植物诱抗蛋白HarpinN_（CSDS001）的基因克隆、表达和作用机制研究
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摘要
Harpin是一类由hrpN(hrpZ，hrpW)基因编码的能诱导植物产生系统抗性的重要致敏信号蛋白质，广泛存在于植物革兰氏阴性致病细菌和部分动物病原菌中，其基因不仅具有不同程度的同源性，更具有非常丰富的遗传多样性和差异性。本研究从分离携带hrpN基因的目的菌株Erwinia carotovora sp．carotovora CSDS001入手，研究了CSDS001菌株与hrpN基因表达产物相关的生物学特征，构建了CSDS001基因组文库，克隆到hrpN_(CSDS001)基因，实现HarpinN_(CSDS001)蛋白的高效表达，完成了对hrpN_(CSDS001)基因及其表达产物HarpinN_(CSDS001)蛋白的论证，采用多项控制性实验对HarpinN_(CSDS001)蛋白及HarpinN_(CSDS001)-Zn络合物的生物学活性和生物学效应进行了比较，通过对HarpinN_(CSDS001)蛋白及HarpinN_(CSDS001)-Zn络合物作用后的拟南芥全基因表达谱分析，探讨了HarpinN_(CSDS001)蛋白及HarpinN_(CSDS001)-Zn络合物诱导植物产生系统获得性抗性以及促进植物生长发育的可能机制。主要结果如下：
1、从感染软腐病大白菜组织中分离纯化到携带hrpN基因的CSDS001菌株，CSDS001具有对大白菜和西芹具有强致病性，分泌高活性Pel、Peh、Cel、Prt等植物细胞壁降解酶类，将2×10~8个／mlCSDS001菌注射接种到非寄主烟草叶片上，能诱导产生过敏反应(Hypersensitive Response，HR)。
2、将CSDS001基因组文库构建于质粒pLARF5，宿主菌为HB101，文库含有1000个菌落，每个菌落含20—30kb染色体DNA，从中筛选出的1.5—kb EcoRI片段含有CSDS001的hrpN_(CSDS001)基因。
3、序列测试表明hrpN_(CSDS001)基因编码区含1071个核苷酸，注册Genebank号AY939927，编码产物HarpinN_(CSDS001)蛋白含356个氨基酸残基，理论分子量35.55kD，理论等电点5.5。
4、将hrpN_(CSDS001)基因克隆到pET—28a(+)载体质粒T7启动子后，构建高效表达质粒pT7-hrpN_(CSDS001)，，将pT7-hrpN_(CSDS001)转入大肠杆菌JM109，完成了基因工程菌的构建，用IPTG能诱导hrpN_(CSDS001)基因高效表达。
5、Western杂交证明所克隆的hrpN_(CSDS001)基因表达产物为HarpinN_(CSDS001)蛋白；Northern杂交证明CSDS001菌株携带的hrpN_(CSDS001)基因表达受外界环境碳源、氮源和pH值等调控。
6、对HarpinN_(CSDS001)蛋白的分离纯化表明，工程菌中高效表达的HarpinN_(CSDS001)蛋白是以包含体形式存在于细胞质中，能诱导烟草叶片产生过敏反应，证明以包含体形式存在的HarpinN_(CSDS001)蛋白具有生物学活性。
7、一定条件下，HarpinN_(CSDS001)蛋白与锌离子发生络合反应，形成的HarpinN_(CSDS001)—Zn络合物与HarpinN_(CSDS001)蛋白具有同样的生物学活性，能诱导烟草叶片产生过敏反应。
8、分别将30mg／L浓度的HarpinN_(CSDS001)—Zn络合物和HarpinN_(CSDS001)蛋白施用到辣椒、茄子、黄瓜、马铃薯和西红柿，进行田间控制性药效试验。结果显示，施用HarpinN_(CSDS001)-Zn络合物使几种蔬菜产量分别增加13.69％、16.52％、18.86％、11.55％、11.64％；施用HarpinN_(CSDS001)蛋白增产分别为11.73％、13.27％、16.3％、8.26％、10.25％。经过对参试作物病虫害、株高、生育期等的调查分析，增产主要原因是HarpinN_(CSDS001)-Zn和HarpinN_(CSDS001)蛋白诱导作物系统抗性，减少作物因病虫害造成的损失，此外，在一定程度上还能促进作物生长发育、增加生物量、诱导提前开花结果、延长结果期，促进增加作物产量。表明HarpinN_(CSDS001)—Zn比HarpinN_(CSDS001)蛋白具有更强的生物学功效。
9、用Cluster 3.0，TreeView 1.60软件聚类分析HarpinN_(CSDS001)蛋白处理后的拟南芥全基因谱，显示在分别处理后第3h、12h、24h、36h和48h，有52个基因(聚为5类)的表达变化与HarpinN_(CSDS001)蛋白作用持续相关，其中第4类和第5类基因功能与植物抗逆性和生长发育相关。
10、对HarpinN_(CSDS001)蛋白诱导拟南芥发生显著表达差异的基因数量分析表明，在作用后第3h、12h、24h、36h和48h时发生显著表达变化(log ratio≥1或≤-1)的基因数分别为912、1787、2393、1833和1755。表明，HarpinN_(CSDS001)作用后的一个时期内，与若干生理过程相关的基因发生显著表达差异，且发生显著表达差异的基因数量逐步上升，在作用后第24h左右诱导发生显著表达差异的基因数量最高，同时，诱导发生显著表达差异的基因数具有时间上的持续性。
11、对被诱导发生显著表达差异的转录因子家族基因分析表明，有13个转录因子家族：ZIM、BES1、TCP、C2C2、AP2／EREBP、WRKY、bHLH、bZIP、GARP、MYB、NAC、HB、C2H2与HarpinN_(CSDS001)蛋白作用相关。这些转录因子主要参与调控植物抗性反应、光合作用、生长发育、开花等生理反应。
