大豆对SMV3号株系的抗性遗传及分子标记研究
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摘要
大豆花叶病毒（SMV）病是世界性大豆病害，在我国各大豆产区普遍发生，对大
豆生产危害严重，利用抗病品种是控制SMV最经济而有效的措施。东北是我国大豆主
产区，SMV3号株系是强毒株系，抗源较少。筛选鉴定和研究新抗源具有重要的理论价
值和实践意义。本研究利用SMV3号株系对大豆资源进行了抗性鉴定，筛选出抗性种
质，并利用农艺性状和RAPD标记分析了部分抗源的遗传关系。通过对抗病×感病杂
交组合后代分析，明确了高抗SMV的大豆品系95-5383的抗性遗传规律。利用BSA法
筛选出与SMV抗性基因紧密连锁的RAPD标记，并将其转化成SCAR标记。利用作图
群体将RAPD标记定位于大豆遗传连锁图上。具体研究结果如下：
1．利用人工汁液摩擦法对不同来源的347份大豆种质资源接种SMV3号株系进行
抗性鉴定。筛选出113份高抗资源，占32.56％；112份材料中抗SMV3，占32.27％；122
份感病，占35.16％。抗源主要来自于东北春作大豆区（辽宁）和黄淮海夏作大豆区（山
东、山西、北京）以及美国和韩国，南方大豆产区抗源较少。不同品种接种后的症状类
型不同，表明品种和株系间存在互作。国外引进资源接种后顶枯症状较多，表明症状反
应和地理来源有一定关系。本文还分析了美国一些抗源对SMV3的抗性反应及抗性基
因的关系。
2．利用筛选出的高抗SMV3号株系的大豆品系95-5383与4个感病品种（品系）
HBI、铁丰21、Amsoy、Williams和抗病品种PI486355配制5个杂交组合，对各组合
的F1、F2代接种SMV鉴定抗性。结果表明，95-5383与各感病品种杂交组合的F1代
表现为感病，F2群体分离比例为3感（花叶+顶枯）:1抗，表明95-5383对SMV3号
株系的抗性受一对隐性基因控制。95-5383×PI486355的F2代接种后有感病植株分离，
表明二者对SMV3的抗性基因是不等位的。利用BSA法对95-5383×HBI的F2代进行
鉴定，筛选出与SMV抗病基因紧密连锁的共显性RAPD标记OPNII_(980/1070)。RAPD引
物OPNII在95-5383和抗池扩增出OPNII_(980)片段，在HBI和感池扩增出OPNII_(1070)片
段，在F1同时扩增出OPNII_(980)和OPNII_(1070)。用该引物分析95-5383×HBI的F2个体，
OPNII_(980/1070)与95-5383抗病基因的遗传距离为2.1cM。
3．从95-5383和HBI中分别回收OPNII_(980)和OPNII_(1070)特异片段，克隆在PGEM-
T easy vector中，并进行序列分析。结果表明OPNII_(980)和OPNII_(1070)的实际长度分别
为986bp和1084bp，同源性为93.4％，主要差别是在两个不同区段插入（缺失）54bp
和35bp的碱基。用克隆的RAPD片段OPNII_(980)作探针（SOPNII）对亲本基因组DNA
中国农业科学陨博十学位论义
进行 Southern 65交，结果表明 OPN。。和 OPN川。。在人豆基因组中是以低拷贝形式
存在。用n PD引物OPN和y卜于探针9口PN！1分析科丰1。南农1！3吕毛重组自交
系作图群体，将 Opp和 SOPN定位于 F迹锁群的抗病基冈簇上．根据克隆片段
OPN。羽IOPNll；。。的序列设计合成了一对SCAR引物SCNll，SCNll在95·5383中
扩增出与 OPN。人小相同的片段 SCN。，在 HB中扩增出与 OPN。大小相同
的片段SCN。，。，在FI同时扩增出SCN！l＿、。SCNn对95·5383 X HBI的FZ群体
扩增结果与RAPD引物OPNll分析结果完全吻台。SCAR标记SCNll。。，。对于大豆
抗SMV标记辅助选抒育种和抗病性鉴定有很）“泛的应用前景。
4．选用已经鉴定出的68份高抗SMV3的大亘资源，利用农艺性状和nPD分析
品种间的遗传关系。从 60个 nPD引物中筛选出 11个多态性引物，共扩增出刃个片
段，多态性片段为33条。扩增产物的有无分别以1和0记录，选用J＿。rd公式计算
品种之间的相似系数，利用UPGMA法构建了树状图。RAPD聚类结果和农艺性状聚
类结果具有一定的吻合性。聚类结果表明亲源关系相同、地理来额相同的品种多数聚在
一类。分聚在不同组间的人豆品种迢传背景存在差异，可能具有不同的抗病基因。用
OPN分析了68份抗性资源，其中有25份扩增出OPN。，衰明这些品种可能携带
与95－5383相同的抗性基因，而其它扩增出OPN！l。；。。和OPNll。。的抗性品种可能
携带不同于95．5383的抗性基因。
Soybean mosaic virus (SMV) occurs worldwide and is a major soybean disease
in China, resulting in yield losses and seed-quality deterioration . Using resistant
varieties is the most economic and effective method to control SMV. Northeast
China is the main soybean production region in China and the virulence of SMV 3
is the strongest. It抯 very important to identify and study new resistant
germplasms. We have screened resistant soybean germplasm by inoculation of
SMV3 and investigated the genetic diversity of part of the resistant varieties with
RAPD and morphological characters. We have demostrated the inheritance of
