烟草花叶病毒弱毒疫苗的研制及其分子生物学
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摘要
烟草花叶病毒（Tobacco mosaic virus，TMV）是烟草花叶病毒属
（Tobamovirus）的代表种，给许多种植物造成严重损失。福建农林大学植物病
毒研究所对福建烟草病毒种群和田间发生频率的调查研究表明，TMV在福
建几个主要烟区占有很高的比例，其中以TMV普通株系为优势株系。利用
病毒的弱毒株的交互保护作用是防治病毒病较为理想的手段之一。为此本文
对TMV弱毒株的获得方法、生物学特性、分子生物学方面进行了较系统的
研究。
从福建烟区分离、纯化、鉴定到TMV普通株系。通过高温处理、亚硝
酸诱变及两者的复合处理对强毒株TMV-W进行诱变，筛选获得了两个弱毒
株，记为TMV-017（高温处理）、TMV-152（复合处理）。应用正交设计法考察
病毒接种浓度、接种间隔时间、接种部位以及普通烟生育期等因素对弱毒株
交互保护作用的影响，结果表明烟苗生育期和接种间隔时间两个因素影响最
大。弱毒株接种12d后才有较好的保护作用。浸根法可适用于弱毒株的大量
接种。
考察了弱毒株本身对烟草植株的影响。在普通烟上分别接种TMV-W、
TMV-017、TMV-152，考察3者在接种叶上的病毒的增殖情况以及叶绿素a、
叶绿素b、叶绿素、可溶性蛋白含量的变化。结果表明弱毒株的增殖速率、
在烟草植株体内的稳定浓度低于强毒株。分别受强、弱毒株侵染的烟草叶片
中的叶绿素a、叶绿素b、叶绿素含量均有所下降，但强毒株引起的下降幅
度最大。可溶性蛋白含量变化幅度不太明显。
观察了分别受TMV-W、TMV-017侵染的烟叶细胞病理的变化。健康烟
叶的叶绿体具双层膜，基粒片层紧密整齐，内含大量淀粉粒。受TMV-017
侵染的烟草细胞中，叶绿体数量较少，叶绿体具双层膜结构，基粒片层排列
整齐，有的叶绿体膨大，内含少量小的淀粉粒。叶绿体周围经常出现4-6个
线粒体。细胞质中出现结晶状内含体，经常存在于叶绿体周围。受TMV-W
浸染的烟草细胞中，有的叶绿体破裂，基粒松散。细胞质中的线粒体增多，
增大，有的形状不规则。细胞质中也出现晶体状内含体，分布在叶绿体周围。
测定了TMV-W、TMV-017、TMV-152的基因组cDNA全序列（GenBank
福建农林大学博士学位论文 ＊
接收号分别为：AF395127、AF395128、AF395129X 结果表明，强、弱毒株
基因组均为 6395个核昔酸，具有 4个开放阅读框（ORF），分别编码 126kDa
蛋白（69－3419nt，ill6ed和183kDa 通读蛋白（69－4919nt，1616eq、运动蛋白
（4903－5709m，268盼）、外壳蛋白（5712－619Im，159eq。5’端非编码区爬）
为68nt＊，通读蛋白和运动蛋白基因有17个碱基重叠，运动蛋白和外
壳蛋白基因间隔区为Znt（5710－57lint），3’端NR区为204nt（6192－6395nt）。
TMV－W和TMV－UI相比，基因组同源率达到98％，TMV．W为普通株系。
TMV，of 7、TMV．152基因组核苦酸发生变异的部位主要在126／183kDa蛋白
ORF。和 TMV．W相比，TMV．of 7仅在 126kDa蛋白 ORF中有 18个碱基发
生变异，m个引起氨基酸的变异；而TMVJ 在126门83kD蛋白OM中
有 16个碱基发生变异，其中有 11个引起氨基酸的变异。TMV．of 7和 TMV．
152有 11个共同变异的碱基，其中 8个引起氨基酸的变异。TMV－152在 MP
Ony中有1个碱基引起氨基酸的变异。其它区域，三者完全相同。
首次尝试将弱毒株的缺失复制酶基因转化烟草。构建了含TMV．W、
TMV－of 7各2个126kDa 蛋白O肚的部分片段，EIEZ（873－2813nt）、
RI EZ的928 3nt）的植物表达载体。经农杆菌 Ti质粒介导转化烟草。再生植
株总DNA的Southern点杂交检测结果初步表明，一个片段均己整合到烟草
基因组中。再生植株总RNA的RT－PCR Southern点杂交检测结果初步表明，
4个片段均有转录。用含TMV－W EIEZ的重组克隆和载体pGEX上，构建
了重组原核表达载体，诱导表达出融合蛋白O4kDal，制备了抗血清。免疫
斑点杂交检测抗血清的适宜工作浓度为 1：100。转化植株能延迟、减轻攻
击病毒oMV－W）的症状表现，含 TMV－W EI EZ片段的转化植株中有 1株抗
性水平高。
Tobacco mosaic virus(TMV), a type member of Tobamovirus, has caused great losses to many plants yields. The previous studies on the population and occurrence frequency of tobacco virus disease in Fujian in our institute(Institute of Plant Virology, Fujian Agriculture and Forestry University) suggested that the ratio of TMV was high in a few main tobacco fields and TMV common strain was dominant strain. Using the cross-protection of attenuated strain to protect plant from infecting by virulent strain of the same virus was proved one of ideal means. On the basis of these results, we studied the methods of obtaining the mild strains of TMV as well as their biology and molecular biology features.
