中国辣椒叶斑病病原学及分子检测研究
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摘要
辣椒叶斑病(leaf spot of pepper)是我国新报道发生病害,在保护地辣椒生产中,病害发生面积及其危害呈上升趋势。目前对于该病的发生危害、病原学、致病机理、病害发生规律乃至分子检测等均缺乏系统的研究,制约着病害防治策略的制定及病害的防控。本文针对这些问题,以辣椒叶斑病为研究对象,对其致病病原进行了鉴定,从病菌生物学特性、致病力分化、种群遗传多样性和侵染特性4个方面进行了病原学的系统研究；对其病原菌致病机理进行了初步探讨；并建立了辣椒叶斑病菌的分子检测技术,以期为病害的早期诊断提供有效方法。
1.首次鉴定明确了我国辣椒叶斑病的致病菌
2009年6月在辽宁省大连瓦房店市首次发现辣椒叶斑病。2009年至2012年,全面调查了辽宁省各辣椒栽培地区辣椒叶斑病的发生危害情况。通过组织分离和显微形态鉴定,分离获得病原菌22株。经常规的形态学观察和柯赫氏法则证病试验,并采用分子技术——多基因联合分析方法,对病菌的rDNA-ITS、β-tubulin和Actin基因序列进行比对,鉴定明确了我国辣椒叶斑病的致病菌为尖孢枝孢Cladosporium oxysporum。
2.系统测定了辣椒叶斑病致病菌尖孢枝孢的生物学特性
通过采用不同的营养、温度、pH值、光照等条件,对病菌菌丝生长及孢子萌发影响因素测定试验结果表明,对于病菌菌丝生长,以V8汁培养基为最适；能有效利用多种糖和氮源,分别以山梨醇和麦芽浸膏最佳；适宜温度为20～25℃；最适pH值为6～8；光照可促进菌丝生长；菌丝致死温度为77℃。而对于病菌分生孢子萌发,对碳源无特殊需要,在清水中萌发率最高；氮源培养液严重抑制孢子萌发；20～30℃为孢子萌发适宜温度；最适pH值为7；光照对孢子萌发无明显的促进作用；分生孢子致死温度为48℃。
3.测定探讨了辽宁省不同地理来源的辣椒叶斑病菌的致病力
采用人工接种方法,选用4个辣椒栽培品种(迅驰、沈椒4号、杭椒一号和新金号角),对采自辽宁省不同地区的辣椒叶斑病菌22个菌株,进行了致病力测定。根据供试菌株对4个供试辣椒栽培品种的抗感反应,将22个菌株划分为3个致病类型,即强致病类型、中等致病类型和弱致病类型。表明我国辣椒叶斑病菌存在明显的致病力分化现象,但地理来源与致病力分化之间无明显的相关性,相同地理来源的病原菌群体中致病力存在强、中、弱的差异。
4.测定分析了辣椒叶斑病菌的遗传多样性
应用ISSR和SRAP两种分子标记技术,对供试辣椒叶斑病菌22个菌株进行了遗传多样性分析,依据不同地理来源尖孢枝孢的ISSR和SRAP指纹图谱分别构建UPGMA系统发育树,从分子水平探究致病菌的遗传演变和进化。测定结果表明,ISSR标记的菌株间相似系数在0.56～0.93之间,在相似系数0.65处被划分成4个类群；SRAP标记的菌株间相似系数在0.59～0.90之间,在相似系数约0.63处被划分为3个类群。表明供试辣椒叶斑病菌群体具有丰富的遗传多样性和明显的遗传分化现象,遗传聚类组群与菌株的地理来源具有一定的相关性,而与菌株的致病性无明显的相关性。
5.初步探讨了辣椒叶斑病的侵染循环
试验研究结果表明,辣椒叶斑病菌尖孢枝孢以菌丝体在植株残体上潜伏,随病残体残落于土壤越冬,作为病害翌年危害的初侵染源。温度和湿度是影响病害发生的最主要因素,病菌侵染植株的最适温度为20～25℃,潜育期为7～8d；在适温条件下,保湿时间越长,病害发病率越高。罹病辣椒叶片的显微及超微观察结果显示,病菌以菌丝通过表皮和气孔2种途径侵入寄主组织；接种后8h,即可观察到分生孢子萌发产生芽管,进而形成菌丝,在叶片组织中穿行扩展；老熟后的菌丝上产生分生孢子梗,其上产生大量产生分生孢子；分生孢子成熟后自然脱落,可经多次再侵染反复循环传播；病菌菌丝在寄主组织中不断扩展,导致受侵细胞被破坏,最终崩解。
6.初步研究了辣椒叶斑病菌的致病机理
辣椒叶斑病菌在人工培养条件下可以产生毒素,对辣椒叶片具有明显的毒害作用,引致与田间症状类似的病斑,而且可导致幼苗萎蔫,抑制种子萌发和胚根生长。病菌最佳产毒条件为采用pH值为7的Richard培养液,置于25℃黑暗条件下,持续振荡培养14d。活体内外辣椒叶斑病菌产生的细胞壁降解酶活性测定结果表明,病菌能产生果胶酶(PG、PMG、PGTE、PMTE)和纤维素酶(Cx、β-葡萄糖苷酶),人工培养条件下,果胶酶活性明显高于纤维素酶,是活体外病菌产生的主要细胞壁降解酶；辣椒叶片接种病菌后,果胶酶活性显著提高,在接种3～4d后达到活性高峰,推测其在病菌致病过程中先起作用,而纤维素酶活性逐渐增强,推测其参与病菌在寄主组织内的扩展。
7.初步建立了辣椒叶斑病菌巢式PCR分子检测体系
根据尖孢枝孢rDNA-ITS区序列,设计了特异性引物Clad-F/Clad-R,在常规PCR反应的基础上,以Clad-F/Clad-R为外引物,设计合成了内引物Clad-NF/Clad-NR,建立了辣椒叶斑病菌巢式PCR分子检测体系。分子检测结果表明,人工接种发病情况下,该方法对病原菌的最低检测量为1fgDNA/μL。应用巢氏PCR方法对人工接种辣椒叶斑病菌处于不同潜育期的辣椒叶片进行检测,结果表明,接种后第二天的叶片即可检测到病原菌。在辣椒生产田中,按五点取样法对自然侵染的辣椒叶片在显症前进行随机取样,巢式PCR检测病叶带菌率为52%,离体叶片培养调查发病率为32%。本研究初步建立的分子检测方法,具有比传统方法更高的灵敏度,可用以在病害潜育期即未显症期对其进行早期诊断。