12、利用http://www.genome.ad.jp/提供的KEGG2分析工具，对HarpinN_(CSDS001)蛋白作用后类黄酮合成相关基因表达分析，揭示了查尔酮合成酶和黄烷酮—3—羟基化酶基因表达上调，从转录水平证明HarpinN_(CSDS001)蛋白诱导类黄酮合成。
13、HarpinN_(CSDS001)—Zn作用于拟南芥后第24h，诱导发生显著表达变化基因数达到2741，更有35个转录因子家族基因的表达变化与HarpinN_(CSDS001)—Zn的作用相关，且查尔酮合成酶和黄烷酮—3—羟基化酶基因表达也上调。这些结果表明，可能HarpinN_(CSDS001)—Zn诱导的与若干生理过程相关基因的显著表达差异是更为复杂的过程，较之HarpinN_(CSDS001)蛋白，明显具有更强生物学功效，这与田间控制性试验结果相一致。
Harpins are a class of proteins which is ecoded by hrpN (hrpZ,hrpW) andproduced in nature by certain plant Gram-negative bacteria and some animalpathogens. They cause hypersensitive response (HR) of plants and elicit a complexnatural defense mechanism. There are homolog and wide diversity among the hrpgenes. This work began with the isolation and characterization of wild type bacteriaErwinia carotovora subsp.carotovora strains CSDS001, then the genomic libraries ofthe CSDS001 was constructed and the hrpN_(CSDS001) gene was cloned. HarpinN_(CSDS001)protein was overproduced in E.coli JM109. The biological activities andmultifunctionality of HarpinN_(CSDS001) and HarpinN_(CSDS001)-Zn were tested and analyzedby series controlled experiments. Potential mechanism of inducing systemic acquiredresistance (SAR) and promoting plant growth by HarpinN_(CSDS001) andHarpinN_(CSDS001)-Zn was explored by analyzing Arabidopsis treated by the two proteinsby Genechip. The main results are:
1、CSDS001 was isolated from soft-rotting Chinese cabbage. It caused soft rot inChinese cabbage and celery, and produced exoenzymes such as Pel、Peh、Cel、Prt. 2×10~8 cfu/ml CSDS001 inoculated into tobacco leaves causing HR.
2、CSDS001 genomic library was constructed in vector pLARF5, harbored in E.colistrain HB101 and contained 1000 colonies. Each colony possessed 20—30kb ofchromosomal DNA. Southern blot revealed 1.5-kb EcoRI segments containinghrpN_(CSDS001).
3、The hrpN_(CSDS001) nucleotide sequence is 1071bp. Registered number is AY939927.HarpinN_(CSDS001) contains 556 amino acids and its molecular weight is 35.55kD andpI is 5.5.
4、pT7-hrpN_(CSDS001), in protein over expression vector, was constructed by cloninghrpN_(CSDS001) into pET—28a (+). Then the pT7-hrpN_(CSDS001) was transferred intoE.coli strain JM109. IPTG induced hrpN_(CSDS001) to over express.
5、The over expressed protein was identified as HarpinN_(CSDS001) by Western blot.Northern blot analysis showed the expression of hrpN_(CSDS001) were influenced by pH、carbon and nitrogen source.