resistence of 95-5383 by studying the progenies of the crosses of
resisantxsusceptible . We have identified one RAPD marker closely linked to
SMV resistance gene by BSA and converted it into SCAR marker. The RAPD
marker have been mapped in soybean molecular linkage group by using a mapping
populations. The main results are summarized as followings:
1. 347 soybean accessions have been identified for the resistance to SMV3
by artificially inoculation with SMV 3, a strong virulent strain in Northeast
China. The results showed that 113 accessions were highly resistant to SMV3.
account for 32.56%; 112 accessions were moderately resistant to SMV3. account
for 32.27%; 122 accessions were susceptible to SMV3, account for 35.16%.
Resistant accessions were mainly originated from Northeast China (Liaoning),
Huang Huaihai areas (Shandong, Shanxi province and Beijing), U.S. and South
Korea. There were few resistant accessions from South China in our evaluation.
Symptoms are different in different varieties when inoculated with the same SMV
strain, showing there are interaction between soybean varieties and SMV strains.
Top necrosis symptoms were observed in introduced varieties from abroad,
indicating symptoms may relate to geographical origin of the accessions. Some
varieties with known resistance gene introduced from U.S. were identified for the
resistance to SMV3 and the relationship of resistance gene between Chinese
accessions and U.S. accessions was discussed.
2 One highly resistant soybean line (95-5383) was crossed with four
susceptible soybean varieties/line (HB 1, Tiefeng2 1, Amsoy, Williams) and one
resistant variety (P1486355) Their Fl and F2 individuals were identified for
SMV resistance by inoculation with SMV3. The results showed that in the four
crosses of resistantxsusceptible, Fl were susceptible and F2 populations
segregated in a 1 resistant: 3 susceptible (mosaic and necrosis) ratio, indicating
that 95-53 83 carries one recessive gene that confer resistance to SMV. There are
segregations of susceptibility in F2 progenies from the cross of 95-5383x
P1486355, indicating that the SMV resistance gene in 95-5383 is located at
different loci from that in P14863 55. By bulked segregant analysis (BSA) in F2
progenies of 95-5383xHB1, we have identified one codominant RAPD marker
OPNI 1980/1070 closely linked to SMV resistance gene. DNA fragment OPN1 1980
was amplified in 95-5383 and resistant bulk, OPN1 1)070 was amplified in HBI and
susceptible bulk. OPN1 1980,10,0 was amplified in Fl. Identification of the markers
in F2 plants of 95-5383xHB1 showed that the codominant marker OPN1 1930,10,0 is
closely linked to the resistance gene in 95-5383, with genetic distance of 2.1cM.
