人参锈腐病化学诱导抗性机制及其病原菌分子检测技术研究
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摘要
人参(Panax ginseng C.A. Meyer)为五加科人参属多年生草本药用植物,是我国传统的名贵药材,被誉为“药中之王”。人参锈腐病是由人参锈腐病菌(Cylindrocarpon destructans (Zinns.) Scholten)引起的根部病害之一,平均发病率20%-30%,严重影响人参产量和加工质量,造成极大的经济损失。随着国际市场人参需求量的不断上升,人类健康意识和环保意识的增强,通过降低人参的农药残留、倡导安全无污染的GAP栽培模式来提高我国人参的市场竞争力已成为必然的发展趋势。利用植物诱导抗病性的原理来提高人参植株的抗性,利用分子检测手段对病害进行早期诊断和病菌种群动态监测,可能为人参锈腐病的防治提供一条新途径。本文以植物诱导抗病性为着眼点,从生理生化反应、差异蛋白质组学、分子检测等角度系统研究了外源化学诱导因子诱导人参抗锈腐病的机制,建立了人参锈腐病菌的分子检测体系,主要研究结果如下：
1.明确了水杨酸、肉桂酸、阿魏酸、苯甲酸、对羟基苯甲酸和茉莉酸甲酯6种化学诱导因子对人参锈腐病菌的影响。通过在培养基内人工添加不同化学诱导因子培养人参锈腐病菌,来考察这些化学因子对人参锈腐病菌的影响。结果表明肉桂酸、阿魏酸、苯甲酸显著抑制人参锈腐病菌的菌丝生长和孢子萌发；水杨酸、茉莉酸甲酯在低浓度时不影响锈腐病菌的生长,浓度高于200μg mL-1时显著抑制其生长；而对羟基苯甲酸则对人参锈腐病菌没有影响。这些物质在人参锈腐病的化学诱导抗病性中发挥着重要作用。
2.室内接种试验证明了水杨酸和茉莉酸甲酯在低浓度处理人参时可以对人参锈腐病菌产生诱导抗性。在所选用的6种化学诱导因子中,水杨酸和茉莉酸甲酯的诱抗效果最为显著。经200μg mL-1的水杨酸和茉莉酸甲酯处理后,不仅对人参生长有一定的促进作用,而且对人参锈腐病的防效分别达到了67.9%和56.6%。而其他几种化学因子对锈腐病的防效不显著,并会对人参生长产生不同程度的不良影响。因此,本文筛选水杨酸和茉莉酸甲酯作为进一步研究的化学诱导因子。
3.首次证明了外源水杨酸(SA)和茉莉酸甲酯(MeJA)处理可以显著提高人参植株的诱导抗病性,并明确了其诱导人参抗人参锈腐病的生理生化机制。通过温室盆栽试验测定与抗性相关的生理生化指标发现,外源SA. MeJA能有效降低人参根内MDA含量和细胞膜电解质外渗率,提高脯氨酸和可溶性糖的含量及总酚含量,人参根系PAL、CAT、PPO、POD活性较对照均上升,p-1,3-葡聚糖酶和几丁质酶活性也较对照增强,说明SA. MeJA诱导的植物抗病性可能与植物的系统获得抗性有关。外源施入SA和MeJA可减轻人参锈腐病的发病率和病害严重度,发病率分别下降了39.1%和34.8%,明显提高了人参植株抗性。SA诱导人参抗人参锈腐病的效果较MeJA更为显著,因此选择SA作为进一步研究人参锈腐病化学诱导抗性的诱导因子。
4.采用植物组培技术,成功诱导出人参根愈伤组织并建立了生长旺盛的人参悬浮细胞系,为下一步运用SA诱导人参抗人参锈腐病的差异蛋白质组学研究提供了稳定、均一的试验试材。愈伤组织诱导选用MS培养基,外源激素种类及浓度为3mg/L2,4-D和0.2mg/L KT,培养基蔗糖浓度为0.8%,pH值在5.8-6.0之间。继代培养添加的激素配比及浓度为2mg/L2,4-D和0.5mg/L BA。悬浮细胞液体培养基的激素浓度及配比与愈伤组织继代培养基的激素水平相同,人参悬浮细胞于摇床悬浮振荡培养,培养条件为110rpm,24±2℃
5.首次采用蛋白质组学分析方法对SA诱导人参抗人参锈腐病的差异蛋白组进行了分析,明确了SA对人参抗锈腐病蛋白表达的调控作用。建立了适合人参悬浮细胞蛋白质组学分析的高通量、高分辨率的双向电泳技术体系：采用改良酚提取法制备人参悬浮细胞总蛋白,裂解缓冲液为9M尿素、2M硫脲、2%IPG Buffer、4%CHAPS、1%TBP.65mMDTT, pH5-8IPG胶条,SDS-PAGE分离胶浓度为12%。通过双向电泳技术获得了清水对照、SA处理、C. destructans处理及SA+C. destructans处理的人参悬浮细胞蛋白2-DE图谱。在凝胶上分别检测到800多个蛋白点。对蛋白质丰度变化在2倍以上、重复性好的24个差异表达蛋白点进行MALDI-TOF-MS质谱鉴定分析,其中共成功鉴定了23个蛋白点,去掉重复的蛋白共鉴定出22种蛋白质。在这些鉴定的蛋白点中,以对照为参考胶,只在SA处理中上调表达的蛋白点有12个,下调表达的蛋白点2个,特异表达的蛋白点1个；只在SA+C.destructans处理中上调表达的蛋白点2个,下调表达的蛋白点1个；,在三个处理中都上调表达的蛋白点5个,下调表达的蛋白点1个。这些特异表达的蛋白涉及植物自身的防卫反应、信号转导、能量代谢和转录调控等方面。其中热激蛋白60、肉桂醇脱氢酶、单脱水抗坏血酸还原酶、甲氧基转移酶都与防御反应正相关。
