腺枝葡萄鉴定与评价及遗传多样性研究
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摘要
腺枝葡萄(Vitis adenoclada Hand.-Mazz)是一种中国特有野生葡萄种类,广泛分布于湖南省、福建省、广西区等长江以南省区。本论文以湖南农业大学园艺园林学院葡萄教学科研基地收集的腺枝葡萄的不同类型及引自中国农科院郑州果树研究所的腺枝葡萄等为试材,对其生物学特性进行了观测,进行扦插生根与嫁接亲和力及光合特性研究,并利用EST-SSR分子标记技术进行遗传多样性与亲缘关系分析。对腺枝葡萄抗葡萄黑痘病和霜霉病进行鉴定,为进一步开展葡萄抗性育种以及探明腺枝葡萄对黑痘病和霜霉病的抗病机理提供理论依据。主要研究结果如下：
1.腺枝葡萄根据花器官的形态,分为三种类型,即两性花类型、雄花类型和雌花类型；最主要的特征是嫩枝具有紫红色腺毛。在外观形态上,三种类型没有明显的差别,雄株不着生果粒是其主要区别。
2.腺枝葡萄的三种不同类型的物候期基本相同,在湖南省长沙市,4月中旬开始萌芽,6月上、中旬开始开花,9月下旬果实充分成熟,11月中旬至12月上旬植株落叶,进入休眠。
3.硬枝扦插成活率腺枝葡萄较华东葡萄高,但较巨峰(成活率95%左右)及栽培葡萄品种低。插穗直径以8～10mm成活率最高。红宝石无核、红地球、美人指、粉红亚都蜜等栽培品种以腺枝葡萄为砧木,嫁接成活率为40%-70%,明显低于5BB、贝达、刺葡萄、华佳8号等砧木。嫁接后成枝率、结果枝率、接穗叶面积也低于其他砧木,但抗病性明显强于5BB、贝达、华佳8号等三种砧木。
4.在相同的有效辐射强度下,腺枝葡萄的光合能力比毛葡萄弱。腺枝葡萄的光合“午休现象”不明显,其光饱和点为1500μmolm-2s-1,光补偿点为90μmolm-2s-1。腺枝葡萄的胞间C02浓度较高,在较低浓度C02的条件下,C02的利用能力相对较强。
5.腺枝葡萄对葡萄黑痘病和葡萄霜霉病的抗性较强；在与抗霜霉病相关的生化指标中,腺枝葡萄叶片内可溶性糖、可溶性蛋白质含量随发病时间逐渐推移而逐渐下降,超氧化物歧化酶活性则随发病时间逐渐推移呈上升趋势；与抗黑痘病的相关的生化指标中,腺枝葡萄叶片内可溶性糖含量随着发病时间逐渐推移逐渐上升,而过氧化物酶和超氧化物歧化酶的活性则随发病时间逐渐推移呈先上升,随后下降的趋势。
6.以腺枝葡萄等19份葡萄材料叶片的8个主要性状,进行聚类分析,种间分布在0.0705～2.3762,变化幅度很大,反映出腺枝葡萄种内的形态多样性,也充分反映了葡萄属植物种类的形态多样性。
7.建立了EST-SSR反应体系,适合于腺枝葡萄。用形态学差异较大的维多利亚,腺枝葡萄-1,户太8号,贝达等4个葡萄种类和品种,用16对EST-SSR引物进行筛选,一共筛选出8对引物,其多态性较好,再进一步进行扩增,共扩增出49条DNA带,有37条具有多态性,占总数的75.51%。
8.本研究对腺枝葡萄等23份葡萄资源与品种进行EST-SSR分析,进行聚类分析,结果表明：当相似系数0.8为临界值时,可将试验材料分为8个类群。腺枝葡萄与毛葡萄的遗传相似系数为0.9556,亲缘关系比较近,腺枝葡萄与毛葡萄归为一组。
Vitis adenoclada Hand.-Mazz is a special wild grape resource which distributes in Hunan and other provinces of the southern area of Yangtze River. In the paper is an attempt to study the biological characteristics, cutting rooting capability, graft compatibility, photosynthetic characteristics, resistance to Sphaceloma ampelinum and Plasmopara viticola of V. adenoclada Hand.-Mazz collections in Hunan Agricultural University and from Zhengzhou Fruit Research Institute of the Chinese Academy of Agricultural Sciences. Meanwhile, genetic diversity and phylogenetic relationship were analyzed by EST-SSR method. The main results are as below:
1. V. adenoclada Hand.-Mazz divided into three types including hermaphrodite flower, female flower and male flower types. The common character was their glandular trichomes on twigs. However, the obvious difference in morphology among the three types was not observed. In a word, the three types differ in flower organ and fruiting ability.