TMV commom strain was isolated from Fujian tobacco region, then purified and identified. Two symptomless TMV mutants named TMV-O 17 and TMV- 152 respectively were obtained by treatment of heating, heating and nitrous acid combination. The effects on the effectiveness of TMV mild strain cross-protection, of the inoculation concentration of mild and virulent strains, inoculation interval time between mild and virulent strains, the inoculation position of the virulent strain and the age of tobacco, were observed by orthogonal designing. The inoculation interval time and the age of tobacco were the two most important factors. The cross-protection of mild strains were obvious when inoculation interval time between two strains over 12 days. The root soak was a method to mass inoculation of mild strains.
We examined the effects of mild strains of TMV on the tobacco plants. The rate of virus multiplication, the conents of chlorophyll a, chlorophyll b, total chlorophyll and soluble protein were detected, in the inoculated leaves of the tobacco plants infected by virulent and two mild strains of TMV respectively. The rates of replication and the steady concentrations of the two mild strains were less than those of the virulent strain. The contents of chlorophyll a, chlorophyll b, total chlorophyll all decreased in the leaves infected by the virulent and two mild strains respectively, but those of the virulent strain declined the most. The
contents of soluble protein all changed unconspicuously.
With electron microscope techniques, the cytopathological changes of the tobacco leaves infected by TMV-W and TMV-O 17 were observed. The results showed that the chloroplasts of health plant had double-membrance, and the stroma lamellae arranged tight and regular, containing a few of large starch granules. The quantity of chloroplasts was decrease of the leaves infected by TMV-O1 7. The chloroplasts had shaped membrance structure. The stroma lamellae also arranged tight and regular. Some chloroplasts enlarged, containing few little starch granules. The quantity of mitochondrias was increased to 4-6 around the chloroplasts. TMV-O 17 induced ciystal structure in cytoplasts, which appeared around the chloroplasts. Some chloroplasts of the leaves infected by TMV-W were destroyed, the stroma lainellae were caused to be loosed. The quantity of mitochondrias was also increased and enlarged, some even irregular. The rhombic crystal structure were also observed in cytoplasts around the chloroplasts.