6.首次建立了人参锈腐病菌分子检测体系,为人参锈腐病的早期诊断和病原菌种群动态监测奠定了理论基础。该体系在对人参锈腐病菌ITS区进行PCR扩增、测序及序列分析基础上,设计了特异性引物CD-F/CD-R,可以用于人参主要病原真菌及常见土壤习居菌的PCR检测。只有以C. destructans基因组DNA为模板的体系中能扩增出一条450bp左右的条带,而其他菌株及阴性对照均无特异性条带产生。利用该特异性引物能从罹病组织和人工接种C. destructans的土壤DNA中扩增出特异性条带,而对照及健康人参组织均无扩增产物,表明该特异性引物可以用于人参锈腐病菌的分子检测和早期诊断。
Ginseng (Panax ginseng C.A. Meyer) is a perennial herbaceous plant in the family Araliaceae which has been grown in northeast China for centuries, is known as the king of medicine. Cylindrocarpon root rot of ginseng caused by Cylindrocarpon destructans is the main disease which led to serious crop yield losses, and the average loss is up to20%-30%. The soilborne disease is very difficult to control. The use of high toxic pesticide is forbidden because of GAP cultivation. It maybe provide a new way to control Cylindrocarpon root rot of ginseng by using the principle of plant induced disease resistance to improve the resistance of ginseng plant and using molecular detection method to monitor pathogen population. The inducing mechanism of Cylindrocarpon root rot was studied and the pathogen molecular detection system was established based on plant induced disease resistance from the physiological and biochemical reaction, proteomics, and molecular detection. The results were as follows:
1. The effects of salicylic acid, cinnamic acid, ferulic acid, benzoic acid, p-hydroxybenzoic acid and methyl jasmonate on C. destructans were studied. These chemicals were added into media to incubate C. destructans to investigate their effects on the pathogen. The results indicated that the growth and spore germination of C. destructans were suppressed by cinnamic acid, ferulic acid, benzoic acid, while p-hydroxybenzoic acid had no influence on the pathogen. At low concentration of salicylic acid and methyl jasmonate, the growth of C. destructans was hardly affected, while at higher concentrations (>200μg mL-1) the growth of pathogen was prohibited.