6、Over expressed HarpinN_(CSDS001) formed as inclusion body and can elicit HR intobacco leaves. It means this protein in inclusion body form also had biologicalactivity.
7、In some conditions, HarpinN_(CSDS001) chelated with Zn~(2+) and formedHarpinN_(CSDS001)-Zn, which had the same biological activity-eliciting HR intobacco leaves.
8、30mg/L HarpinN_(CSDS001)-Zn and 30mg/L HarpinN_(CSDS001) were applied oncapsicum、aubergine、cucumber、potato、and tomato respectively. The resultsshowed that the yields increased by HarpinN_(CSDS001)—Zn were 13.69%、16.52%、18.86%、11.55%、11.64% respectively and by HarpinN_(CSDS001) were 11.73%、13.27%、16.3%、8.26%、10.25%. Statistical analysis of the plant disease、growthand development showed that both HarpinN_(CSDS001)—Zn and HarpinN_(CSDS001) mayinduce plant defense, boost growth and promote development, moreoverHarpinN_(CSDS001)-Zn had more powerful biological functions than HarpinN_(CSDS001).
9、Based on the cluster analysis of mRNA level tested by Genechip of Arabidopsistreated by HarpinN_(CSDS001), 52 genes (clustered into five groups) were regulated in3h、12h、24h、36h and 48h, and the fourth and fifth group genes related to thefunctions of plant defense、growth and development.
10、Analysis of genes which expression changed significantly (log ratio≤1 or≥1) showed that after treatment with HarpinN_(CSDS001) at 3h、12h、24h、36h and 48h,there were 912、1787、2393、1833 and 1755 genes correlatively changedsignificantly. The results revealed that the level of mRNA regulation byHarpinN_(CSDS001) increased gradually, most of the reaction happened at 24h andcontinual physiological responses also were drawn out.
11、Analysis of transcription factors (TF) families showed that 13 TF familiesresponded to HarpinN_(CSDS001). They were ZIM、BES1、TCP、C2C2、AP2/EREBP、WRKY、bHLH、bZIP、GARP、MYB、NAC、HB、C2H2. The families mainlyregulate reactions about plant defense, photosynthesis, growth, development andflowering.
12、Used KEGG2, provided by http://www.genome.ad.jp/ to analyze theflavonoid biosynthesis pathway revealing that chalcone synthase (CHS) gene andflavanone 3-hydroxylase (F3H) gene expression increased.
13、24h after treatment by HarpinN_(CSDS001)-Zn to Arabidopsis, there were 2741genes regulated significantly, 35 TF families responding to HarpinN_(CSDS001)-Zn andthe chalcone synthase gene and flavanone 3-hydroxylase gene expression alsoincreased. The results indicated that HarpinN_(CSDS001)-Zn had more powerfulbiological function than HarpinN_(CSDS001). It was consistent with the results ofcontrolled experiments in the fields.
1、从感染软腐病大白菜组织中分离纯化到携带hrpN基因的CSDS001菌株，CSDS001具有对大白菜和西芹具有强致病性，分泌高活性Pel、Peh、Cel、Prt等植物细胞壁降解酶类，将2×10~8个／mlCSDS001菌注射接种到非寄主烟草叶片上，能诱导产生过敏反应(Hypersensitive Response，HR)。
2、将CSDS001基因组文库构建于质粒pLARF5，宿主菌为HB101，文库含有1000个菌落，每个菌落含20—30kb染色体DNA，从中筛选出的1.5—kb EcoRI片段含有CSDS001的hrpN_(CSDS001)基因。
3、序列测试表明hrpN_(CSDS001)基因编码区含1071个核苷酸，注册Genebank号AY939927，编码产物HarpinN_(CSDS001)蛋白含356个氨基酸残基，理论分子量35.55kD，理论等电点5.5。