2. The phenological phases of three types of V. adenoclada Hand.-Mazz were basically consistent, budding at mid-April, blossoming at early and mid-June, ripening at late-September, defoliating at mid-November to early-December in Changsha, Hunan province.
3. The hard wood cutting survival ratio of V. adenoclada Hand.-Mazz, reaching to the highest with8-10mm diameter, was much higher than that of V. pseudoreticulata W.T.Wang but lower than that of Kyoho(V. vinifera, about95%). The influence of V. adenoclada Hand.-Mazz as rootstock of scion cultivars was investigated as well. Results showed that the survival rate of Ruby Seedless, Red Globe, Manicure Finger and Yatomi Rosa grafted on V. adenoclada Hand.-Mazz were40%-70%, apparently lower than that grafted on5BB, Beta, and NO.8Huajia rootstocks, so as the twig formation rate, fruit branch rate and scion leaf area. However the disease resistance of scion cultivars grafted on V. adenoclada Hand.-Mazz was obviously stronger than that grafted on5BB, Beta, and NO.8Huajia rootstocks.
4. The photosynthetic capacity of V.adenoclada Hand.-Mazz was weaker than that of V. quinquangularis under the same intensity of the photosynthetic active radiation. No typical midday depression of photosynthesis is there in leaves of V.adenoclada Hand.-Mazz, with light saturation points of1500μmol·m-2·s-1, light compensation points of90μmol·m-2·s-1, high intercellular CO2concentration and better CO2utilization capacity under lower CO2concentration.
5. It was observed that V.adenoclona Hand.-Mazz has a stronger resistance to Sphaceloma ampelinum and Plasmopara viticola. The soluble protein and the soluble sugar content in the leaves dropped gradually but the superoxide dismutase enzyme activity escalated, along with the developing of Plasmopara viticola. On the other hand, the soluble sugar content in the leaves increased and the activities of superoxide dismutase enzyme and peroxide enzyme increased first and then decreased, following the developing of Sphaceloma ampelinum.
6. The diversity of morphological traits among19grape resources was observed using phylogenetic cluster analysis of8leaf traits, differing greatly in genetic distance of species from0.0705to2.3762.
7. An effective and reproductive PCR reaction system for detecting V.adenoclona Hand.-Mazz EST-SSR was established.8pairs of informative polymorphism primers were selected from the group consisting of16pairs of EST-SSR primers using V.adenoclona Hand.-Mazz-1, Bate, Hutai no.8and Victoria as test materials.8pairs of informative polymorphism primers produced49DNA bands, of which37DNA bands were polymorphic,75.5%of the total DNA bands.
8. Total23related grape materials were analyzed by EST-SSR method. The phylogenetic clustering analysis showed that0.8for critical value,23samples can be divided into eight clusters and the genetic coefficient between V. adenoclada and V. quinquangularis was0.9556, suggesting a relatively close genetic relationship between the two species.
1.腺枝葡萄根据花器官的形态,分为三种类型,即两性花类型、雄花类型和雌花类型；最主要的特征是嫩枝具有紫红色腺毛。在外观形态上,三种类型没有明显的差别,雄株不着生果粒是其主要区别。
2.腺枝葡萄的三种不同类型的物候期基本相同,在湖南省长沙市,4月中旬开始萌芽,6月上、中旬开始开花,9月下旬果实充分成熟,11月中旬至12月上旬植株落叶,进入休眠。
3.硬枝扦插成活率腺枝葡萄较华东葡萄高,但较巨峰(成活率95%左右)及栽培葡萄品种低。插穗直径以8～10mm成活率最高。红宝石无核、红地球、美人指、粉红亚都蜜等栽培品种以腺枝葡萄为砧木,嫁接成活率为40%-70%,明显低于5BB、贝达、刺葡萄、华佳8号等砧木。嫁接后成枝率、结果枝率、接穗叶面积也低于其他砧木,但抗病性明显强于5BB、贝达、华佳8号等三种砧木。
4.在相同的有效辐射强度下,腺枝葡萄的光合能力比毛葡萄弱。腺枝葡萄的光合“午休现象”不明显,其光饱和点为1500μmolm-2s-1,光补偿点为90μmolm-2s-1。腺枝葡萄的胞间C02浓度较高,在较低浓度C02的条件下,C02的利用能力相对较强。
5.腺枝葡萄对葡萄黑痘病和葡萄霜霉病的抗性较强；在与抗霜霉病相关的生化指标中,腺枝葡萄叶片内可溶性糖、可溶性蛋白质含量随发病时间逐渐推移而逐渐下降,超氧化物歧化酶活性则随发病时间逐渐推移呈上升趋势；与抗黑痘病的相关的生化指标中,腺枝葡萄叶片内可溶性糖含量随着发病时间逐渐推移逐渐上升,而过氧化物酶和超氧化物歧化酶的活性则随发病时间逐渐推移呈先上升,随后下降的趋势。
6.以腺枝葡萄等19份葡萄材料叶片的8个主要性状,进行聚类分析,种间分布在0.0705～2.3762,变化幅度很大,反映出腺枝葡萄种内的形态多样性,也充分反映了葡萄属植物种类的形态多样性。
7.建立了EST-SSR反应体系,适合于腺枝葡萄。用形态学差异较大的维多利亚,腺枝葡萄-1,户太8号,贝达等4个葡萄种类和品种,用16对EST-SSR引物进行筛选,一共筛选出8对引物,其多态性较好,再进一步进行扩增,共扩增出49条DNA带,有37条具有多态性,占总数的75.51%。
8.本研究对腺枝葡萄等23份葡萄资源与品种进行EST-SSR分析,进行聚类分析,结果表明：当相似系数0.8为临界值时,可将试验材料分为8个类群。腺枝葡萄与毛葡萄的遗传相似系数为0.9556,亲缘关系比较近,腺枝葡萄与毛葡萄归为一组。