2. The resistance of ginseng to C. destructans could be induced by low concentrations of SA and MeJA. Of these6kinds of chemicals, the induced effects of SA and MeJA were better than other chemicals. It showed that ginseng treated with low concentration of salicylic acid and jasmonic acid methyl jasmonate could enhance the resistance of ginseng plant itself, the control effects were up to67.9%and56.6%, respectively, and the two kinds of chemical factors in low concentration had promoting effects on the growth of ginseng. However, the other chemicals had harmful effects on ginseng growth. Therefore, SA and MeJA were chosen for next study.
3. The resistance of ginseng induced by SA and MeJA was firstly demonstrated. The physiological and biochemical mechanisms of ginseng resistant to Cylindrocarpon root rot induced by exogenous salicylic acid and methyl jasmonate were studied. SA and MeJA treatment could effectively decrease the contents of MDA and relative electricity conductivity, but increase the contents of proline, soluble sugar and total phenols. The activities of PAL, CAT, PPO, POD,(β-1,3-glucanase and chitinase could be increased significantly in the ginseng roots induced by SA and MeJA than those in control. The incidence and disease severity were reduced with ginseng plant resistance increased obviously due to exogenous applied SA and MeJA, and the incidences were reduced by39.1%and34.8%, respectively. It demonstrated that SA and MeJA could induce the system resistance of ginseng to C. destructans. In addition, the impact of SA was better than MeJA. Thus, SA is chosen as the chemical factor for further study.4. A stable and vigorous growth of ginseng suspension cell lines was established using plant tissue culture technology. The establishment of the system provides a stable and uniform test material in ginseng proteomics research. MS culture medium was used in callus induction, supplement with3mg/L2,4-D and0.2mg/L KT,0.8%sucrose, pH5.8to6.0. Successive transfer culture added2mg/L2,4-D and0.5mg/L BA. The hormone of suspension cell liquid medium was at the same levels of successive transfer culture. The culture condition of callus is25℃, dark culture. The suspension cell masses were grown in the flasks on a rotary shaker (110rpm)at24±2℃.
5. The proteome of salicylic acid inducing ginseng suspension cell was firstly studied by two-dimensional gel electrophoresis technology. A high-throughput and high-resolution two-dimensional gel electrophoresis system was established for suspension-cultured ginseng cells proteins. The proteins were extracted by using the phenol extraction, the elements of lysis buffer were composed of9M urea,2M Thiourea,2%IPG buffer,4%CHAPS,65mM DTT and1%TBP. The pH5-8IPG strip was used. SDS-PAGE was performed in12%polyacrylamide gels. The proteins related to resistance were identified.2-DE was adopted to separate the differentially expressed proteins from suspension-cultured ginseng cells induced by SA. About800protein spots were detected on the2-DE gels. Twenty-four differentially expressed proteins that changes in protein abundance in more than2times and good repeatability were analyzed by MALDI-TOF-MS, of23proteins were successfully identified. In the absence of pathogen,12proteins had increased levels, two proteins had decreased levels and one protein was newly induced in suspension cultures treated directly with SA, in comparison to the control. Two proteins were up-regulated and one protein was down-regulated after SA treatment in the presence of C. destructans. Five proteins were up-regulated and only one protein was down-regulated in suspension cultures were differentially expressed after both SA treatment and pathogen infection. These identified proteins were predicted to be involved in defense and stress responses, energy and metabolism, signal transduction/transcription, protein synthesis and metabolism, photosynthesis and others. The proteins related to defense responses, such as Chaperonin60, monodehydroascorbate reductase, CAD and orcinol O-methyltransferase, were more abundant in suspension cultures of ginseng roots after application of S A.
