中国甘薯种质资源遗传多样性分析及高淀粉轮回选择群体改良研究
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摘要
“甘薯——21世纪人类的健康、能源植物”。优质高产多抗专用型甘薯新品种选育是目前和将来众多的育种工作者主攻方向，而甘薯种质资源的发掘、创新和利用是获得突破性甘薯新品种的基础。因此，本研究应用农艺性状、农艺性状与形态特征的合并性状和RAPD标记对我国常用甘薯种质资源的遗传多样性进行分析；在国内率先开展了高淀粉甘薯轮回选择群体改良工作，明确了轮回选择对主要农艺性状改良效果，并结合RAPD标记对基础群体和轮回选择群体进行遗传多样性比较。其主要结果如下：
1、我国甘薯种质资源的遗传多样性
(1)基于农艺性状分析甘薯的遗传多样性：对36份种质资源的农艺性状的研究结果表明引进资源的蔓长、藤叶重、鲜薯重的级差、标准差大以及变异系数均最大，而烘干率和淀粉率、茎粗的级差、标准差小，变异系数最小，该类型资源存在丰富的遗传多样性。主推品种的重要农艺性状如蔓长、分枝数、藤叶重和鲜薯重的级差、标准差，变异系数均小于引进资源和国内资源。从农艺性状的角度讲，引进资源的遗传多样性(特别是与蔓长、藤叶重和鲜薯重的遗传多样性)显著高于国内的种质资源。因此引进资源在甘薯育种中将会发挥十分重要的作用。
(2)基于RAPD标记分析甘薯的遗传多样性：本研究从50条RAPD引物中筛选出带型清晰、稳定的20条引物对36份材料的DNA进行PCR扩增。结果发现20条引物在供试材料中共扩增出177条带，其中172条带具有多态性，多态性频率达97.18％，每个引物扩增出4～15条多态性带，平均为8.7条。不难看出36份供试材料在DNA水平上具有丰富的多样性。
(3)基于农艺性状的聚类图不能与所用材料的系谱相吻合，而RAPD标记产生的聚类图与系谱关系吻合性好，合并性状的聚类图与系谱关系吻合性较好，与RAPD标记的遗传聚类图基本一致。这表明利用RAPD标记在甘薯育种中进行辅助选择是可靠的。
(4)根据农艺性状、合并性状得出的遗传距离来看，引进资源品种(系)间的平均遗传距离最大，其次是国内资源，主推品种间变异最小。引进资源与国内资源间的平均遗传距离最大，主推品种与引进资源的平均遗传距离次之，主推品种与国内资源平均遗传距离最小。根据RAPD标记显示的遗传距离来看，主推品种内的遗传差异最大，其次是引进资源，国内资源变异最小；类型之间主推品种与引进资源的遗传差异最大，其次是主推品种与国内资源之间，国内资源与引进资源之间遗传差异最小。
(5)优异种质资源的筛选与鉴定：本研究发现国内资源中的H11-67、川薯294、绵粉1号、绵薯早秋、川薯27、肇源16、主推品种中的丰收白、潮薯1号、豫薯7号和引进资源中的台农27、内源、澳洲黄，它们的RAPD分子标记的平均遗传距离大于骨干亲本，其中H11-67含近缘野生种一半的血缘，和其它资源的遗传差异最大，具有极大的应用价值。可见，进一步加强近缘野生种资源的研究和利用，对解决甘薯育种中存在的基因源日趋狭窄等问题具有重大意义。
2、高淀粉甘薯轮回选择群体的改良效果及遗传多样性分析。
(1)在国内率先开展了高淀粉甘薯轮回选择群体改良研究，确定了基础群体构建和轮回选择的基本原则和方法，即以基础亲本群体的构建为起点，轮回选择为中心，改良创新种质资源、选育优质高效专用型品种为目的，组建优良基因数目多、均值高、遗传变异丰富的高淀粉甘薯品种(系)的基础群体，进行人工混合授粉、集团杂交，并采用轮回选择方法，不断改良群体和筛选优系，试图打破不利基因连锁，增加有利基因重组的频率，使品质性状有新的突破，为选育突破性高淀粉甘薯新品种提供新的种质和技术支撑。
(2)高淀粉甘薯轮回选择群体改良的过程中，第一轮群体(C1)总的水平比基础群体C0下降，部分性状下降显著。通过第二轮选择后，第二轮群体(C2)水平大幅度提高，远远高于C1，多数性状的平均值达极显著水平，特别是薯干、鲜薯重增益幅度最大，并且超过CO群体，鲜薯增益(10.48％)，薯干增益(10.61％)达显著水平，另外藤叶产量增益(29.7％)达极显著。而品质性状的烘干率和淀粉率的变化和其它性状变化不一致，即C1群体退化不严重，C2提高但没有达到C0水平。虽然由此看出要尽快改良提高烘干率和淀粉率的难度较大，但轮回选择群体烘干率和淀粉率与单株薯干重遗传相关系数由基础群体的较强负相关变为弱负相关和正相关。
(3)随着选择轮次的增加，单株鲜薯重和薯干重得到了显著改进，变异系数降低，同时群体方差也下降，但不显著。烘干率和淀粉率变异不大。
(4)利用RAPD标记对基础群体及轮回选择群体间(内)材料进行遗传差异研究发现，通过第一轮轮回选择后，群体内的遗传差异变化较小，第二轮选择后，群体内的遗传差异变化明显增加。从聚类结果看出，RAPD分子标记可以较准确地反应和确立各群体间(内)基因型之间的遗传差异和亲缘关系。这进一步表明，利用RAPD标记在甘薯育种中进行辅助选择是合理的、可靠的。
"Sweet potato is the health and energy plants of human in 21st century." To develop new good-quality, high-yielding, resistance and specialized type varieties is the main aim of breeders at anytime. Moreover, exploration, innovation and utilization of the sweet potato germplasm resources are the basis of gained new breakthrough sweet potato varieties. Therefore, the agronomic traits, the integrated index of agronomic and morphological traits and the RAPD markers were used for genetic diversity of the Chinese sweet potato cuurent germplasm. This study was to use recurrent selection in population improvement of high starch sweet potato in China. The results showed the there were obvious effect of recurrent selection method on agronomic traits population improvement effect. At the same time, foundation populations were compared to recurrent selection population in genetic diversity with RAPD markers. The main results were as following:
1. Genetic diversity of Chinese sweet potato germplasm resources.
(1) Genetic diversity analysis of 36 germplasm resources based on agronomic traits showed that the introduction resources existed a longest vine leaves, the largest fresh weight of relativity, standard deviation and coefficient of variation and the smallest drying rate and the rate of starch, stem diameter of relativity, the small standard deviation, coefficient of variation, which suggest that they have abundant genetic diversity. Wide planting varieties of important agronomic traits such as a long, the number of branches, leaves and vine stem heavy differential, standard deviation, the coefficient of variation was less than both the domestic and the introduction resources. As far as agronomic traits (especially with a long, vine leaves and fresh weight of the genetic diversity) are concerned, the genetic diversity of the introduced resources was significantly higher than that of domestic germplasm. Therefore, the introduced resources will play a very important role in sweet potato breeding.
(2) Genetic diversity analysis based on RAPD analysis:Twenty out of 50 stripes of RAPD primers that can produce polymorphic, clear and bright bands, with reliable repeatability, have been used for DNA amplification of 36 genotypes. The results showed that total 177 bands were amplified in these materials and 172 bands were polymorphic among them. The polymorphism frequency was up to 97.18%. Each primer amplified 4～15 polymorphic bands, with an average of 8.7. It was obvious that there were abundant diversity in the 36 materials on the DNA level.
(3) The cluster dendrogram from the agronomic traits was not consistent with the pedigree of the materials, while the cluster dendrogram based on RAPD markers was not consistent with it. The cluster dendrogram based on the integrated index was also consistent with the pedigree and the cluster dendrogram based on RAPD markers. The results showed that it was reliable to utilize RAPD markers for assisted selection in sweet potato breeding.
(4) The results derived from agronomic traits and integrated index showed the largest genetic distance was in introduced resources, and the next was in domestic resources, while the smallest was in main planting varieties. Moreover, the largest average genetic distance was between introduced resources and domestic resources, and the second between introduced resources and main planting varieties, while the smallest between domestic resources and main planting varieties. RAPD analysis showed that the largest genetic diversity exists within in main planting varieties, and the next within introduced resources, while the smallest in domestic resources. Moreover, the largest genetic difference existed within types was between main planting varieties and introduced resources, and the next between main planting varieties and domestic resources, while the smallest between domestic resources and introduced resources.
