烟草抗马铃薯γ病毒（PVY）育种、抗性基因的克隆与功能分析
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摘要
马铃薯Y病毒(Potato virus Y, PVY)是我国烟叶生产中的主要病害之一,近年来，随着农村保护地蔬菜栽培和马铃薯种植面积的扩大，烟草PVY病毒病的发生与危害呈现急剧上升趋势，在黑龙江、吉林、辽宁、山东、安徽和云南相继出现大面积爆发流行，造成了严重的经济损失。针对我国烟区缺乏抗PVY品种的生产实际，本研究通过筛选烟草PVY抗源，并对主要抗源材料进行抗性遗传分析，克隆其抗性基因，同时开展抗PVY烟草育种工作,主要包括以下内容:
1.采用人工接种方式，在温室和田间病圃中分别对17份和114份烟草种质进行PVY抗病性鉴定，筛选出高抗马铃薯Y病毒的烟草种质资源C151、CV91，达到中抗水平的有CV87，87414和NC55,表现多基因水平抗性。
2.对C151、CV91、CV87和NC55等抗源材料进行了抗性遗传分析，证实C151为少数基因控制，不完全显性，CV91为少数隐性基因控制，易受显性感病基因影响，与没有显性感病基因品种杂交表现抗性。CV87分别与IslandGold和龙江851杂交组合的F2代抗感比例不同且表现连续分布，证实为多数微效基因控制，且为加性基因控制。NC55×9625组合F2代中无病株-病轻株-病重株≈8-5-3，基本表现连续分布，NC55是多数微效抗性基因。
3.利用高抗PVY的CV91做亲本，选育出了烤烟常规纯系品种龙江911和杂种优势利用品种LJ237，其中龙江911为中抗马铃薯Y病毒病，LJ237高抗马铃薯Y病毒病，利用CV87育成了抗马铃薯Y病毒的烤烟品种LJ981。
4.提出微效水平抗性基因富集思路，利用中等水平抗性资源,选配适当的杂交组合，在杂交后代材料中富集、浓缩抗性基因，得到高于抗性亲本抗性的品种，重视水平抗性资源的开发与利用。
5.以高抗PVY的C151为试验材料，利用抑制差减杂交技术（Suppression SubtractiveHybridization，SSH）构建了抗PVY相关基因的烟草叶片SSH文库，对烟草叶片接种PVY前后的差异表达基因片段进行测序，得到280多条EST序列。利用NCBI上的GenBank等公共数据库对所获得的280多条EST序列进行生物信息分析和功能预测，其中与光合作用有关的RuBP羧化酶相关蛋白基因片段96条，林烟草和绒毛烟草叶绿体DNA片段12条，与抗病性有关的有：脂质转化酶基因片段15条，突变的过氧化氢酶基因片段8条，蛋白酶抑制剂基因片段32条，铁氧还蛋白和铁硫蛋白基因片段26条，糖基转移酶3条，富含甘氨酸蛋白质前体蛋白9条。其他序列86条，未知基因片段71条。
6、利用荧光定量PCR对5个可能与抗病性有关诱导表达片段的从开始接种到接种后72小时之间的0小时，12小时，24小时，36小时和72小时5个时间点基因的差异表达，并证实了所获得的基因序列为真实诱导的表达序列。对此序列进行RACE，最终获得一个与PVY抗病性有关的全长cDNA序列。
7、采用同源序列法，根据抗PVY的辣椒品种中的eIF4E基因序列设计扩增引物，在抗PVY的CV91、C151等烟草品种扩增晒选抗性基因，在CV91品种中克隆出了与马铃薯Y病毒抗性有关的翻译起始因子eIF4E基因。
8、通过比对该eIF4E基因的核酸序列，发现与感病品种有两个突变位点，导致病毒基因不能翻译，产生抗病性。
Potato virus Y (PVY) is one of the most important tobacco diseases in China, As the expand ofgreenhouse vegetable cultivation and the potato planting area in recently years, the occurrenceand damage of PVY to tobacco production have appeared sharp upward trend. A greateconomic loss was taken as the outbreak epidemic of PVY in Heilongjiang, Liaoning,Shandong, Anhui and Yunnan tobacco planting areas successively. In view of the actualsituation of lack of PVY resistance tobacco variety in our country, the PVY resistance sourceselection, genetic analysis for PVY resistance, cloning of PVY resistance gene and PVYresistance breeding have been done in this research, which contents including followings:
1.17and114tobacco germ plasma resources were identified for PVY-resistance usingthe artificial inoculation methods in greenhouse and field disease nursery respectively. Thetobacco varieties, C151and CV91, were selected out as high resistance to potato virus Y, andthe varieties, including CV87，87414and NC55were confirmed medium resistance to PVY,which were appeared to horizontal resistance controlled by multiple genes.