3 Both fragments of OPNI ~ and OPNI 110,0?amplified from 95-5383 and
HB 1. respectively, were purified and cloned into PGEM-T easy vector and
sequenced.
豆生产危害严重，利用抗病品种是控制SMV最经济而有效的措施。东北是我国大豆主
产区，SMV3号株系是强毒株系，抗源较少。筛选鉴定和研究新抗源具有重要的理论价
值和实践意义。本研究利用SMV3号株系对大豆资源进行了抗性鉴定，筛选出抗性种
质，并利用农艺性状和RAPD标记分析了部分抗源的遗传关系。通过对抗病×感病杂
交组合后代分析，明确了高抗SMV的大豆品系95-5383的抗性遗传规律。利用BSA法
筛选出与SMV抗性基因紧密连锁的RAPD标记，并将其转化成SCAR标记。利用作图
群体将RAPD标记定位于大豆遗传连锁图上。具体研究结果如下：
1．利用人工汁液摩擦法对不同来源的347份大豆种质资源接种SMV3号株系进行
抗性鉴定。筛选出113份高抗资源，占32.56％；112份材料中抗SMV3，占32.27％；122
份感病，占35.16％。抗源主要来自于东北春作大豆区（辽宁）和黄淮海夏作大豆区（山
东、山西、北京）以及美国和韩国，南方大豆产区抗源较少。不同品种接种后的症状类
型不同，表明品种和株系间存在互作。国外引进资源接种后顶枯症状较多，表明症状反
应和地理来源有一定关系。本文还分析了美国一些抗源对SMV3的抗性反应及抗性基
因的关系。
2．利用筛选出的高抗SMV3号株系的大豆品系95-5383与4个感病品种（品系）
HBI、铁丰21、Amsoy、Williams和抗病品种PI486355配制5个杂交组合，对各组合
的F1、F2代接种SMV鉴定抗性。结果表明，95-5383与各感病品种杂交组合的F1代
表现为感病，F2群体分离比例为3感（花叶+顶枯）:1抗，表明95-5383对SMV3号
株系的抗性受一对隐性基因控制。95-5383×PI486355的F2代接种后有感病植株分离，
表明二者对SMV3的抗性基因是不等位的。利用BSA法对95-5383×HBI的F2代进行
鉴定，筛选出与SMV抗病基因紧密连锁的共显性RAPD标记OPNII_(980/1070)。RAPD引
物OPNII在95-5383和抗池扩增出OPNII_(980)片段，在HBI和感池扩增出OPNII_(1070)片
段，在F1同时扩增出OPNII_(980)和OPNII_(1070)。用该引物分析95-5383×HBI的F2个体，
OPNII_(980/1070)与95-5383抗病基因的遗传距离为2.1cM。
3．从95-5383和HBI中分别回收OPNII_(980)和OPNII_(1070)特异片段，克隆在PGEM-
T easy vector中，并进行序列分析。结果表明OPNII_(980)和OPNII_(1070)的实际长度分别
为986bp和1084bp，同源性为93.4％，主要差别是在两个不同区段插入（缺失）54bp
和35bp的碱基。用克隆的RAPD片段OPNII_(980)作探针（SOPNII）对亲本基因组DNA
中国农业科学陨博十学位论义
进行 Southern 65交，结果表明 OPN。。和 OPN川。。在人豆基因组中是以低拷贝形式
存在。用n PD引物OPN和y卜于探针9口PN！1分析科丰1。南农1！3吕毛重组自交
系作图群体，将 Opp和 SOPN定位于 F迹锁群的抗病基冈簇上．根据克隆片段
OPN。羽IOPNll；。。的序列设计合成了一对SCAR引物SCNll，SCNll在95·5383中
扩增出与 OPN。人小相同的片段 SCN。，在 HB中扩增出与 OPN。大小相同
的片段SCN。，。，在FI同时扩增出SCN！l＿、。SCNn对95·5383 X HBI的FZ群体
扩增结果与RAPD引物OPNll分析结果完全吻台。SCAR标记SCNll。。，。对于大豆
抗SMV标记辅助选抒育种和抗病性鉴定有很）“泛的应用前景。
4．选用已经鉴定出的68份高抗SMV3的大亘资源，利用农艺性状和nPD分析
品种间的遗传关系。从 60个 nPD引物中筛选出 11个多态性引物，共扩增出刃个片
段，多态性片段为33条。扩增产物的有无分别以1和0记录，选用J＿。rd公式计算
品种之间的相似系数，利用UPGMA法构建了树状图。RAPD聚类结果和农艺性状聚
类结果具有一定的吻合性。聚类结果表明亲源关系相同、地理来额相同的品种多数聚在
一类。分聚在不同组间的人豆品种迢传背景存在差异，可能具有不同的抗病基因。用
OPN分析了68份抗性资源，其中有25份扩增出OPN。，衰明这些品种可能携带
与95－5383相同的抗性基因，而其它扩增出OPN！l。；。。和OPNll。。的抗性品种可能
携带不同于95．5383的抗性基因。