The newly occurred pepper disease, pepper leaf spot, observed in greenhouse of Liaoning Province has become more and more serious in China in recent years. Because limited knowledgement is known on the etiology, pathogenic mechanism, occurrence regularity and molecular detection of the disease, the effective control strategies have not been established by now. In order to solve these problems, firstly the pathogen was identified in this study. In the aspect of etiology research, biological characteristics, pathogenicity, genetic diversity and infection characteristics were studied systematicly. Then a preliminary study on pathogenic mechanism of the pathogen was carried on. Besides, a molecular technigue for early diagnosis was established.
1. The First Identification of Pepper Leaf Spot Pathogen in China
In June2009, for the first time, leaf spotting was observed on pepper leaves in Wafangdian County of Liaoning Province. During2009to2012, a comprehensive survey of the occurrence and damage of pepper leaf spot was conducted in pepper cultivation areas of Liaoning Province. By tissue isolation and morphological identification,22strains of the pathogenic were collected. The pathogen was identified as Cladosporium oxysporum according to morphology and Koch's postulates testing, in combination with rDNA-ITS, β-tubulin, Actin genes sequences data sets.
2. Systematic Determination of Biological Characteristics of C. oxysporum
The results of effect factors on mycelium growth and conidium germination of the pathogen showed that the best medium for mycelium growth was V8juice. The pathogen could effectively use of sugar and nitrogen sources. Sorbitol and malt extract were used as better carbon and nitrogen sources for mycelium growth, respectively. The optimum temperature for mycelium growth was20to25℃and the optimum pH was6to8. Light could promote mycelium growth. The mycelium lethal temperature was77℃. Water was best for conidium germination without special carbon sources. But nitrogen sources seriously inhibited conidium germination. The optimum temperature for conidium germination was20to30℃and the optimum pH was7. Light had no obvious promoting effect on conidium germination. The conidium lethal temperature was48℃.
