CN115160419B - 盔形毕赤酵母Thioredoxin类分泌蛋白及其应用 - Google Patents
盔形毕赤酵母Thioredoxin类分泌蛋白及其应用 Download PDFInfo
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Abstract
本发明公开了氨基酸序列如SEQ ID No.1、SEQ ID No.2、SEQ ID No.3或SEQ ID No.4所示的盔形毕赤酵母Thioredoxin类分泌蛋白及其在制备诱导柑橘果实抗病性的保鲜剂中的应用,研究结果证实,农杆菌介导的遗传转化在柑橘果实上瞬时表达Thioredoxin类分泌蛋白,能一定程度抑制柑橘果实绿霉病的发病和病斑直径的增长;本发明从盔形毕赤酵母中挖掘出具有诱导柑橘果实抗病性的Thioredoxin类分泌蛋白,不仅丰富了盔形毕赤酵母诱导柑橘果实抗病性的机理,拓宽了拮抗酵母在果实保鲜领域的应用方式,也为柑橘果实采后绿霉病的防治提供了新的解决方法。
Description
技术领域
本发明属于基因工程技术和果实采后病害生物防治技术领域,涉及盔形毕赤酵母(Pichia galeiformis)Thioredoxin类分泌蛋白及其应用。
背景技术
拮抗酵母是果实采后病害防治中常用的生物防治拮抗菌。已有大量文献报道拮抗酵母能诱导果实抗病性,涉及果实多种抗病基因的表达、激素合成和信号转导途径、抗病相关物质的合成等。然而,拮抗酵母分泌蛋白对果实抗病性的作用还不清楚。
柑橘是芸香科柑橘亚科柑橘属植物,是世界和中国产量最多的水果。新鲜柑橘果实在采后运输、贮藏等过程中极易受微生物侵染,损失可达总产量的30%以上。由真菌指状青霉(Penicillium digitatum)引起的绿霉病是柑橘果实贮藏过程中主要的侵染性病害。目前产业上主要采用咪鲜胺等化学杀菌剂控制果实采后病害,但存在杀菌剂残留、微生物产生耐药性等问题。
发明内容
本发明的目的在于探究和明确拮抗酵母分泌蛋白在诱导果实抗病性中的作用,以期为果实采后病害的防治提供新的解决方案。
经研究,本发明提供如下技术方案:
1.盔形毕赤酵母Thioredoxin类分泌蛋白,氨基酸序列如SEQ ID No.1、SEQ IDNo.2、SEQ ID No.3或SEQ ID No.4所示。
2.盔形毕赤酵母Thioredoxin类分泌蛋白的编码基因。
进一步,所述编码基因的核苷酸序列如SEQ ID No.5、SEQ ID No.6、SEQ ID No.7或SEQ ID No.8所示。
3.含有盔形毕赤酵母Thioredoxin类分泌蛋白编码基因的重组表达载体。
进一步,所述重组表达载体是将核苷酸序列如SEQ ID No.5、SEQ ID No.6、SEQ IDNo.7或SEQ ID No.8所示的盔形毕赤酵母Thioredoxin类分泌蛋白编码基因克隆入植物表达载体pCAMBIA2300的多克隆位点KpnI和PstI之间而得到。
4.含有上述重组表达载体的工程菌。
进一步,所述工程菌是将含有盔形毕赤酵母Thioredoxin类分泌蛋白编码基因的重组表达载体转入农杆菌GV3101中而得到。
5.盔形毕赤酵母Thioredoxin类分泌蛋白在制备诱导柑橘果实抗病性的保鲜剂中的应用。
进一步,所述诱导柑橘果实抗病性为诱导柑橘果实绿霉病抗病性。
6.所述工程菌在制备诱导柑橘果实抗病性的保鲜剂中的应用。
进一步,所述诱导柑橘果实抗病性为诱导柑橘果实绿霉病抗病性。
本发明的有益效果在于:本发明提供了盔形毕赤酵母Thioredoxin类分泌蛋白及其在制备诱导柑橘果实抗病性的保鲜剂中的应用,研究结果证实,农杆菌介导的遗传转化在柑橘果实上瞬时表达Thioredoxin类分泌蛋白,能一定程度抑制柑橘果实绿霉病的发病和病斑直径的增长。本发明从盔形毕赤酵母中挖掘出具有诱导柑橘果实抗病性的Thioredoxin类分泌蛋白,不仅丰富了盔形毕赤酵母诱导柑橘果实抗病性的机理,拓宽了拮抗酵母在果实保鲜领域的应用方式,也为柑橘果实采后绿霉病的防治提供了新的解决方法。
附图说明
图1为盔形毕赤酵母Thioredoxin类分泌蛋白PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、PgThioredoxin4编码基因的电泳图,其中M为DNA分子量标准。
图2为重组载体pCAMBIA2300-PgThioredoxin1、pCAMBIA2300-PgThioredoxin2、pCAMBIA2300-PgThioredoxin3、pCAMBIA2300-PgThioredoxin4的电泳图,其中M为DNA分子量标准。
图3为含有PgThioredoxin1、PgThioredoxin2、PgThioredoxin3或PgThioredoxin4编码基因的农杆菌工程菌的电泳图,其中M为DNA分子量标准。
图4为盔形毕赤酵母Thioredoxin类分泌蛋白PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、PgThioredoxin4对柑橘果实采后绿霉病的诱导效果;A为发病率,B为病斑直径,*表示与对照组(Control)相比差异显著(P<0.05);C为柑橘果实在接种指状青霉后于25℃贮藏6天后的绿霉病发病症状。
具体实施方式