6. The molecular detection of Cylindrocarpon root rot of ginseng was established. A pair of primers specific for C. destructans was designed from comparisons of the internal transcribed spacer (ITS) regions1and2of isolates of diverse origins with sequences from commonly encountered fungi in soil. Under stringent PCR conditions, primers CD-F/CD-R amplify a450bp fragment from C. destructans DNA but not from other species or negative control. And C. destructans could be specifically detected with CD-F/CD-R from ginseng tissues and soil samples inoculated artificially.
1.明确了水杨酸、肉桂酸、阿魏酸、苯甲酸、对羟基苯甲酸和茉莉酸甲酯6种化学诱导因子对人参锈腐病菌的影响。通过在培养基内人工添加不同化学诱导因子培养人参锈腐病菌,来考察这些化学因子对人参锈腐病菌的影响。结果表明肉桂酸、阿魏酸、苯甲酸显著抑制人参锈腐病菌的菌丝生长和孢子萌发；水杨酸、茉莉酸甲酯在低浓度时不影响锈腐病菌的生长,浓度高于200μg mL-1时显著抑制其生长；而对羟基苯甲酸则对人参锈腐病菌没有影响。这些物质在人参锈腐病的化学诱导抗病性中发挥着重要作用。
2.室内接种试验证明了水杨酸和茉莉酸甲酯在低浓度处理人参时可以对人参锈腐病菌产生诱导抗性。在所选用的6种化学诱导因子中,水杨酸和茉莉酸甲酯的诱抗效果最为显著。经200μg mL-1的水杨酸和茉莉酸甲酯处理后,不仅对人参生长有一定的促进作用,而且对人参锈腐病的防效分别达到了67.9%和56.6%。而其他几种化学因子对锈腐病的防效不显著,并会对人参生长产生不同程度的不良影响。因此,本文筛选水杨酸和茉莉酸甲酯作为进一步研究的化学诱导因子。
3.首次证明了外源水杨酸(SA)和茉莉酸甲酯(MeJA)处理可以显著提高人参植株的诱导抗病性,并明确了其诱导人参抗人参锈腐病的生理生化机制。通过温室盆栽试验测定与抗性相关的生理生化指标发现,外源SA. MeJA能有效降低人参根内MDA含量和细胞膜电解质外渗率,提高脯氨酸和可溶性糖的含量及总酚含量,人参根系PAL、CAT、PPO、POD活性较对照均上升,p-1,3-葡聚糖酶和几丁质酶活性也较对照增强,说明SA. MeJA诱导的植物抗病性可能与植物的系统获得抗性有关。外源施入SA和MeJA可减轻人参锈腐病的发病率和病害严重度,发病率分别下降了39.1%和34.8%,明显提高了人参植株抗性。SA诱导人参抗人参锈腐病的效果较MeJA更为显著,因此选择SA作为进一步研究人参锈腐病化学诱导抗性的诱导因子。
4.采用植物组培技术,成功诱导出人参根愈伤组织并建立了生长旺盛的人参悬浮细胞系,为下一步运用SA诱导人参抗人参锈腐病的差异蛋白质组学研究提供了稳定、均一的试验试材。愈伤组织诱导选用MS培养基,外源激素种类及浓度为3mg/L2,4-D和0.2mg/L KT,培养基蔗糖浓度为0.8%,pH值在5.8-6.0之间。继代培养添加的激素配比及浓度为2mg/L2,4-D和0.5mg/L BA。悬浮细胞液体培养基的激素浓度及配比与愈伤组织继代培养基的激素水平相同,人参悬浮细胞于摇床悬浮振荡培养,培养条件为110rpm,24±2℃
5.首次采用蛋白质组学分析方法对SA诱导人参抗人参锈腐病的差异蛋白组进行了分析,明确了SA对人参抗锈腐病蛋白表达的调控作用。建立了适合人参悬浮细胞蛋白质组学分析的高通量、高分辨率的双向电泳技术体系：采用改良酚提取法制备人参悬浮细胞总蛋白,裂解缓冲液为9M尿素、2M硫脲、2%IPG Buffer、4%CHAPS、1%TBP.65mMDTT, pH5-8IPG胶条,SDS-PAGE分离胶浓度为12%。通过双向电泳技术获得了清水对照、SA处理、C. destructans处理及SA+C. destructans处理的人参悬浮细胞蛋白2-DE图谱。在凝胶上分别检测到800多个蛋白点。对蛋白质丰度变化在2倍以上、重复性好的24个差异表达蛋白点进行MALDI-TOF-MS质谱鉴定分析,其中共成功鉴定了23个蛋白点,去掉重复的蛋白共鉴定出22种蛋白质。在这些鉴定的蛋白点中,以对照为参考胶,只在SA处理中上调表达的蛋白点有12个,下调表达的蛋白点2个,特异表达的蛋白点1个；只在SA+C.destructans处理中上调表达的蛋白点2个,下调表达的蛋白点1个；,在三个处理中都上调表达的蛋白点5个,下调表达的蛋白点1个。这些特异表达的蛋白涉及植物自身的防卫反应、信号转导、能量代谢和转录调控等方面。其中热激蛋白60、肉桂醇脱氢酶、单脱水抗坏血酸还原酶、甲氧基转移酶都与防御反应正相关。