3. Determination of the Pathogenicity of C. oxysporum from Different Geographic Origins in Liaoning Province
Pathogenicity was determined of C. oxysporum from different geographic origins in Liaoning Province on four pepper cultivars (Xunchi,4th Shen pepper,1st Hang pepper and New golden horn) by artifical inoculation. The results showed22strains could be classified into three pathogenic types, which were strong pathogenic type, moderate pathogenic type and weak pathogenic type. The C. oxysporum strains showed significant difference in pathogenicity, while there was no significant correlation between the pathogenic types of strains and their geographic origins. The strains from the same geographic origin also exhibited different levels of pathogenicity.
4. Aanlysis of the Genetic Diversity of C. oxysporum
In order to explore heredity variation and evolution of the pathogen on molecular level, genetic diversity of all C. oxysporum strains was analyzed by ISSR and SRAP technique. The phylogenetic tree based on ISSR markers revealed that similarity coefficient of22strains was from0.56to0.93, and the strains were divided into four clusters at the threshold of genetic similar coefficient0.65by UPGMA. The phylogenetic tree based on SRAP markers revealed that similarity coefficient of22strains was from0.59to0.90, and the strains were divided into three clusters at the threshold of genetic similar coefficient approximately0.63by UPGMA. Both ISSR and SRAP analysis showed that there were abundant genetic diversity and remarkable genetic differentiation among C. oxysporum strains. The genetic clusters of C. oxysporum strains were associated with their geographic origins, but there was no significant correlation between the genetic relationships and pathogenicity of the strains.
5. Preliminary Study on the Infection Cycle of Pepper Leaf Spot
The results showed that C. oxysporum adhering to plant residues could overwinter successfully and become primary infection sources of the disease. Temperature and humidity were two main factors on disease occurring. The optimum temperature for infection was20to25℃when incubation period was7to8days. Under optimum temperature, keeping moist increased incidence of the disease. The infection process of C. oxysporum on pepper leaves was studied micrologically and ultramicrologically. It showed that the pathogen was able to penetrate host through two paths such as epidermis and stomata. The infectious stage started with conidium germination8h after inoculation, subsequently germinated hyphae grew over the tissue and conidiophores emerged through the stomata which produced conidia. Conidia fell off naturally after matured, then repeated infection. A large number of mycelia expanded rapidly within leaf tissue, and infected cells were damaged and died finally.
6. Preliminary Study on the Pathogenic Mechanism of C. oxysporum
The pathogen could produce toxin in fluid media. The biological activity assay of the crude toxin showed that it could induce the characteristic symptom of pepper leaf spot, cause seeding wilting and inhibit seed germination and radicle growth. The optimum conditions of producing toxin were Richard culture filtrate with pH7and sustained oscillation culture for14d in dark at25℃. A series of cell wall degrading enzymes (CWDEs) including pectinase (PG, PMG, PGTE and PMTE) and cellulase (Cx and β-glucosidase) were found in the pathogen. Pectinase produced from the pathogen showed higher activities as compared with cellulase in vitro CWDEs. Activities of pectinase were significantly improved after inoculation and reached the peak in3-4d after inoculation. Perhaps pectinase firstly play a role in pathogenic process. Activities of cellulase were gradually increased after inoculation. Maybe cellulase was conducive to the expansion of pathogen in the host tissues.
7. Establishment of the Nested PCR Detection System of C。 oxysporum
Based on the rDNA-ITS conservative sequence of C. oxysporum, one pair of specific primers Clad-F/Clad-R was designed. Then intermal primers Clad-NF/Clad-NR were designed based on the regular PCR product amplified with the primers Clad-F/Clad-R as outer primers. Combined these two kinds of primers, a nested PCR protocol was established. The sensitivity of this assay was1fg DNA of C. oxysporum. The nested PCR detection results in artificially inoculated pepper leaves with different incubation period showed that pathogen could be detected on the second day after inoculation. For symptomless samples that randomly collected from fields, the efficiency of the nested PCR assay was compared to that of culture in vitro method. As a result, the detection rate of nested PCR was52%, while incidence of leaves culture in vitro was32%. These data showed that the nested PCR had higher sensitivity than traditional method. The nested PCR could be effective to diagnose disease in incubation period when pepper leaves without visible symptoms in field.