为了使本发明的目的、技术方案和有益效果更加清楚,下面对本发明的优选实施例进行详细的描述。
本实施例所用盔形毕赤酵母分离于柑橘果园的柠檬果实表面。
1.引物设计
根据盔形毕赤酵母的基因序列和pCAMBIA2300载体上的多克隆位点KpnI和PstI,设计以下引物,委托生工生物工程(上海)股份有限公司进行合成。
PgThioredoxin1-F:caatttactattctagggtaccATGTTCAATTTCCTTGTTCTCGTAC(SEQID No.9)
PgThioredoxin1-R:gtatgggtatctagactgcagTCAGGCCAACTTGTCCAAG(SEQ IDNo.10)
PgThioredoxin2-F:caatttactattctagggtaccATGCATTTATGGAATGTCTTTGTGG(SEQID No.11)
PgThioredoxin2-R:gtatgggtatctagactgcagTTACAACTCGTCCTTCTCTAGGTT(SEQ IDNo.12)
PgThioredoxin3-F:caatttactattctagggtaccATGCAATTTTCAATCAAGGCCG(SEQ IDNo.13)
PgThioredoxin3-R:gtatgggtatctagactgcagTTATAATTCATCATGAGCATCGGCT(SEQID No.14)
PgThioredoxin4-F:caatttactattctagggtaccATGCGTTTTTCTTTTGCCTT(SEQ IDNo.15)
PgThioredoxin4-R:gtatgggtatctagactgcagCTACAATTCGTCTTTTTCAATTTTC(SEQID No.16)
2.盔形毕赤酵母总RNA提取及cDNA合成
用北京酷来搏科技有限公司的真菌RNA提取试剂盒(RE781-50T)提取盔形毕赤酵母总RNA,具体步骤按照试剂盒说明书进行。
所得盔形毕赤酵母总RNA用PrimeScriptTM RT regent Kit试剂盒(Takara)反转录合成cDNA,具体步骤按照试剂盒说明书进行,用酶标仪Take3检测所得cDNA的浓度和质量,-20℃保存备用。
3.盔形毕赤酵母Thioredoxin类分泌蛋白编码基因序列克隆
以反转录合成的cDNA为模板,使用引物PgThioredoxin1-F和PgThioredoxin1-R,使用诺唯赞高保真酶P505试剂盒,PCR扩增核苷酸序列如SEQ ID No.5所示的PgThioredoxin1编码基因序列,具体步骤按照试剂盒说明书进行。
以反转录合成的cDNA为模板,使用引物PgThioredoxin2-F和PgThioredoxin2-R,使用诺唯赞高保真酶P505试剂盒,PCR扩增核苷酸序列如SEQ ID No.6所示的PgThioredoxin2编码基因序列,具体步骤按照试剂盒说明书进行。
以反转录合成的cDNA为模板,使用引物PgThioredoxin3-F和PgThioredoxin3-R,使用诺唯赞高保真酶P505试剂盒,PCR扩增核苷酸序列如SEQ ID No.7所示的PgThioredoxin3编码基因序列,具体步骤按照试剂盒说明书进行。
以反转录合成的cDNA为模板,使用引物PgThioredoxin4-F和PgThioredoxin4-R,使用诺唯赞高保真酶P505试剂盒,PCR扩增核苷酸序列如SEQ ID No.8所示的PgThioredoxin4编码基因序列,具体步骤按照试剂盒说明书进行。
PCR产物进行1%琼脂糖凝胶电泳(150V,30min),结果如图1所示,用无菌刀片切下目的条带,用胶回收试剂盒进行回收,具体步骤按照试剂盒说明书进行,回收产物-20℃保存备用。
4.质粒pCAMBIA2300的KpnI和PstI双酶切
将质粒pCAMBIA2300进行KpnI和PstI双酶切,酶切反应液置37℃金属浴中30min。
酶切产物进行琼脂糖凝胶电泳,用无菌刀片切下目的条带,用胶回收试剂盒进行回收,具体步骤按照试剂盒说明书进行,回收产物-20℃保存备用。
5.重组载体的构建
将回收的线性化质粒pCAMBIA2300与PgThioredoxin1编码基因序列使用诺唯赞同源重组酶进行连接,具体步骤按照试剂说明书进行,连接反应液置37℃金属浴中30min。同样的方法将PgThioredoxin2、PgThioredoxin3、PgThioredoxin4编码基因序列分别与线性化质粒pCAMBIA2300连接。
连接产物转化大肠杆菌感受态细胞。用质粒测序引物对转化子进行菌液PCR验证。验证结果为阳性的转化子委托生工生物工程(上海)股份有限公司进行测序。测序结果比对正确的重组质粒即为成功构建的重组载体pCAMBIA2300-PgThioredoxin1、pCAMBIA2300-PgThioredoxin2、pCAMBIA2300-PgThioredoxin3、pCAMBIA2300-PgThioredoxin4。
6.农杆菌工程菌的构建