6.首次建立了人参锈腐病菌分子检测体系,为人参锈腐病的早期诊断和病原菌种群动态监测奠定了理论基础。该体系在对人参锈腐病菌ITS区进行PCR扩增、测序及序列分析基础上,设计了特异性引物CD-F/CD-R,可以用于人参主要病原真菌及常见土壤习居菌的PCR检测。只有以C. destructans基因组DNA为模板的体系中能扩增出一条450bp左右的条带,而其他菌株及阴性对照均无特异性条带产生。利用该特异性引物能从罹病组织和人工接种C. destructans的土壤DNA中扩增出特异性条带,而对照及健康人参组织均无扩增产物,表明该特异性引物可以用于人参锈腐病菌的分子检测和早期诊断。
Ginseng (Panax ginseng C.A. Meyer) is a perennial herbaceous plant in the family Araliaceae which has been grown in northeast China for centuries, is known as the king of medicine. Cylindrocarpon root rot of ginseng caused by Cylindrocarpon destructans is the main disease which led to serious crop yield losses, and the average loss is up to20%-30%. The soilborne disease is very difficult to control. The use of high toxic pesticide is forbidden because of GAP cultivation. It maybe provide a new way to control Cylindrocarpon root rot of ginseng by using the principle of plant induced disease resistance to improve the resistance of ginseng plant and using molecular detection method to monitor pathogen population. The inducing mechanism of Cylindrocarpon root rot was studied and the pathogen molecular detection system was established based on plant induced disease resistance from the physiological and biochemical reaction, proteomics, and molecular detection. The results were as follows:
1. The effects of salicylic acid, cinnamic acid, ferulic acid, benzoic acid, p-hydroxybenzoic acid and methyl jasmonate on C. destructans were studied. These chemicals were added into media to incubate C. destructans to investigate their effects on the pathogen. The results indicated that the growth and spore germination of C. destructans were suppressed by cinnamic acid, ferulic acid, benzoic acid, while p-hydroxybenzoic acid had no influence on the pathogen. At low concentration of salicylic acid and methyl jasmonate, the growth of C. destructans was hardly affected, while at higher concentrations (>200μg mL-1) the growth of pathogen was prohibited.