2. The PVY resistance varieties including C151、CV91、CV87and NC55were geneticanalyzed using hybrid with other materials, the results indicate that C151`PVY-resistance wascontrolled by few genes, which belong to incomplete dominant. CV91`PVY-resistance wascontrolled by few recessive gene，which is impressible of dominant gene susceptible to PVY，it will be appeared resistance hydride with varieties without dominant gene susceptible to PVY.F2generation of CV87Hybrid combination with IslandGold and Longjiang851respectivelyappeared difference in the rate, none disease: slight disease: heavy disease, which performanceof continuous distribution. Which controlled by the majority of minor genes，should be additivegene. The plants rate of none disease: slight disease: heavy disease in the F2generation ofNC55×9625combination≈8-5-3，which performance of continuous distribution. NC55wasshowed controlled by the majority of minor genes.
3. Using CV91as a parents, a high resistance to PVY variety, flue-cured tobacco varietiesLongjiang911and hybride LJ237were bred, and LJ237was confirmed as high resistance toPVY and longjiang911was confirmed as medium resistance PVY. In the same time, LJ981have been bred using CV87as a parents.
4. The idea concentrating more minor horizontal resistance genes in a variety was putforward in this article. Making use of medium horizontal resistance genes, setting properhybrid groups, concentrating more resistance genes and getting stronger resistance variety thanits parents is possible in their hybrid latter generations. The use of horizontal resistance germplasma resources should be appreciable.
5. C151, a variety high resistance to PVY, were used as the experimental materials, Thetobacco leaf SSH Library induced by PVY has been build usingSuppression Subtractive Hybridization, The differential expression fragments in the tobaccoleaf inoculated by PVY were sequenced more than280ESTs. The obtained280ESTs weresequence analyzed and function predicted through blasting in NCBI’s Blast Web site, amongof them,96fragments were predicted to be RuBP carboxylase related to photosynthesis,12fragments were believed to have relative with the chloroplast DNA in Nicotiana sylvestris andNicotiana tomentosiformis，and there are78fragments were predicted to be relate to PVY resistance,：15fragments for Lipid invertase gene,8fragments for mutant catalase gene，26fragments for protease inhibitor gene，26fragments for Ferredoxin gene and Iron sulfur proteingene,3fragments for Glycosyltransferase genes，9fragments for Glycine rich proteinprecursor protein,86fragments for other function genes。
6. Real-time PCR analysis was applied to analyze the dynamic expression patterns of fivegenes at five times after inoculation of PVY1-72hours. The results showed that all of that fivesequences were induced expression fragments, the full-length gene relative to PVY resistancewas obtained by the strategy of Rapid amplification of cDNA ends to the five sequences.
7. Using homologous sequence method, the translation initiation factor eIF4E which wasrelative to PVY resistance was cloned in tobacco variety CV91.
8. Through blast the eIF4E (above) gene sequence with others, two mutant sits have beenfound, which are not happened in susceptibility varieties, which possibly influences virustranslating and then cause the variety to occur resistance.
1.采用人工接种方式，在温室和田间病圃中分别对17份和114份烟草种质进行PVY抗病性鉴定，筛选出高抗马铃薯Y病毒的烟草种质资源C151、CV91，达到中抗水平的有CV87，87414和NC55,表现多基因水平抗性。
2.对C151、CV91、CV87和NC55等抗源材料进行了抗性遗传分析，证实C151为少数基因控制，不完全显性，CV91为少数隐性基因控制，易受显性感病基因影响，与没有显性感病基因品种杂交表现抗性。CV87分别与IslandGold和龙江851杂交组合的F2代抗感比例不同且表现连续分布，证实为多数微效基因控制，且为加性基因控制。NC55×9625组合F2代中无病株-病轻株-病重株≈8-5-3，基本表现连续分布，NC55是多数微效抗性基因。
3.利用高抗PVY的CV91做亲本，选育出了烤烟常规纯系品种龙江911和杂种优势利用品种LJ237，其中龙江911为中抗马铃薯Y病毒病，LJ237高抗马铃薯Y病毒病，利用CV87育成了抗马铃薯Y病毒的烤烟品种LJ981。
4.提出微效水平抗性基因富集思路，利用中等水平抗性资源,选配适当的杂交组合，在杂交后代材料中富集、浓缩抗性基因，得到高于抗性亲本抗性的品种，重视水平抗性资源的开发与利用。
5.以高抗PVY的C151为试验材料，利用抑制差减杂交技术（Suppression SubtractiveHybridization，SSH）构建了抗PVY相关基因的烟草叶片SSH文库，对烟草叶片接种PVY前后的差异表达基因片段进行测序，得到280多条EST序列。利用NCBI上的GenBank等公共数据库对所获得的280多条EST序列进行生物信息分析和功能预测，其中与光合作用有关的RuBP羧化酶相关蛋白基因片段96条，林烟草和绒毛烟草叶绿体DNA片段12条，与抗病性有关的有：脂质转化酶基因片段15条，突变的过氧化氢酶基因片段8条，蛋白酶抑制剂基因片段32条，铁氧还蛋白和铁硫蛋白基因片段26条，糖基转移酶3条，富含甘氨酸蛋白质前体蛋白9条。其他序列86条，未知基因片段71条。
6、利用荧光定量PCR对5个可能与抗病性有关诱导表达片段的从开始接种到接种后72小时之间的0小时，12小时，24小时，36小时和72小时5个时间点基因的差异表达，并证实了所获得的基因序列为真实诱导的表达序列。对此序列进行RACE，最终获得一个与PVY抗病性有关的全长cDNA序列。
7、采用同源序列法，根据抗PVY的辣椒品种中的eIF4E基因序列设计扩增引物，在抗PVY的CV91、C151等烟草品种扩增晒选抗性基因，在CV91品种中克隆出了与马铃薯Y病毒抗性有关的翻译起始因子eIF4E基因。
8、通过比对该eIF4E基因的核酸序列，发现与感病品种有两个突变位点，导致病毒基因不能翻译，产生抗病性。