(5) The results showed that the average genetic distances, based on the RAPD markers with the genotypes including H11-67, chuanshu 294, mianfen 1, mainshuzaoqiu, chuanshu 27 and zhapyuan 16 in domestic resources, fengshoubai, chaoshu 1 and yushu 7 in main planting resources, and tainong 27, neiyuan, and aozhouhuang in introduced resources, were larger than their main parents. Among them, H11-67 was with half blood of the wild species, and was large difference from other resources. This showed that it was significant to further strengthen the study and utilization of the wild relatives resources to broaden the genetic basis in the sweet potato breeding.
2 Improvement effect and genetic diversity analysis of the high starch sweet potato recurrent selection populations
(1) To determine basic principles and methods of the construction of high starch sweet potato basic populations and the recurrent selection, it commenced from the construction of basic parents populations, centered the recurrent selection, and aimed at improving innovation germplasm resources and identification of high sweet potato starch based groups and the construction of recurrent selection of the basic principles and breeding good-quality, high-yielding, resistance and specialized varieties. The basic populations of excellent genes, high mean, and rich genetic variation high starch sweet potato varieties or lines had been constructed, with artificial mixture pollination or group hybridizing, recurrent selection to break the linked negative gene, enhance favorable gene recombination frequency and create new breakthrough quality traits.
(2) During recurrent selecting, the first selection population (C_1) had the reduced agronomic traits compared with the basic population. After the second selecting, the agronomic traits especially the weight of dried potato and fresh potato of the population (C_2) were infinitely improved compared with C_1. The change tendency of quality traits such as the drying rate and starch rate were different from others. Results of selection showed that after a slightly degradation of C_1, C_2 had the improvement but they had not reached the level of C_0. Although it is rather difficult to enhance the drying rate and starch rate in a short period, the genetic correlation coefficient between the drying and starch rate and the dry weight per plant was changed from the strong negative correlation to the weak negative or positive correlation with recurrent selecting.
(3) Along with the increase of selection times, fresh weight and dry weight per plant were significantly improved with the slightly lowered variation coefficient, and there was the small variation in drying and starch rate.
(4) RAPD analysis results showed that the genetic difference within populations reduced after the first selection and it was saliently increased after the second selection. Cluster analyses showed that RAPD markers could accurately response and determine the genetic variations and the pedigree relations. The result further indicated that RAPD markers are reasonable and reliable markers in assisted selection breeding of sweet potato.
1、我国甘薯种质资源的遗传多样性
(1)基于农艺性状分析甘薯的遗传多样性：对36份种质资源的农艺性状的研究结果表明引进资源的蔓长、藤叶重、鲜薯重的级差、标准差大以及变异系数均最大，而烘干率和淀粉率、茎粗的级差、标准差小，变异系数最小，该类型资源存在丰富的遗传多样性。主推品种的重要农艺性状如蔓长、分枝数、藤叶重和鲜薯重的级差、标准差，变异系数均小于引进资源和国内资源。从农艺性状的角度讲，引进资源的遗传多样性(特别是与蔓长、藤叶重和鲜薯重的遗传多样性)显著高于国内的种质资源。因此引进资源在甘薯育种中将会发挥十分重要的作用。