4、将hrpN_(CSDS001)基因克隆到pET—28a(+)载体质粒T7启动子后，构建高效表达质粒pT7-hrpN_(CSDS001)，，将pT7-hrpN_(CSDS001)转入大肠杆菌JM109，完成了基因工程菌的构建，用IPTG能诱导hrpN_(CSDS001)基因高效表达。
5、Western杂交证明所克隆的hrpN_(CSDS001)基因表达产物为HarpinN_(CSDS001)蛋白；Northern杂交证明CSDS001菌株携带的hrpN_(CSDS001)基因表达受外界环境碳源、氮源和pH值等调控。
6、对HarpinN_(CSDS001)蛋白的分离纯化表明，工程菌中高效表达的HarpinN_(CSDS001)蛋白是以包含体形式存在于细胞质中，能诱导烟草叶片产生过敏反应，证明以包含体形式存在的HarpinN_(CSDS001)蛋白具有生物学活性。
7、一定条件下，HarpinN_(CSDS001)蛋白与锌离子发生络合反应，形成的HarpinN_(CSDS001)—Zn络合物与HarpinN_(CSDS001)蛋白具有同样的生物学活性，能诱导烟草叶片产生过敏反应。
8、分别将30mg／L浓度的HarpinN_(CSDS001)—Zn络合物和HarpinN_(CSDS001)蛋白施用到辣椒、茄子、黄瓜、马铃薯和西红柿，进行田间控制性药效试验。结果显示，施用HarpinN_(CSDS001)-Zn络合物使几种蔬菜产量分别增加13.69％、16.52％、18.86％、11.55％、11.64％；施用HarpinN_(CSDS001)蛋白增产分别为11.73％、13.27％、16.3％、8.26％、10.25％。经过对参试作物病虫害、株高、生育期等的调查分析，增产主要原因是HarpinN_(CSDS001)-Zn和HarpinN_(CSDS001)蛋白诱导作物系统抗性，减少作物因病虫害造成的损失，此外，在一定程度上还能促进作物生长发育、增加生物量、诱导提前开花结果、延长结果期，促进增加作物产量。表明HarpinN_(CSDS001)—Zn比HarpinN_(CSDS001)蛋白具有更强的生物学功效。
9、用Cluster 3.0，TreeView 1.60软件聚类分析HarpinN_(CSDS001)蛋白处理后的拟南芥全基因谱，显示在分别处理后第3h、12h、24h、36h和48h，有52个基因(聚为5类)的表达变化与HarpinN_(CSDS001)蛋白作用持续相关，其中第4类和第5类基因功能与植物抗逆性和生长发育相关。
10、对HarpinN_(CSDS001)蛋白诱导拟南芥发生显著表达差异的基因数量分析表明，在作用后第3h、12h、24h、36h和48h时发生显著表达变化(log ratio≥1或≤-1)的基因数分别为912、1787、2393、1833和1755。表明，HarpinN_(CSDS001)作用后的一个时期内，与若干生理过程相关的基因发生显著表达差异，且发生显著表达差异的基因数量逐步上升，在作用后第24h左右诱导发生显著表达差异的基因数量最高，同时，诱导发生显著表达差异的基因数具有时间上的持续性。
11、对被诱导发生显著表达差异的转录因子家族基因分析表明，有13个转录因子家族：ZIM、BES1、TCP、C2C2、AP2／EREBP、WRKY、bHLH、bZIP、GARP、MYB、NAC、HB、C2H2与HarpinN_(CSDS001)蛋白作用相关。这些转录因子主要参与调控植物抗性反应、光合作用、生长发育、开花等生理反应。
12、利用http://www.genome.ad.jp/提供的KEGG2分析工具，对HarpinN_(CSDS001)蛋白作用后类黄酮合成相关基因表达分析，揭示了查尔酮合成酶和黄烷酮—3—羟基化酶基因表达上调，从转录水平证明HarpinN_(CSDS001)蛋白诱导类黄酮合成。
13、HarpinN_(CSDS001)—Zn作用于拟南芥后第24h，诱导发生显著表达变化基因数达到2741，更有35个转录因子家族基因的表达变化与HarpinN_(CSDS001)—Zn的作用相关，且查尔酮合成酶和黄烷酮—3—羟基化酶基因表达也上调。这些结果表明，可能HarpinN_(CSDS001)—Zn诱导的与若干生理过程相关基因的显著表达差异是更为复杂的过程，较之HarpinN_(CSDS001)蛋白，明显具有更强生物学功效，这与田间控制性试验结果相一致。
Harpins are a class of proteins which is ecoded by hrpN (hrpZ,hrpW) andproduced in nature by certain plant Gram-negative bacteria and some animalpathogens. They cause hypersensitive response (HR) of plants and elicit a complexnatural defense mechanism. There are homolog and wide diversity among the hrpgenes. This work began with the isolation and characterization of wild type bacteriaErwinia carotovora subsp.carotovora strains CSDS001, then the genomic libraries ofthe CSDS001 was constructed and the hrpN_(CSDS001) gene was cloned. HarpinN_(CSDS001)protein was overproduced in E.coli JM109. The biological activities andmultifunctionality of HarpinN_(CSDS001) and HarpinN_(CSDS001)-Zn were tested and analyzedby series controlled experiments. Potential mechanism of inducing systemic acquiredresistance (SAR) and promoting plant growth by HarpinN_(CSDS001) andHarpinN_(CSDS001)-Zn was explored by analyzing Arabidopsis treated by the two proteinsby Genechip. The main results are:
1、CSDS001 was isolated from soft-rotting Chinese cabbage. It caused soft rot inChinese cabbage and celery, and produced exoenzymes such as Pel、Peh、Cel、Prt. 2×10~8 cfu/ml CSDS001 inoculated into tobacco leaves causing HR.