提取重组载体pCAMBIA2300-PgThioredoxin1、pCAMBIA2300-PgThioredoxin2、pCAMBIA2300-PgThioredoxin3、pCAMBIA2300-PgThioredoxin4(电泳结果如图2所示,以质粒pCAMBIA2300为对照),分别加入农杆菌GV3101感受态细胞中,轻弹混匀,置冰上5min,液氮5min,37℃5min,冰上5min;加无抗LB液体培养基600μL,28℃、200rpm培育2~3h,4000rpm离心2min,弃上清,将余液均匀涂布于LB平板(含有25μg/L利福平和50μg/L卡那霉素);挑取单菌落至500μL LB液体培养基(含有25μg/L利福平和50μg/L卡那霉素)中,28℃、200rpm摇菌培养16h后检测目的片段。结果如图3所示,含有PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、Thioredoxin4编码基因的农杆菌即为成功构建的PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、PgThioredoxin4农杆菌工程菌。
7.农杆菌工程菌在柑橘果实上瞬时表达分泌蛋白诱导柑橘果实绿霉病抗病性的效果
分别将PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、PgThioredoxin4的农杆菌工程菌在LB培养基(含有25μg/L利福平和50μg/L卡那霉素)中培养过夜,当菌液OD600约为1时,4000rpm离心5min,将菌体用孵育液(1L孵育液含有5g葡萄糖,1.0663g MES(使用浓度5mM),0.760g Na3PO4·12H2O(使用浓度2mM),0.785mL 127.4mM AS母液(使用浓度0.1mM))重悬并调整OD600=0.8,等量混合基因沉默抑制子P19,28℃避光孵育2~3h,得农杆菌工程菌液;挑选大小均匀一致、色泽均匀、无机械伤和疤痕的柑橘果实,用2%次氯酸钠浸泡2min后用清水冲洗干净,自然风干,用75%酒精擦果实赤道部位,待酒精风干后,用1mL无菌枪头在果实赤道部位对立面各打一个孔,用1mL注射器(去掉针头)向孔中注射约0.5mL农杆菌工程菌液,以含有pCAMBIA2300质粒的农杆菌为对照(control);接种农杆菌工程菌1d后,在每个孔的右方1cm处再打一个孔,用移液枪接种10μL 1×104CFU/mL的指状青霉(Penicillium digitatum)孢子悬浮液,待菌液吸收后,单果套袋,避光放置于25℃的环境中,从第3天开始统计发病率和病斑直径。
结果如图4所示,含有PgThioredoxin1、PgThioredoxin2、PgThioredoxin3、PgThioredoxin4编码基因的农杆菌工程菌在柑橘果实上瞬时表达分泌蛋白PgThioredoxin1(氨基酸序列如SEQ ID No.1所示)、PgThioredoxin2(氨基酸序列如SEQ IDNo.2所示)、PgThioredoxin3(氨基酸序列如SEQ ID No.3所示)、PgThioredoxin4(氨基酸序列如SEQ ID No.4所示),能一定程度抑制柑橘果实绿霉病的发病和病斑直径的增长。
最后说明的是,以上优选实施例仅用以说明本发明的技术方案而非限制,尽管通过上述优选实施例已经对本发明进行了详细的描述,但本领域技术人员应当理解,可以在形式上和细节上对其作出各种各样的改变,而不偏离本发明权利要求书所限定的范围。
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agttcaggaa ttgttgagac catgaaagga actgttaaat cttacactaa aaaggtttct 480
ctttttaaat taccaaaatt tttatccaca agagatgaaa atgctctacc gcgtgtactt 540
ttcattactg ataaacttca aaattctcca atgtacaaaa tcttagctgt ggacttcaaa 600
ggaactttag aattcttcca tataggggta actgatgctt cccaaaagga gaaggtgaaa 660
agcttacttc ctgaattatc tgactccttt gaaatcccac aactcatagt gataagtcct 720
accgagggca ttgttcctta tgatggtgaa ttgaaaaaaa ccccaatctc ggaatttctt 780
actagatttg gtgcccctgt tgaaggtgat tttagtgaaa gaaatgagat tatccaaggc 840
ataaaaaagg gaacttacaa gagcttcaaa gactatcaca ggaagaaggc caagaaggcc 900
aagaaggaaa acctagagaa ggacgagttg taa 933
<210> 7
<211> 1593
<212> DNA
<213> 盔形毕赤酵母(Pichia galeiformis)
<400> 7
atgcaatttt caatcaaggc cgttgccgct atactggcag caatcacaaa tattgctgcc 60
gtctccgctg aaggcgcagt tgcccctgaa gattccgccg ttgttaaact tacggaggaa 120
accttcaagg acttcattgc caacaacgaa tatgttttag cagaattctt tgcaccatgg 180