1.首次鉴定明确了我国辣椒叶斑病的致病菌
2009年6月在辽宁省大连瓦房店市首次发现辣椒叶斑病。2009年至2012年,全面调查了辽宁省各辣椒栽培地区辣椒叶斑病的发生危害情况。通过组织分离和显微形态鉴定,分离获得病原菌22株。经常规的形态学观察和柯赫氏法则证病试验,并采用分子技术——多基因联合分析方法,对病菌的rDNA-ITS、β-tubulin和Actin基因序列进行比对,鉴定明确了我国辣椒叶斑病的致病菌为尖孢枝孢Cladosporium oxysporum。
2.系统测定了辣椒叶斑病致病菌尖孢枝孢的生物学特性
通过采用不同的营养、温度、pH值、光照等条件,对病菌菌丝生长及孢子萌发影响因素测定试验结果表明,对于病菌菌丝生长,以V8汁培养基为最适；能有效利用多种糖和氮源,分别以山梨醇和麦芽浸膏最佳；适宜温度为20～25℃；最适pH值为6～8；光照可促进菌丝生长；菌丝致死温度为77℃。而对于病菌分生孢子萌发,对碳源无特殊需要,在清水中萌发率最高；氮源培养液严重抑制孢子萌发；20～30℃为孢子萌发适宜温度；最适pH值为7；光照对孢子萌发无明显的促进作用；分生孢子致死温度为48℃。
3.测定探讨了辽宁省不同地理来源的辣椒叶斑病菌的致病力
采用人工接种方法,选用4个辣椒栽培品种(迅驰、沈椒4号、杭椒一号和新金号角),对采自辽宁省不同地区的辣椒叶斑病菌22个菌株,进行了致病力测定。根据供试菌株对4个供试辣椒栽培品种的抗感反应,将22个菌株划分为3个致病类型,即强致病类型、中等致病类型和弱致病类型。表明我国辣椒叶斑病菌存在明显的致病力分化现象,但地理来源与致病力分化之间无明显的相关性,相同地理来源的病原菌群体中致病力存在强、中、弱的差异。
4.测定分析了辣椒叶斑病菌的遗传多样性
应用ISSR和SRAP两种分子标记技术,对供试辣椒叶斑病菌22个菌株进行了遗传多样性分析,依据不同地理来源尖孢枝孢的ISSR和SRAP指纹图谱分别构建UPGMA系统发育树,从分子水平探究致病菌的遗传演变和进化。测定结果表明,ISSR标记的菌株间相似系数在0.56～0.93之间,在相似系数0.65处被划分成4个类群；SRAP标记的菌株间相似系数在0.59～0.90之间,在相似系数约0.63处被划分为3个类群。表明供试辣椒叶斑病菌群体具有丰富的遗传多样性和明显的遗传分化现象,遗传聚类组群与菌株的地理来源具有一定的相关性,而与菌株的致病性无明显的相关性。
5.初步探讨了辣椒叶斑病的侵染循环
试验研究结果表明,辣椒叶斑病菌尖孢枝孢以菌丝体在植株残体上潜伏,随病残体残落于土壤越冬,作为病害翌年危害的初侵染源。温度和湿度是影响病害发生的最主要因素,病菌侵染植株的最适温度为20～25℃,潜育期为7～8d；在适温条件下,保湿时间越长,病害发病率越高。罹病辣椒叶片的显微及超微观察结果显示,病菌以菌丝通过表皮和气孔2种途径侵入寄主组织；接种后8h,即可观察到分生孢子萌发产生芽管,进而形成菌丝,在叶片组织中穿行扩展；老熟后的菌丝上产生分生孢子梗,其上产生大量产生分生孢子；分生孢子成熟后自然脱落,可经多次再侵染反复循环传播；病菌菌丝在寄主组织中不断扩展,导致受侵细胞被破坏,最终崩解。
6.初步研究了辣椒叶斑病菌的致病机理
辣椒叶斑病菌在人工培养条件下可以产生毒素,对辣椒叶片具有明显的毒害作用,引致与田间症状类似的病斑,而且可导致幼苗萎蔫,抑制种子萌发和胚根生长。病菌最佳产毒条件为采用pH值为7的Richard培养液,置于25℃黑暗条件下,持续振荡培养14d。活体内外辣椒叶斑病菌产生的细胞壁降解酶活性测定结果表明,病菌能产生果胶酶(PG、PMG、PGTE、PMTE)和纤维素酶(Cx、β-葡萄糖苷酶),人工培养条件下,果胶酶活性明显高于纤维素酶,是活体外病菌产生的主要细胞壁降解酶；辣椒叶片接种病菌后,果胶酶活性显著提高,在接种3～4d后达到活性高峰,推测其在病菌致病过程中先起作用,而纤维素酶活性逐渐增强,推测其参与病菌在寄主组织内的扩展。
7.初步建立了辣椒叶斑病菌巢式PCR分子检测体系
根据尖孢枝孢rDNA-ITS区序列,设计了特异性引物Clad-F/Clad-R,在常规PCR反应的基础上,以Clad-F/Clad-R为外引物,设计合成了内引物Clad-NF/Clad-NR,建立了辣椒叶斑病菌巢式PCR分子检测体系。分子检测结果表明,人工接种发病情况下,该方法对病原菌的最低检测量为1fgDNA/μL。应用巢氏PCR方法对人工接种辣椒叶斑病菌处于不同潜育期的辣椒叶片进行检测,结果表明,接种后第二天的叶片即可检测到病原菌。在辣椒生产田中,按五点取样法对自然侵染的辣椒叶片在显症前进行随机取样,巢式PCR检测病叶带菌率为52%,离体叶片培养调查发病率为32%。本研究初步建立的分子检测方法,具有比传统方法更高的灵敏度,可用以在病害潜育期即未显症期对其进行早期诊断。