2、CSDS001 genomic library was constructed in vector pLARF5, harbored in E.colistrain HB101 and contained 1000 colonies. Each colony possessed 20—30kb ofchromosomal DNA. Southern blot revealed 1.5-kb EcoRI segments containinghrpN_(CSDS001).
3、The hrpN_(CSDS001) nucleotide sequence is 1071bp. Registered number is AY939927.HarpinN_(CSDS001) contains 556 amino acids and its molecular weight is 35.55kD andpI is 5.5.
4、pT7-hrpN_(CSDS001), in protein over expression vector, was constructed by cloninghrpN_(CSDS001) into pET—28a (+). Then the pT7-hrpN_(CSDS001) was transferred intoE.coli strain JM109. IPTG induced hrpN_(CSDS001) to over express.
5、The over expressed protein was identified as HarpinN_(CSDS001) by Western blot.Northern blot analysis showed the expression of hrpN_(CSDS001) were influenced by pH、carbon and nitrogen source.
6、Over expressed HarpinN_(CSDS001) formed as inclusion body and can elicit HR intobacco leaves. It means this protein in inclusion body form also had biologicalactivity.
7、In some conditions, HarpinN_(CSDS001) chelated with Zn~(2+) and formedHarpinN_(CSDS001)-Zn, which had the same biological activity-eliciting HR intobacco leaves.
8、30mg/L HarpinN_(CSDS001)-Zn and 30mg/L HarpinN_(CSDS001) were applied oncapsicum、aubergine、cucumber、potato、and tomato respectively. The resultsshowed that the yields increased by HarpinN_(CSDS001)—Zn were 13.69%、16.52%、18.86%、11.55%、11.64% respectively and by HarpinN_(CSDS001) were 11.73%、13.27%、16.3%、8.26%、10.25%. Statistical analysis of the plant disease、growthand development showed that both HarpinN_(CSDS001)—Zn and HarpinN_(CSDS001) mayinduce plant defense, boost growth and promote development, moreoverHarpinN_(CSDS001)-Zn had more powerful biological functions than HarpinN_(CSDS001).
9、Based on the cluster analysis of mRNA level tested by Genechip of Arabidopsistreated by HarpinN_(CSDS001), 52 genes (clustered into five groups) were regulated in3h、12h、24h、36h and 48h, and the fourth and fifth group genes related to thefunctions of plant defense、growth and development.
10、Analysis of genes which expression changed significantly (log ratio≤1 or≥1) showed that after treatment with HarpinN_(CSDS001) at 3h、12h、24h、36h and 48h,there were 912、1787、2393、1833 and 1755 genes correlatively changedsignificantly. The results revealed that the level of mRNA regulation byHarpinN_(CSDS001) increased gradually, most of the reaction happened at 24h andcontinual physiological responses also were drawn out.
11、Analysis of transcription factors (TF) families showed that 13 TF familiesresponded to HarpinN_(CSDS001). They were ZIM、BES1、TCP、C2C2、AP2/EREBP、WRKY、bHLH、bZIP、GARP、MYB、NAC、HB、C2H2. The families mainlyregulate reactions about plant defense, photosynthesis, growth, development andflowering.
12、Used KEGG2, provided by http://www.genome.ad.jp/ to analyze theflavonoid biosynthesis pathway revealing that chalcone synthase (CHS) gene andflavanone 3-hydroxylase (F3H) gene expression increased.
13、24h after treatment by HarpinN_(CSDS001)-Zn to Arabidopsis, there were 2741genes regulated significantly, 35 TF families responding to HarpinN_(CSDS001)-Zn andthe chalcone synthase gene and flavanone 3-hydroxylase gene expression alsoincreased. The results indicated that HarpinN_(CSDS001)-Zn had more powerfulbiological function than HarpinN_(CSDS001). It was consistent with the results ofcontrolled experiments in the fields.
引文
方中达。植病研究方法．中国农业出版社
方中达．1996年，中国农业植物病害，中国农业出版社
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