Soybean mosaic virus (SMV) occurs worldwide and is a major soybean disease
in China, resulting in yield losses and seed-quality deterioration . Using resistant
varieties is the most economic and effective method to control SMV. Northeast
China is the main soybean production region in China and the virulence of SMV 3
is the strongest. It抯 very important to identify and study new resistant
germplasms. We have screened resistant soybean germplasm by inoculation of
SMV3 and investigated the genetic diversity of part of the resistant varieties with
RAPD and morphological characters. We have demostrated the inheritance of
resistence of 95-5383 by studying the progenies of the crosses of
resisantxsusceptible . We have identified one RAPD marker closely linked to
SMV resistance gene by BSA and converted it into SCAR marker. The RAPD
marker have been mapped in soybean molecular linkage group by using a mapping
populations. The main results are summarized as followings:
1. 347 soybean accessions have been identified for the resistance to SMV3
by artificially inoculation with SMV 3, a strong virulent strain in Northeast
China. The results showed that 113 accessions were highly resistant to SMV3.
account for 32.56%; 112 accessions were moderately resistant to SMV3. account
for 32.27%; 122 accessions were susceptible to SMV3, account for 35.16%.
Resistant accessions were mainly originated from Northeast China (Liaoning),
Huang Huaihai areas (Shandong, Shanxi province and Beijing), U.S. and South
Korea. There were few resistant accessions from South China in our evaluation.
Symptoms are different in different varieties when inoculated with the same SMV
strain, showing there are interaction between soybean varieties and SMV strains.
Top necrosis symptoms were observed in introduced varieties from abroad,
indicating symptoms may relate to geographical origin of the accessions. Some
varieties with known resistance gene introduced from U.S. were identified for the
resistance to SMV3 and the relationship of resistance gene between Chinese
accessions and U.S. accessions was discussed.