Vitis adenoclada Hand.-Mazz is a special wild grape resource which distributes in Hunan and other provinces of the southern area of Yangtze River. In the paper is an attempt to study the biological characteristics, cutting rooting capability, graft compatibility, photosynthetic characteristics, resistance to Sphaceloma ampelinum and Plasmopara viticola of V. adenoclada Hand.-Mazz collections in Hunan Agricultural University and from Zhengzhou Fruit Research Institute of the Chinese Academy of Agricultural Sciences. Meanwhile, genetic diversity and phylogenetic relationship were analyzed by EST-SSR method. The main results are as below:
1. V. adenoclada Hand.-Mazz divided into three types including hermaphrodite flower, female flower and male flower types. The common character was their glandular trichomes on twigs. However, the obvious difference in morphology among the three types was not observed. In a word, the three types differ in flower organ and fruiting ability.
2. The phenological phases of three types of V. adenoclada Hand.-Mazz were basically consistent, budding at mid-April, blossoming at early and mid-June, ripening at late-September, defoliating at mid-November to early-December in Changsha, Hunan province.
3. The hard wood cutting survival ratio of V. adenoclada Hand.-Mazz, reaching to the highest with8-10mm diameter, was much higher than that of V. pseudoreticulata W.T.Wang but lower than that of Kyoho(V. vinifera, about95%). The influence of V. adenoclada Hand.-Mazz as rootstock of scion cultivars was investigated as well. Results showed that the survival rate of Ruby Seedless, Red Globe, Manicure Finger and Yatomi Rosa grafted on V. adenoclada Hand.-Mazz were40%-70%, apparently lower than that grafted on5BB, Beta, and NO.8Huajia rootstocks, so as the twig formation rate, fruit branch rate and scion leaf area. However the disease resistance of scion cultivars grafted on V. adenoclada Hand.-Mazz was obviously stronger than that grafted on5BB, Beta, and NO.8Huajia rootstocks.
4. The photosynthetic capacity of V.adenoclada Hand.-Mazz was weaker than that of V. quinquangularis under the same intensity of the photosynthetic active radiation. No typical midday depression of photosynthesis is there in leaves of V.adenoclada Hand.-Mazz, with light saturation points of1500μmol·m-2·s-1, light compensation points of90μmol·m-2·s-1, high intercellular CO2concentration and better CO2utilization capacity under lower CO2concentration.
5. It was observed that V.adenoclona Hand.-Mazz has a stronger resistance to Sphaceloma ampelinum and Plasmopara viticola. The soluble protein and the soluble sugar content in the leaves dropped gradually but the superoxide dismutase enzyme activity escalated, along with the developing of Plasmopara viticola. On the other hand, the soluble sugar content in the leaves increased and the activities of superoxide dismutase enzyme and peroxide enzyme increased first and then decreased, following the developing of Sphaceloma ampelinum.
6. The diversity of morphological traits among19grape resources was observed using phylogenetic cluster analysis of8leaf traits, differing greatly in genetic distance of species from0.0705to2.3762.
7. An effective and reproductive PCR reaction system for detecting V.adenoclona Hand.-Mazz EST-SSR was established.8pairs of informative polymorphism primers were selected from the group consisting of16pairs of EST-SSR primers using V.adenoclona Hand.-Mazz-1, Bate, Hutai no.8and Victoria as test materials.8pairs of informative polymorphism primers produced49DNA bands, of which37DNA bands were polymorphic,75.5%of the total DNA bands.
8. Total23related grape materials were analyzed by EST-SSR method. The phylogenetic clustering analysis showed that0.8for critical value,23samples can be divided into eight clusters and the genetic coefficient between V. adenoclada and V. quinquangularis was0.9556, suggesting a relatively close genetic relationship between the two species.
引文
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