(2)基于RAPD标记分析甘薯的遗传多样性：本研究从50条RAPD引物中筛选出带型清晰、稳定的20条引物对36份材料的DNA进行PCR扩增。结果发现20条引物在供试材料中共扩增出177条带，其中172条带具有多态性，多态性频率达97.18％，每个引物扩增出4～15条多态性带，平均为8.7条。不难看出36份供试材料在DNA水平上具有丰富的多样性。
(3)基于农艺性状的聚类图不能与所用材料的系谱相吻合，而RAPD标记产生的聚类图与系谱关系吻合性好，合并性状的聚类图与系谱关系吻合性较好，与RAPD标记的遗传聚类图基本一致。这表明利用RAPD标记在甘薯育种中进行辅助选择是可靠的。
(4)根据农艺性状、合并性状得出的遗传距离来看，引进资源品种(系)间的平均遗传距离最大，其次是国内资源，主推品种间变异最小。引进资源与国内资源间的平均遗传距离最大，主推品种与引进资源的平均遗传距离次之，主推品种与国内资源平均遗传距离最小。根据RAPD标记显示的遗传距离来看，主推品种内的遗传差异最大，其次是引进资源，国内资源变异最小；类型之间主推品种与引进资源的遗传差异最大，其次是主推品种与国内资源之间，国内资源与引进资源之间遗传差异最小。
(5)优异种质资源的筛选与鉴定：本研究发现国内资源中的H11-67、川薯294、绵粉1号、绵薯早秋、川薯27、肇源16、主推品种中的丰收白、潮薯1号、豫薯7号和引进资源中的台农27、内源、澳洲黄，它们的RAPD分子标记的平均遗传距离大于骨干亲本，其中H11-67含近缘野生种一半的血缘，和其它资源的遗传差异最大，具有极大的应用价值。可见，进一步加强近缘野生种资源的研究和利用，对解决甘薯育种中存在的基因源日趋狭窄等问题具有重大意义。
2、高淀粉甘薯轮回选择群体的改良效果及遗传多样性分析。
(1)在国内率先开展了高淀粉甘薯轮回选择群体改良研究，确定了基础群体构建和轮回选择的基本原则和方法，即以基础亲本群体的构建为起点，轮回选择为中心，改良创新种质资源、选育优质高效专用型品种为目的，组建优良基因数目多、均值高、遗传变异丰富的高淀粉甘薯品种(系)的基础群体，进行人工混合授粉、集团杂交，并采用轮回选择方法，不断改良群体和筛选优系，试图打破不利基因连锁，增加有利基因重组的频率，使品质性状有新的突破，为选育突破性高淀粉甘薯新品种提供新的种质和技术支撑。
(2)高淀粉甘薯轮回选择群体改良的过程中，第一轮群体(C1)总的水平比基础群体C0下降，部分性状下降显著。通过第二轮选择后，第二轮群体(C2)水平大幅度提高，远远高于C1，多数性状的平均值达极显著水平，特别是薯干、鲜薯重增益幅度最大，并且超过CO群体，鲜薯增益(10.48％)，薯干增益(10.61％)达显著水平，另外藤叶产量增益(29.7％)达极显著。而品质性状的烘干率和淀粉率的变化和其它性状变化不一致，即C1群体退化不严重，C2提高但没有达到C0水平。虽然由此看出要尽快改良提高烘干率和淀粉率的难度较大，但轮回选择群体烘干率和淀粉率与单株薯干重遗传相关系数由基础群体的较强负相关变为弱负相关和正相关。
(3)随着选择轮次的增加，单株鲜薯重和薯干重得到了显著改进，变异系数降低，同时群体方差也下降，但不显著。烘干率和淀粉率变异不大。
(4)利用RAPD标记对基础群体及轮回选择群体间(内)材料进行遗传差异研究发现，通过第一轮轮回选择后，群体内的遗传差异变化较小，第二轮选择后，群体内的遗传差异变化明显增加。从聚类结果看出，RAPD分子标记可以较准确地反应和确立各群体间(内)基因型之间的遗传差异和亲缘关系。这进一步表明，利用RAPD标记在甘薯育种中进行辅助选择是合理的、可靠的。
"Sweet potato is the health and energy plants of human in 21st century." To develop new good-quality, high-yielding, resistance and specialized type varieties is the main aim of breeders at anytime. Moreover, exploration, innovation and utilization of the sweet potato germplasm resources are the basis of gained new breakthrough sweet potato varieties. Therefore, the agronomic traits, the integrated index of agronomic and morphological traits and the RAPD markers were used for genetic diversity of the Chinese sweet potato cuurent germplasm. This study was to use recurrent selection in population improvement of high starch sweet potato in China. The results showed the there were obvious effect of recurrent selection method on agronomic traits population improvement effect. At the same time, foundation populations were compared to recurrent selection population in genetic diversity with RAPD markers. The main results were as following:
1. Genetic diversity of Chinese sweet potato germplasm resources.
(1) Genetic diversity analysis of 36 germplasm resources based on agronomic traits showed that the introduction resources existed a longest vine leaves, the largest fresh weight of relativity, standard deviation and coefficient of variation and the smallest drying rate and the rate of starch, stem diameter of relativity, the small standard deviation, coefficient of variation, which suggest that they have abundant genetic diversity. Wide planting varieties of important agronomic traits such as a long, the number of branches, leaves and vine stem heavy differential, standard deviation, the coefficient of variation was less than both the domestic and the introduction resources. As far as agronomic traits (especially with a long, vine leaves and fresh weight of the genetic diversity) are concerned, the genetic diversity of the introduced resources was significantly higher than that of domestic germplasm. Therefore, the introduced resources will play a very important role in sweet potato breeding.