2 One highly resistant soybean line (95-5383) was crossed with four
susceptible soybean varieties/line (HB 1, Tiefeng2 1, Amsoy, Williams) and one
resistant variety (P1486355) Their Fl and F2 individuals were identified for
SMV resistance by inoculation with SMV3. The results showed that in the four
crosses of resistantxsusceptible, Fl were susceptible and F2 populations
segregated in a 1 resistant: 3 susceptible (mosaic and necrosis) ratio, indicating
that 95-53 83 carries one recessive gene that confer resistance to SMV. There are
segregations of susceptibility in F2 progenies from the cross of 95-5383x
P1486355, indicating that the SMV resistance gene in 95-5383 is located at
different loci from that in P14863 55. By bulked segregant analysis (BSA) in F2
progenies of 95-5383xHB1, we have identified one codominant RAPD marker
OPNI 1980/1070 closely linked to SMV resistance gene. DNA fragment OPN1 1980
was amplified in 95-5383 and resistant bulk, OPN1 1)070 was amplified in HBI and
susceptible bulk. OPN1 1980,10,0 was amplified in Fl. Identification of the markers
in F2 plants of 95-5383xHB1 showed that the codominant marker OPN1 1930,10,0 is
closely linked to the resistance gene in 95-5383, with genetic distance of 2.1cM.
3 Both fragments of OPNI ~ and OPNI 110,0?amplified from 95-5383 and
HB 1. respectively, were purified and cloned into PGEM-T easy vector and
sequenced.
引文
1．马淑梅．1991．中国大豆品种对大豆花叶病毒(SMV)病抗病性鉴定结果．大豆科学，10(3)：240-244
2．王国勋等．1984．大豆病毒病的抗病育种．中国油料，(3)：7-10
3．长泽次男等．1981．大豆抗病毒病的育种．国外农学-大豆，6：13-16
4．刘玉芝等．1997．对大豆花叶病毒病抗源的筛选．吉林农业科学，(1)：30-34
5．刘丽君等．1994．大豆SMV病源相关蛋白的诱导与耐病性获得．大豆科学，13(4)：345-348
6．刘俊君等．1994．大豆花叶病毒NIb基因的cDNA克隆和序列分析．中国科学，B辑24(3)：265-71
7．刘俊君等，1993．大豆花叶病毒外壳蛋白基因的克隆和其在大肠杆菌中的表达．生物工程学报，9(3)：198-203
8．刘德璞等．1997，导入外源DNA获得抗SMV大豆品系．大豆科学，16(4)：277-282
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10．吕文清等．1991．六个SMV毒株在大豆不同品种上病毒浓度的变化与比较．大豆科学，10(4)：277-284
11．吕文清等．1985．东北三省大豆花叶病毒(SMV)株系的种类与分布．植物病理学报，15(4)：225-228
12．孙大敏等．1990．东北(SMV)三株系在大豆杂交亲本选配及后代选择应用．大豆科学，9(1)：60-64
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16．庄炳昌等．1993．接种大豆花叶病毒后，大豆叶片超氧物歧化酶，过氧化物酶和蛋白组份的变化．植物病理学报，23(3)：261-265
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19．余子林等．1986．湖北地区大豆花叶病毒的研究．全国大豆病害学术讨论会论文摘要汇编
20．吴宗璞等．1986．大豆品种对SMV不同毒株抗性反应与种粒斑驳关系的研究．大豆科学，5(2)：153-160
21．张玉东等．1989．大豆对二个大豆花叶病毒本地株系的抗性遗传研究．作物学报，15(3)：213-219
22．张明厚等．1998．东北亚四国(地区)SMV株系毒力比较．大豆科学，17(2)：101-107
23．张明厚等．1998．我国东北部五省市对大豆主栽品种的毒力测定．植物病理学报，28(3)：237-242
24．李延华，吕文清．1989．大豆花叶病毒—强株系在抗性不同大豆品种上的症状及病毒浓度．大豆科学，8(4)：395-399
25．李默然等．1986．筛选大豆花叶病毒种传率低品种的研究．大豆科学，5(3)：245-248
26．杨崇良等．1993．大豆品种生育特性对大豆花叶病毒种传率的影响．山东农业科学，4：9-11
27．杨崇良等．1997．北方大豆品种资源抗大豆花叶病毒筛选．植物病理学报，27(1)：27-28
28．沈文飚等．1997．大豆花叶病毒(SMV)的侵染对大豆叶片过氧化物酶及其同工酶的影响．南京农业大学学报，20(4)：88-92
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