The newly occurred pepper disease, pepper leaf spot, observed in greenhouse of Liaoning Province has become more and more serious in China in recent years. Because limited knowledgement is known on the etiology, pathogenic mechanism, occurrence regularity and molecular detection of the disease, the effective control strategies have not been established by now. In order to solve these problems, firstly the pathogen was identified in this study. In the aspect of etiology research, biological characteristics, pathogenicity, genetic diversity and infection characteristics were studied systematicly. Then a preliminary study on pathogenic mechanism of the pathogen was carried on. Besides, a molecular technigue for early diagnosis was established.
1. The First Identification of Pepper Leaf Spot Pathogen in China
In June2009, for the first time, leaf spotting was observed on pepper leaves in Wafangdian County of Liaoning Province. During2009to2012, a comprehensive survey of the occurrence and damage of pepper leaf spot was conducted in pepper cultivation areas of Liaoning Province. By tissue isolation and morphological identification,22strains of the pathogenic were collected. The pathogen was identified as Cladosporium oxysporum according to morphology and Koch's postulates testing, in combination with rDNA-ITS, β-tubulin, Actin genes sequences data sets.
2. Systematic Determination of Biological Characteristics of C. oxysporum
The results of effect factors on mycelium growth and conidium germination of the pathogen showed that the best medium for mycelium growth was V8juice. The pathogen could effectively use of sugar and nitrogen sources. Sorbitol and malt extract were used as better carbon and nitrogen sources for mycelium growth, respectively. The optimum temperature for mycelium growth was20to25℃and the optimum pH was6to8. Light could promote mycelium growth. The mycelium lethal temperature was77℃. Water was best for conidium germination without special carbon sources. But nitrogen sources seriously inhibited conidium germination. The optimum temperature for conidium germination was20to30℃and the optimum pH was7. Light had no obvious promoting effect on conidium germination. The conidium lethal temperature was48℃.