Potato virus Y (PVY) is one of the most important tobacco diseases in China, As the expand ofgreenhouse vegetable cultivation and the potato planting area in recently years, the occurrenceand damage of PVY to tobacco production have appeared sharp upward trend. A greateconomic loss was taken as the outbreak epidemic of PVY in Heilongjiang, Liaoning,Shandong, Anhui and Yunnan tobacco planting areas successively. In view of the actualsituation of lack of PVY resistance tobacco variety in our country, the PVY resistance sourceselection, genetic analysis for PVY resistance, cloning of PVY resistance gene and PVYresistance breeding have been done in this research, which contents including followings:
1.17and114tobacco germ plasma resources were identified for PVY-resistance usingthe artificial inoculation methods in greenhouse and field disease nursery respectively. Thetobacco varieties, C151and CV91, were selected out as high resistance to potato virus Y, andthe varieties, including CV87，87414and NC55were confirmed medium resistance to PVY,which were appeared to horizontal resistance controlled by multiple genes.
2. The PVY resistance varieties including C151、CV91、CV87and NC55were geneticanalyzed using hybrid with other materials, the results indicate that C151`PVY-resistance wascontrolled by few genes, which belong to incomplete dominant. CV91`PVY-resistance wascontrolled by few recessive gene，which is impressible of dominant gene susceptible to PVY，it will be appeared resistance hydride with varieties without dominant gene susceptible to PVY.F2generation of CV87Hybrid combination with IslandGold and Longjiang851respectivelyappeared difference in the rate, none disease: slight disease: heavy disease, which performanceof continuous distribution. Which controlled by the majority of minor genes，should be additivegene. The plants rate of none disease: slight disease: heavy disease in the F2generation ofNC55×9625combination≈8-5-3，which performance of continuous distribution. NC55wasshowed controlled by the majority of minor genes.
3. Using CV91as a parents, a high resistance to PVY variety, flue-cured tobacco varietiesLongjiang911and hybride LJ237were bred, and LJ237was confirmed as high resistance toPVY and longjiang911was confirmed as medium resistance PVY. In the same time, LJ981have been bred using CV87as a parents.
4. The idea concentrating more minor horizontal resistance genes in a variety was putforward in this article. Making use of medium horizontal resistance genes, setting properhybrid groups, concentrating more resistance genes and getting stronger resistance variety thanits parents is possible in their hybrid latter generations. The use of horizontal resistance germplasma resources should be appreciable.
5. C151, a variety high resistance to PVY, were used as the experimental materials, Thetobacco leaf SSH Library induced by PVY has been build usingSuppression Subtractive Hybridization, The differential expression fragments in the tobaccoleaf inoculated by PVY were sequenced more than280ESTs. The obtained280ESTs weresequence analyzed and function predicted through blasting in NCBI’s Blast Web site, amongof them,96fragments were predicted to be RuBP carboxylase related to photosynthesis,12fragments were believed to have relative with the chloroplast DNA in Nicotiana sylvestris andNicotiana tomentosiformis，and there are78fragments were predicted to be relate to PVY resistance,：15fragments for Lipid invertase gene,8fragments for mutant catalase gene，26fragments for protease inhibitor gene，26fragments for Ferredoxin gene and Iron sulfur proteingene,3fragments for Glycosyltransferase genes，9fragments for Glycine rich proteinprecursor protein,86fragments for other function genes。
6. Real-time PCR analysis was applied to analyze the dynamic expression patterns of fivegenes at five times after inoculation of PVY1-72hours. The results showed that all of that fivesequences were induced expression fragments, the full-length gene relative to PVY resistancewas obtained by the strategy of Rapid amplification of cDNA ends to the five sequences.
7. Using homologous sequence method, the translation initiation factor eIF4E which wasrelative to PVY resistance was cloned in tobacco variety CV91.
8. Through blast the eIF4E (above) gene sequence with others, two mutant sits have beenfound, which are not happened in susceptibility varieties, which possibly influences virustranslating and then cause the variety to occur resistance.
引文
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