tgtggccact gtaagaagtt gggccctgaa tttgcttccg ctgcagacag cttggccact 240
tccaacccag atatcaagtt ggctcaagtc gactgtactg aagacagaga cttgtgtgcc 300
gaattcgata tcagaggtta cccaacaatg aagatcttca gaggagaagt ctccactcct 360
tcggactact caggtcaaag acaatccgac gcaatcatca actacatggt caaacttacc 420
ttgccttctg tccagacctt cgaagatgca aaggctttag aagacgcttt ggatgatctt 480
tctgacactt tgatccttca agtcctccca gaaggtgttg agggcagtcc tgcaaatgca 540
actttctacg aagttgctga cagattaagg gaaaccttca ccttcggttc cacttctgat 600
gacaaatacg tctccaagta cgctaaatct tccaacaagc cagcatacgt cattttcaga 660
aacggtgaag aactcgatga tgcttccgtt tacaagggca aggacatcgg cgaggatccg 720
gaattacttg tcgatttcat tgacgtcgaa tctaaacctc ttttcggtga aatttctggt 780
gcaacgtacc aaagctatac ctccgccaat atcccattgg cttactactt ctacagcacc 840
aaggaagaac gtgatgctgc agacccattc atcaagaagt tagcaagaaa atacagaggt 900
gaaatcaact ttgtcggttt agacgccact caattcggta tgcacgctca aaacttgaac 960
atgcaggaag aattcccact ctttgtcatt cacgacctcg aattcaacaa gaagtatggt 1020
attgaccaag ctaagccact cgacaacaat gaaatcgcac aattcgtgca aaagttcaag 1080
gctggtaagt tggagccaat cgttaaatcc gaagcaattc ctgaggtaca aaactcaact 1140
ctttaccatc ttgttggtgc agaacatgac aaggtcatca aatctggtaa ggatgtcttt 1200
gtgaaatact ttgctccatg gtgcggtcac tgtaagagat tagctcctat tttcgaggaa 1260
cttgccgaac tttacgatgg caaagatgtc attgttgctg aaatggatca caccttgaac 1320
gatgtcgaag gcgttgaaat caagggttat ccaactctgg ttcttttccc agctgatggt 1380
tctgatccaa tctactatga tgaagcaaga actttagagg caatggccga cttcatcaaa 1440
gagaagggtt ccttgaaaat cgatgcgtta gcagattctg cagaagaagc tgcagtcgag 1500
gaaaccgcat cttccactgt gtcttccgaa actgagaaga agacggaaac ctccgctcca 1560
gcgactgaag ccgatgctca tgatgaatta taa 1593
<210> 8
<211> 1191
<212> DNA
<213> 盔形毕赤酵母(Pichia galeiformis)
<400> 8
atgcgttttt cttttgcctt gtcactactg agcttgtccg ctgcatgggc aagtaacatc 60
attgttgtaa atgatgagaa tttcgatgat gttgtcttga actccgataa gacatctttt 120
gttaaatttt ttgctgattg gtgcacacat tgtaaacaga tggttccaga atgggagaaa 180
ttagctgact cctacgcgga tgttgaagac gttcaaattg ttgaaattga tagtgacaag 240
tccagaacaa tcggcaagag gtacaatatt gcatcttacc caactcttaa actgttccgt 300
gctgatgctt tatcggaccc agttgacttt gatggaaaaa gagaatatga atattttgca 360
aactttttgc tgaatcaggt tggtgctaag ggtaagaaag tcggaccacc atcaaaggtt 420
gctcaagttc atgatggtaa tattgagaaa ttagtggaaa acaaggacag gtacgcaatg 480