2. The resistance of ginseng to C. destructans could be induced by low concentrations of SA and MeJA. Of these6kinds of chemicals, the induced effects of SA and MeJA were better than other chemicals. It showed that ginseng treated with low concentration of salicylic acid and jasmonic acid methyl jasmonate could enhance the resistance of ginseng plant itself, the control effects were up to67.9%and56.6%, respectively, and the two kinds of chemical factors in low concentration had promoting effects on the growth of ginseng. However, the other chemicals had harmful effects on ginseng growth. Therefore, SA and MeJA were chosen for next study.
3. The resistance of ginseng induced by SA and MeJA was firstly demonstrated. The physiological and biochemical mechanisms of ginseng resistant to Cylindrocarpon root rot induced by exogenous salicylic acid and methyl jasmonate were studied. SA and MeJA treatment could effectively decrease the contents of MDA and relative electricity conductivity, but increase the contents of proline, soluble sugar and total phenols. The activities of PAL, CAT, PPO, POD,(β-1,3-glucanase and chitinase could be increased significantly in the ginseng roots induced by SA and MeJA than those in control. The incidence and disease severity were reduced with ginseng plant resistance increased obviously due to exogenous applied SA and MeJA, and the incidences were reduced by39.1%and34.8%, respectively. It demonstrated that SA and MeJA could induce the system resistance of ginseng to C. destructans. In addition, the impact of SA was better than MeJA. Thus, SA is chosen as the chemical factor for further study.4. A stable and vigorous growth of ginseng suspension cell lines was established using plant tissue culture technology. The establishment of the system provides a stable and uniform test material in ginseng proteomics research. MS culture medium was used in callus induction, supplement with3mg/L2,4-D and0.2mg/L KT,0.8%sucrose, pH5.8to6.0. Successive transfer culture added2mg/L2,4-D and0.5mg/L BA. The hormone of suspension cell liquid medium was at the same levels of successive transfer culture. The culture condition of callus is25℃, dark culture. The suspension cell masses were grown in the flasks on a rotary shaker (110rpm)at24±2℃.
5. The proteome of salicylic acid inducing ginseng suspension cell was firstly studied by two-dimensional gel electrophoresis technology. A high-throughput and high-resolution two-dimensional gel electrophoresis system was established for suspension-cultured ginseng cells proteins. The proteins were extracted by using the phenol extraction, the elements of lysis buffer were composed of9M urea,2M Thiourea,2%IPG buffer,4%CHAPS,65mM DTT and1%TBP. The pH5-8IPG strip was used. SDS-PAGE was performed in12%polyacrylamide gels. The proteins related to resistance were identified.2-DE was adopted to separate the differentially expressed proteins from suspension-cultured ginseng cells induced by SA. About800protein spots were detected on the2-DE gels. Twenty-four differentially expressed proteins that changes in protein abundance in more than2times and good repeatability were analyzed by MALDI-TOF-MS, of23proteins were successfully identified. In the absence of pathogen,12proteins had increased levels, two proteins had decreased levels and one protein was newly induced in suspension cultures treated directly with SA, in comparison to the control. Two proteins were up-regulated and one protein was down-regulated after SA treatment in the presence of C. destructans. Five proteins were up-regulated and only one protein was down-regulated in suspension cultures were differentially expressed after both SA treatment and pathogen infection. These identified proteins were predicted to be involved in defense and stress responses, energy and metabolism, signal transduction/transcription, protein synthesis and metabolism, photosynthesis and others. The proteins related to defense responses, such as Chaperonin60, monodehydroascorbate reductase, CAD and orcinol O-methyltransferase, were more abundant in suspension cultures of ginseng roots after application of S A.
6. The molecular detection of Cylindrocarpon root rot of ginseng was established. A pair of primers specific for C. destructans was designed from comparisons of the internal transcribed spacer (ITS) regions1and2of isolates of diverse origins with sequences from commonly encountered fungi in soil. Under stringent PCR conditions, primers CD-F/CD-R amplify a450bp fragment from C. destructans DNA but not from other species or negative control. And C. destructans could be specifically detected with CD-F/CD-R from ginseng tissues and soil samples inoculated artificially.
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