(2) Genetic diversity analysis based on RAPD analysis:Twenty out of 50 stripes of RAPD primers that can produce polymorphic, clear and bright bands, with reliable repeatability, have been used for DNA amplification of 36 genotypes. The results showed that total 177 bands were amplified in these materials and 172 bands were polymorphic among them. The polymorphism frequency was up to 97.18%. Each primer amplified 4～15 polymorphic bands, with an average of 8.7. It was obvious that there were abundant diversity in the 36 materials on the DNA level.
(3) The cluster dendrogram from the agronomic traits was not consistent with the pedigree of the materials, while the cluster dendrogram based on RAPD markers was not consistent with it. The cluster dendrogram based on the integrated index was also consistent with the pedigree and the cluster dendrogram based on RAPD markers. The results showed that it was reliable to utilize RAPD markers for assisted selection in sweet potato breeding.
(4) The results derived from agronomic traits and integrated index showed the largest genetic distance was in introduced resources, and the next was in domestic resources, while the smallest was in main planting varieties. Moreover, the largest average genetic distance was between introduced resources and domestic resources, and the second between introduced resources and main planting varieties, while the smallest between domestic resources and main planting varieties. RAPD analysis showed that the largest genetic diversity exists within in main planting varieties, and the next within introduced resources, while the smallest in domestic resources. Moreover, the largest genetic difference existed within types was between main planting varieties and introduced resources, and the next between main planting varieties and domestic resources, while the smallest between domestic resources and introduced resources.
(5) The results showed that the average genetic distances, based on the RAPD markers with the genotypes including H11-67, chuanshu 294, mianfen 1, mainshuzaoqiu, chuanshu 27 and zhapyuan 16 in domestic resources, fengshoubai, chaoshu 1 and yushu 7 in main planting resources, and tainong 27, neiyuan, and aozhouhuang in introduced resources, were larger than their main parents. Among them, H11-67 was with half blood of the wild species, and was large difference from other resources. This showed that it was significant to further strengthen the study and utilization of the wild relatives resources to broaden the genetic basis in the sweet potato breeding.
2 Improvement effect and genetic diversity analysis of the high starch sweet potato recurrent selection populations
(1) To determine basic principles and methods of the construction of high starch sweet potato basic populations and the recurrent selection, it commenced from the construction of basic parents populations, centered the recurrent selection, and aimed at improving innovation germplasm resources and identification of high sweet potato starch based groups and the construction of recurrent selection of the basic principles and breeding good-quality, high-yielding, resistance and specialized varieties. The basic populations of excellent genes, high mean, and rich genetic variation high starch sweet potato varieties or lines had been constructed, with artificial mixture pollination or group hybridizing, recurrent selection to break the linked negative gene, enhance favorable gene recombination frequency and create new breakthrough quality traits.
(2) During recurrent selecting, the first selection population (C_1) had the reduced agronomic traits compared with the basic population. After the second selecting, the agronomic traits especially the weight of dried potato and fresh potato of the population (C_2) were infinitely improved compared with C_1. The change tendency of quality traits such as the drying rate and starch rate were different from others. Results of selection showed that after a slightly degradation of C_1, C_2 had the improvement but they had not reached the level of C_0. Although it is rather difficult to enhance the drying rate and starch rate in a short period, the genetic correlation coefficient between the drying and starch rate and the dry weight per plant was changed from the strong negative correlation to the weak negative or positive correlation with recurrent selecting.
(3) Along with the increase of selection times, fresh weight and dry weight per plant were significantly improved with the slightly lowered variation coefficient, and there was the small variation in drying and starch rate.
(4) RAPD analysis results showed that the genetic difference within populations reduced after the first selection and it was saliently increased after the second selection. Cluster analyses showed that RAPD markers could accurately response and determine the genetic variations and the pedigree relations. The result further indicated that RAPD markers are reasonable and reliable markers in assisted selection breeding of sweet potato.
引文
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