cttcttttca caaaagagag ggactgtgtt gaatgcattg acgccaagaa agcttttgat 540
gagttatccc aagcattcca caaagaattg gataaaatca ttattggtga agtcaagaag 600
aatggtgatg aaccaacaga ttggactaga gagttatttg ccatttctga atatcctgcc 660
attatttttg tggaaaaggg tgacttacag aagtacgaag tttatggtgg aggaatttct 720
ggacctgaac tggtcaagtt cgtcaataac aggcttggaa ctaagagagc taccaatggg 780
ctgttggatt ctcaagctgg tattatccca gagatggaag aagcactagc gaagttcatt 840
ggctctaata ttgttgatag aagagcatac gtaaccgaat ttattgaaaa cttgaagaag 900
atcgacgacg ttttattcaa gaacgaagtt aagtattatg ccctgattgt caatcaattt 960
attgcaggta atatcgaata tgttgaatct gagttgacta agtttgagaa gaagattgca 1020
gacaagacag ttgattctgt tgaaaaagac ttagtaaaca tgaagctcaa cttattgaag 1080
cacatccaag aattgactac tccagaaact tatagagacg ctaaagaagt tatgaaggaa 1140
aaggaggagg ctaaggaagc ttcaaagaaa attgaaaaag acgaattgta g 1191
<210> 9
<211> 47
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 9
caatttacta ttctagggta ccatgttcaa tttccttgtt ctcgtac 47
<210> 10
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 10
gtatgggtat ctagactgca gtcaggccaa cttgtccaag 40
<210> 11
<211> 47
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
caatttacta ttctagggta ccatgcattt atggaatgtc tttgtgg 47
<210> 12
<211> 45
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
gtatgggtat ctagactgca gttacaactc gtccttctct aggtt 45
<210> 13
<211> 44
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 13
caatttacta ttctagggta ccatgcaatt ttcaatcaag gccg 44
<210> 14
<211> 46
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 14
gtatgggtat ctagactgca gttataattc atcatgagca tcggct 46
<210> 15
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 15
caatttacta ttctagggta ccatgcgttt ttcttttgcc tt 42
<210> 16
<211> 46
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 16
gtatgggtat ctagactgca gctacaattc gtctttttca attttc 46
Claims (8)
1.盔形毕赤酵母Thioredoxin类分泌蛋白,其特征在于,氨基酸序列如SEQ ID No.1所示。
2.权利要求1所述盔形毕赤酵母Thioredoxin类分泌蛋白的编码基因。
3.如权利要求2所述的编码基因,其特征在于,核苷酸序列如SEQ ID No.5所示。
4.含有权利要求2所述编码基因的重组表达载体。
5.如权利要求4所述的重组表达载体,其特征在于:是将核苷酸序列如SEQ ID No.5所示的盔形毕赤酵母Thioredoxin类分泌蛋白的编码基因克隆入植物表达载体pCAMBIA2300的多克隆位点KpnI和PstI之间而得到。
6.含有权利要求4所述重组表达载体的工程菌。
7.如权利要求6所述的工程菌,其特征在于,是将所述重组表达载体转入农杆菌GV3101中而得到。
8.权利要求1所述的盔形毕赤酵母Thioredoxin类分泌蛋白或权利要求6所述的工程菌在制备诱导柑橘果实绿霉病抗病性的保鲜剂中的应用。
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