胡枝子传粉生物学及分子标记应用的研究
摘要
胡枝子属(Lespedeza Michx)植物是我国重要的生态经济型灌木树种,该属植物具有分布广泛,适应性强,产品用途广等特点,近年来对其开发利用逐渐受到重视。但同国外相比,我国对胡枝子属的开发利用还处于较低的水平。为了更好地开展遗传改良工作,有必要对胡枝子属植物的遗传背景和亲缘关系等生物学特性进行深入系统的研究。本研究以二色胡枝子的24个种源和10种胡枝子为研究材料,围绕二色胡枝子群体遗传结构和部分胡枝子属植物种间系统发育两个问题,采用传粉生物学、分子标记和DNA测序等技术手段,开展了二色胡枝子种群遗传多样性和胡枝子种间部分个体的系统发育研究。此外,还首次开发出胡枝子属物种特异性SSR引物。通过以上研究,得到以下主要结论:
1.开展了花粉活力、柱头可授性和繁育系统研究。筛选出适于二色胡枝子花粉培养的液体培养基,并找到快速测定其花粉活力的染色法。对比不同贮藏条件下花粉活力变化发现,4℃干燥条件下花粉活力可维持一周。通过采用杂交指数(OCI)、花粉-胚珠比(P/O)、去雄—套袋—人工授粉试验检测到二色胡枝子的繁育系统属于部分自交亲和,异交且需要传粉者的类型。二色胡枝子属于虫媒传粉,高频率昆虫访花可明显提高二色胡枝子的结实率。
2.建立并优化了二色胡枝子SRAP反应体系,并对二色胡枝子种源开展遗传多样性研究。筛选出17对SRAP引物组合,对24个种源材料进行了遗传多样性分析,共扩增出464条带,多态性位点百分率为87.28%。种源水平上,每位点的平均等位基因数1.881,平均有效等位基因数1.568,Nei's基因多样性指数为0.331,Shannon信息指数为0.491。种源间遗传距离变幅为0.1029～0.3434,种源间遗传一致度范围为0.709～0.902。种源间的遗传分化系数Gst为0.4157,这表明58.4%的遗传多样性存在于二色胡枝子种源内。种源间基因流Nm为0.7028。基于Nei's遗传距离的聚类结果,二色胡枝子24个种源可以分为五大类。同时应用SRAP标记对10种胡枝子进行了亲缘关系研究,以遗传距离0.52为阀值,可以将10种胡枝子分为三大类,阴山胡枝子和牛枝子首先聚为一类,截叶胡枝子、绒毛胡枝子和多花胡枝子聚到一起,其余的五种胡枝子聚为一类。这一聚类结果并不完全符合传统形态学上的分类情况。
3.结合ISSR和染色体步移技术开发出SSR引物。依据ISSR引物扩增测序结果设计出6对巢式PCR引物,结合巢式PCR结果共开发出20对SSR引物,其中10对可用于胡枝子属内不同种间的转移扩增。同时应用所开发的SSR引物对10种胡枝子的亲缘关系进行了研究,结果表明,10种胡枝子可分为三大类,第一类包括截叶胡枝子和绒毛胡枝子,阴山胡枝子和牛枝子先聚到一起,然后再与多花胡枝子聚为一类,其余的五种胡枝子聚为一类。表明多花胡枝子与阴山胡枝子,以及牛枝子的亲缘关系相对较近。
4.依据测序结果构建了胡枝子属部分种的系统发育树。测序得到了9种胡枝子的ITS序列和10种胡枝子的叶绿体trnL-F序列,同时结合GenBank上近缘物种的相关序列开展胡枝子属系统发育研究。序列分析结果表明,ITS序列中包含61个信息位点,占比对长度的10%,相比之下,trnL-F序列中仅有不到1%的信息位点。对两个片段分别采用MP和NJ法构建了系统发育树,结果表明,对单个片段而言,基于不同方法构建的系统发育树大体上一致,仅是在自展支持率上略有不同,并且NJ树中小支的关系比MP树清楚。对比两个片段构建的系统发育树,显示二者的拓扑结构大体相似,但是多花胡枝子的聚类位置明显有不同。在基于ITS序列的系统发育树中,多花胡枝子和大胡枝子组的物种聚到一起,而在基于tmL-F序列的系统发育树中,它又独立于大胡枝子组和胡枝子组外,单独为一支。
Lespedeza species are an important ecological and economic-type native shrub species in China.In recent years,it has received special attention on its strong adaptability and wide use.Compared with developed countries,utilization of this genus is still in a low level in China.To carry out genetic improvement for Lespedeza genus,it is necessary to unfold detailed researching in genetic background and phylogenetic relationship between them.In this study,24 provenances in Lespedeza bicolor and 10 species in Lespedeza were collected and regarded as tested materials.Population genetic structure of L.bicolor and relationship of some Lespedeza species were investigated using the methods of pollination biology,molecular markers and DNA sequencing.This study firstly developed the SSR-specific primers of Lespedeza genus by combining ISSR markers and chromosomal walking technique.The results are as follows:
1) Pollen viability,stigma receptivity and breeding system for L.bicolor were carried out.The optimum liquid medium for pollen germination of L.bicolor was screened and rapid staining method was found.The changes of pollen viability stored in different conditions suggested that the pollen vitality can last 7 days under 4℃and dry conditions can last 7 days.Results of hybrid index(OCI), pollen-ovule ratio(P/O) and artificial pollination indicated that the breeding system of L.bicolor belonged to outcrossing,part self-compatibility and pollinator-dependence.As an entomophilous plant, the high frequency of visits can significantly increase the seed setting rate of L.bicolor.