3. Determination of the Pathogenicity of C. oxysporum from Different Geographic Origins in Liaoning Province
Pathogenicity was determined of C. oxysporum from different geographic origins in Liaoning Province on four pepper cultivars (Xunchi,4th Shen pepper,1st Hang pepper and New golden horn) by artifical inoculation. The results showed22strains could be classified into three pathogenic types, which were strong pathogenic type, moderate pathogenic type and weak pathogenic type. The C. oxysporum strains showed significant difference in pathogenicity, while there was no significant correlation between the pathogenic types of strains and their geographic origins. The strains from the same geographic origin also exhibited different levels of pathogenicity.
4. Aanlysis of the Genetic Diversity of C. oxysporum
In order to explore heredity variation and evolution of the pathogen on molecular level, genetic diversity of all C. oxysporum strains was analyzed by ISSR and SRAP technique. The phylogenetic tree based on ISSR markers revealed that similarity coefficient of22strains was from0.56to0.93, and the strains were divided into four clusters at the threshold of genetic similar coefficient0.65by UPGMA. The phylogenetic tree based on SRAP markers revealed that similarity coefficient of22strains was from0.59to0.90, and the strains were divided into three clusters at the threshold of genetic similar coefficient approximately0.63by UPGMA. Both ISSR and SRAP analysis showed that there were abundant genetic diversity and remarkable genetic differentiation among C. oxysporum strains. The genetic clusters of C. oxysporum strains were associated with their geographic origins, but there was no significant correlation between the genetic relationships and pathogenicity of the strains.
5. Preliminary Study on the Infection Cycle of Pepper Leaf Spot
The results showed that C. oxysporum adhering to plant residues could overwinter successfully and become primary infection sources of the disease. Temperature and humidity were two main factors on disease occurring. The optimum temperature for infection was20to25℃when incubation period was7to8days. Under optimum temperature, keeping moist increased incidence of the disease. The infection process of C. oxysporum on pepper leaves was studied micrologically and ultramicrologically. It showed that the pathogen was able to penetrate host through two paths such as epidermis and stomata. The infectious stage started with conidium germination8h after inoculation, subsequently germinated hyphae grew over the tissue and conidiophores emerged through the stomata which produced conidia. Conidia fell off naturally after matured, then repeated infection. A large number of mycelia expanded rapidly within leaf tissue, and infected cells were damaged and died finally.
6. Preliminary Study on the Pathogenic Mechanism of C. oxysporum
The pathogen could produce toxin in fluid media. The biological activity assay of the crude toxin showed that it could induce the characteristic symptom of pepper leaf spot, cause seeding wilting and inhibit seed germination and radicle growth. The optimum conditions of producing toxin were Richard culture filtrate with pH7and sustained oscillation culture for14d in dark at25℃. A series of cell wall degrading enzymes (CWDEs) including pectinase (PG, PMG, PGTE and PMTE) and cellulase (Cx and β-glucosidase) were found in the pathogen. Pectinase produced from the pathogen showed higher activities as compared with cellulase in vitro CWDEs. Activities of pectinase were significantly improved after inoculation and reached the peak in3-4d after inoculation. Perhaps pectinase firstly play a role in pathogenic process. Activities of cellulase were gradually increased after inoculation. Maybe cellulase was conducive to the expansion of pathogen in the host tissues.
7. Establishment of the Nested PCR Detection System of C。 oxysporum
Based on the rDNA-ITS conservative sequence of C. oxysporum, one pair of specific primers Clad-F/Clad-R was designed. Then intermal primers Clad-NF/Clad-NR were designed based on the regular PCR product amplified with the primers Clad-F/Clad-R as outer primers. Combined these two kinds of primers, a nested PCR protocol was established. The sensitivity of this assay was1fg DNA of C. oxysporum. The nested PCR detection results in artificially inoculated pepper leaves with different incubation period showed that pathogen could be detected on the second day after inoculation. For symptomless samples that randomly collected from fields, the efficiency of the nested PCR assay was compared to that of culture in vitro method. As a result, the detection rate of nested PCR was52%, while incidence of leaves culture in vitro was32%. These data showed that the nested PCR had higher sensitivity than traditional method. The nested PCR could be effective to diagnose disease in incubation period when pepper leaves without visible symptoms in field.
引文
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