The sequences of genomic cDNA of TMV-W, TMV-O 17 and TMV- 152 were detected(GenBank accession numbers are AF395127, AF395128, AF395129 respectively). Sequence analysis showed that the whole genomes of the three strains were 6395nt with 4 open reading frames(ORF) encoding for l26kDa protein(69-3419nt, with 11 l6aa), l83kDa protein(69-4919nt, with 1616aa), movement protein(4903-5709nt, with 268aa) and coat protein (5712-6191nt, with lS9aa) respectively. They possessed S抰erminal non-coding regions (NR) 68nt in length( 1 -68nt). There were 1 7nt overlap region between 1 83kDa protein ORE and movement protein ORE. Intercistron regions (IR) between movement protein ORE and coat protein ORE is 2nt. 3扤R has 204nt(6192-6395nt). The nucleotide sequence homology between the genome of TMV-W and TMV-U1 was 98%. TMV-W was proved common str
（Tobamovirus）的代表种，给许多种植物造成严重损失。福建农林大学植物病
毒研究所对福建烟草病毒种群和田间发生频率的调查研究表明，TMV在福
建几个主要烟区占有很高的比例，其中以TMV普通株系为优势株系。利用
病毒的弱毒株的交互保护作用是防治病毒病较为理想的手段之一。为此本文
对TMV弱毒株的获得方法、生物学特性、分子生物学方面进行了较系统的
研究。
从福建烟区分离、纯化、鉴定到TMV普通株系。通过高温处理、亚硝
酸诱变及两者的复合处理对强毒株TMV-W进行诱变，筛选获得了两个弱毒
株，记为TMV-017（高温处理）、TMV-152（复合处理）。应用正交设计法考察
病毒接种浓度、接种间隔时间、接种部位以及普通烟生育期等因素对弱毒株
交互保护作用的影响，结果表明烟苗生育期和接种间隔时间两个因素影响最
大。弱毒株接种12d后才有较好的保护作用。浸根法可适用于弱毒株的大量
接种。
考察了弱毒株本身对烟草植株的影响。在普通烟上分别接种TMV-W、
TMV-017、TMV-152，考察3者在接种叶上的病毒的增殖情况以及叶绿素a、
叶绿素b、叶绿素、可溶性蛋白含量的变化。结果表明弱毒株的增殖速率、
在烟草植株体内的稳定浓度低于强毒株。分别受强、弱毒株侵染的烟草叶片
中的叶绿素a、叶绿素b、叶绿素含量均有所下降，但强毒株引起的下降幅
度最大。可溶性蛋白含量变化幅度不太明显。
观察了分别受TMV-W、TMV-017侵染的烟叶细胞病理的变化。健康烟
叶的叶绿体具双层膜，基粒片层紧密整齐，内含大量淀粉粒。受TMV-017
侵染的烟草细胞中，叶绿体数量较少，叶绿体具双层膜结构，基粒片层排列
整齐，有的叶绿体膨大，内含少量小的淀粉粒。叶绿体周围经常出现4-6个
线粒体。细胞质中出现结晶状内含体，经常存在于叶绿体周围。受TMV-W
浸染的烟草细胞中，有的叶绿体破裂，基粒松散。细胞质中的线粒体增多，
增大，有的形状不规则。细胞质中也出现晶体状内含体，分布在叶绿体周围。
测定了TMV-W、TMV-017、TMV-152的基因组cDNA全序列（GenBank
福建农林大学博士学位论文 ＊
接收号分别为：AF395127、AF395128、AF395129X 结果表明，强、弱毒株
基因组均为 6395个核昔酸，具有 4个开放阅读框（ORF），分别编码 126kDa
蛋白（69－3419nt，ill6ed和183kDa 通读蛋白（69－4919nt，1616eq、运动蛋白
（4903－5709m，268盼）、外壳蛋白（5712－619Im，159eq。5’端非编码区爬）
为68nt＊，通读蛋白和运动蛋白基因有17个碱基重叠，运动蛋白和外
壳蛋白基因间隔区为Znt（5710－57lint），3’端NR区为204nt（6192－6395nt）。
TMV－W和TMV－UI相比，基因组同源率达到98％，TMV．W为普通株系。
TMV，of 7、TMV．152基因组核苦酸发生变异的部位主要在126／183kDa蛋白
ORF。和 TMV．W相比，TMV．of 7仅在 126kDa蛋白 ORF中有 18个碱基发
生变异，m个引起氨基酸的变异；而TMVJ 在126门83kD蛋白OM中
有 16个碱基发生变异，其中有 11个引起氨基酸的变异。TMV．of 7和 TMV．
152有 11个共同变异的碱基，其中 8个引起氨基酸的变异。TMV－152在 MP
Ony中有1个碱基引起氨基酸的变异。其它区域，三者完全相同。
首次尝试将弱毒株的缺失复制酶基因转化烟草。构建了含TMV．W、
TMV－of 7各2个126kDa 蛋白O肚的部分片段，EIEZ（873－2813nt）、
RI EZ的928 3nt）的植物表达载体。经农杆菌 Ti质粒介导转化烟草。再生植
株总DNA的Southern点杂交检测结果初步表明，一个片段均己整合到烟草
基因组中。再生植株总RNA的RT－PCR Southern点杂交检测结果初步表明，
4个片段均有转录。用含TMV－W EIEZ的重组克隆和载体pGEX上，构建
了重组原核表达载体，诱导表达出融合蛋白O4kDal，制备了抗血清。免疫
斑点杂交检测抗血清的适宜工作浓度为 1：100。转化植株能延迟、减轻攻
击病毒oMV－W）的症状表现，含 TMV－W EI EZ片段的转化植株中有 1株抗
性水平高。
Tobacco mosaic virus(TMV), a type member of Tobamovirus, has caused great losses to many plants yields. The previous studies on the population and occurrence frequency of tobacco virus disease in Fujian in our institute(Institute of Plant Virology, Fujian Agriculture and Forestry University) suggested that the ratio of TMV was high in a few main tobacco fields and TMV common strain was dominant strain. Using the cross-protection of attenuated strain to protect plant from infecting by virulent strain of the same virus was proved one of ideal means. On the basis of these results, we studied the methods of obtaining the mild strains of TMV as well as their biology and molecular biology features.
