgctggcggt cgcgtgtcgg tgcttaattg tcacaacaat 120
tatcgttccc aattggccaa aggcactgcg gacaacaaat ccggcaaaat gccggctggt 180
agcaatttga tagagctaaa atatttgaat gaacatgaga agggtgcaca gagctgggcc 240
gacggttgct caatgtccca ctcgagttca tcacaacgcc aagggatggg cgagaatttg 300
tacatgtcaa gttcatctac catttcggaa gctgaggcgc tcaaacaagc ttgtgacatg 360
tggtgggcag agctcaaaga gttcggcttt gaccaaagcc tggtgctgaa catgaacgag 420
ttcaacaagg gcatcggaca ttggagccag caagcgtggg ccaaaactgc tcaaattggt 480
tgtgccttgg cacgatgtcc cagctcccaa tggcaaacat gggttgtgtg ccgctacaaa 540
gcggcgggaa atatgttgaa cgaaatggtc tacaaaaagg gaactgcatg cagtggctgt 600
tcagattatt ctggtgcatc gtgcaataat gccaatggtc tgtgcgttgt gcca 654
<210> 3
<211> 270
<212> DNA
<213> Ha-62292基因杂交探针序列
<400> 3
cttcttgctc attgccctaa ttgctattcc ttacaatgtt tgggcacttt ctgctggcgg 60
tcgcgtgtcg gtgcttaatt gtcacaacaa ttatcgttcc caattggcca aaggcactgc 120
ggacaacaaa tccggcaaaa tgccggctgg tagcaatttg atagagctaa aatatttgaa 180
tgaacatgag aagggtgcac agagctgggc cgacggttgc tcaatgtccc actcgagttc 240
atcacaacgc caagggatgg gcgagaattt 270
<210> 4
<211> 267
<212> DNA
<213> Ha-62292基因RNA干扰dsRNA片段序列1
<400> 4
gctcattgcc ctaattgcta ttccttacaa tgtttgggca ctttctgctg gcggtcgcgt 60
gtcggtgctt aattgtcaca acaattatcg ttcccaattg gccaaaggca ctgcggacaa 120
caaatccggc aaaatgccgg ctggtagcaa tttgatagag ctaaaatatt tgaatgaaca 180
tgagaagggt gcacagagct gggccgacgg ttgctcaatg tcccactcga gttcatcaca 240
acgccaaggg atgggcgaga atttgta 267
<210> 5
<211> 181
<212> DNA
<213> Ha-62292基因RNA干扰dsRNA片段序列2
<400> 5
gctttgacca aagcctggtg ctgaacatga acgagttcaa caagggcatc ggacattgga 60
gccagcaagc gtgggccaaa actgctcaaa ttggttgtgc cttggcacga tgtcccagct 120
cccaatggca aacatgggtt gtgtgccgct acaaagcggc gggaaatatg ttgaacgaaa 180
t 181
<210> 6
<211> 255
<212> DNA
<213> Ha-62292基因RNA干扰dsRNA片段序列3
<400> 6
gccctaattg ctattcctta caatgtttgg gcactttctg ctggcggtcg cgtgtcggtg 60
cttaattgtc acaacaatta tcgttcccaa ttggccaaag gcactgcgga caacaaatcc 120
ggcaaaatgc cggctggtag caatttgata gagctaaaat atttgaatga acatgagaag 180
ggtgcacaga gctgggccga cggttgctca atgtcccact cgagttcatc acaacgccaa 240
gggatgggcg agaat 255
<210> 7
<211> 211
<212> DNA
<213> Ha-62292基因RNA干扰dsRNA片段序列4
<400> 7
gcctggtgct gaacatgaac gagttcaaca agggcatcgg acattggagc cagcaagcgt 60
gggccaaaac tgctcaaatt ggttgtgcct tggcacgatg tcccagctcc caatggcaaa 120
catgggttgt gtgccgctac aaagcggcgg gaaatatgtt gaacgaaatg gtctacaaaa 180
agggaactgc atgcagtggc tgttcagatt a 211

Claims (5)

1.一种来源于禾谷孢囊线虫的Ha-62292蛋白，其氨基酸序列如SEQ ID NO：1所示。
2.编码禾谷孢囊线虫的Ha-62292蛋白的基因。
3.根据权利要求1所述的编码禾谷孢囊线虫的Ha-62292蛋白的基因，其核苷酸序列如SEQ ID NO：2所示。
4.权利要求1所述的禾谷孢囊线虫的Ha-62292蛋白在禾谷孢囊线虫防控中的应用。
5.根据权利要求4所述应用，为将Ha-62292蛋白作为禾谷孢囊线虫的寄主防御反应的诱导剂，用于激活寄主免疫反应增强对线虫抗性，或者获得在寄主中的Ha-62292蛋白的受体蛋白，转移到不含有该种受体蛋白的寄主中以获得抗性。
CN201710356195.7A 2017-05-16 2017-05-16 禾谷孢囊线虫的Ha‑62292蛋白、编码基因及其应用 Pending CN107188941A (zh)

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