TMV commom strain was isolated from Fujian tobacco region, then purified and identified. Two symptomless TMV mutants named TMV-O 17 and TMV- 152 respectively were obtained by treatment of heating, heating and nitrous acid combination. The effects on the effectiveness of TMV mild strain cross-protection, of the inoculation concentration of mild and virulent strains, inoculation interval time between mild and virulent strains, the inoculation position of the virulent strain and the age of tobacco, were observed by orthogonal designing. The inoculation interval time and the age of tobacco were the two most important factors. The cross-protection of mild strains were obvious when inoculation interval time between two strains over 12 days. The root soak was a method to mass inoculation of mild strains.
We examined the effects of mild strains of TMV on the tobacco plants. The rate of virus multiplication, the conents of chlorophyll a, chlorophyll b, total chlorophyll and soluble protein were detected, in the inoculated leaves of the tobacco plants infected by virulent and two mild strains of TMV respectively. The rates of replication and the steady concentrations of the two mild strains were less than those of the virulent strain. The contents of chlorophyll a, chlorophyll b, total chlorophyll all decreased in the leaves infected by the virulent and two mild strains respectively, but those of the virulent strain declined the most. The
contents of soluble protein all changed unconspicuously.
With electron microscope techniques, the cytopathological changes of the tobacco leaves infected by TMV-W and TMV-O 17 were observed. The results showed that the chloroplasts of health plant had double-membrance, and the stroma lamellae arranged tight and regular, containing a few of large starch granules. The quantity of chloroplasts was decrease of the leaves infected by TMV-O1 7. The chloroplasts had shaped membrance structure. The stroma lamellae also arranged tight and regular. Some chloroplasts enlarged, containing few little starch granules. The quantity of mitochondrias was increased to 4-6 around the chloroplasts. TMV-O 17 induced ciystal structure in cytoplasts, which appeared around the chloroplasts. Some chloroplasts of the leaves infected by TMV-W were destroyed, the stroma lainellae were caused to be loosed. The quantity of mitochondrias was also increased and enlarged, some even irregular. The rhombic crystal structure were also observed in cytoplasts around the chloroplasts.
The sequences of genomic cDNA of TMV-W, TMV-O 17 and TMV- 152 were detected(GenBank accession numbers are AF395127, AF395128, AF395129 respectively). Sequence analysis showed that the whole genomes of the three strains were 6395nt with 4 open reading frames(ORF) encoding for l26kDa protein(69-3419nt, with 11 l6aa), l83kDa protein(69-4919nt, with 1616aa), movement protein(4903-5709nt, with 268aa) and coat protein (5712-6191nt, with lS9aa) respectively. They possessed S抰erminal non-coding regions (NR) 68nt in length( 1 -68nt). There were 1 7nt overlap region between 1 83kDa protein ORE and movement protein ORE. Intercistron regions (IR) between movement protein ORE and coat protein ORE is 2nt. 3扤R has 204nt(6192-6395nt). The nucleotide sequence homology between the genome of TMV-W and TMV-U1 was 98%. TMV-W was proved common str
引文
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