2) The SRAP reaction system was established and optimized.Senventeen pairs of primer combinations were selected from 484 sets of SRAP primer combinations.Taken 248 individuals as materials,a total of 464 bands were amplified by using 17 pairs of primer combinations,among them 360 bands were polymorphic;the percentage of polymorphic bands was 87.28%.At provenance level, average number of alleles and effective alleles on per locus was 1.881 and 1.568.The Nei's gene diversity index was 0.331,and Shannon's information index was 0.491.Genetic distance between species ranged from 0.1029 to 0.3434,and genetic identity of inter-provenance ranged from 0.709 to 0.902.The coefficient of genetic differentiation was 0.4157,which indicated that 58.4%variabilities occurred in provenance of L.bicolor.The gene flow among provenances(Nm) was 0.7028.According to the UPMGA cluster analysis based on the genetic distance,24 provenances of L.bicolor were divided into five groups.In accordance with cluster figure,10 kinds of Lespedeza were divided into three categories,while L.fomosa showed a higher independence.
3) Chromosomal walking and ISSR technologies were used in developing SSR primers.According to sequencing results of ISSR PCR amplification,6 pairs of newly designed primers were successfully applied to chromosomal walking.Of the 20 obtained SSR primers,10 primers showed high polymorphisms.All of them can amplify in other species of Lespedeza.At the same time,new SSR primers were utilized to the research of the relationship between 10 Lespedeza species.UPGMA cluster analyses based on Nei's genetic distance dividedthe 10 species into three main groups.
4) Phylogenetic trees of Lespedeza genus were constracted based on sequencing results.Nine ITS sequences and 10 trnL-F sequences were amplified and sequenced.The results of sequence analysis showed that ITS sequence had 61 information sites,accounting for more than 10 percent of the sequencing length.Compared with ITS sequencing results,trnL-F sequence only provided less than 1% information sites.Phylogeny trees of Maximum Parsimony(MP) and Neighbor-Joining(NJ) were constructed based on two fragments respectively.For single fragment,phylogenetic trees constructed by different methods were largely consistent and there was only slightly different in bootstrap values.As a whole,NJ tree reflected more clear than the MP tree in the relationships between subgroups. Comparison analysis phylogenetic tree showed that the tapnology of trees based on two fragments are almost identical,but the cluster location of L.floribunda was obvious differences,which clustered with L.bicolor and other Lespedeza species in phylogenetic tree based on ITS sequence,as contrast,it is independent in trnL-F phylogenetic tree.
1.开展了花粉活力、柱头可授性和繁育系统研究。筛选出适于二色胡枝子花粉培养的液体培养基,并找到快速测定其花粉活力的染色法。对比不同贮藏条件下花粉活力变化发现,4℃干燥条件下花粉活力可维持一周。通过采用杂交指数(OCI)、花粉-胚珠比(P/O)、去雄—套袋—人工授粉试验检测到二色胡枝子的繁育系统属于部分自交亲和,异交且需要传粉者的类型。二色胡枝子属于虫媒传粉,高频率昆虫访花可明显提高二色胡枝子的结实率。
2.建立并优化了二色胡枝子SRAP反应体系,并对二色胡枝子种源开展遗传多样性研究。筛选出17对SRAP引物组合,对24个种源材料进行了遗传多样性分析,共扩增出464条带,多态性位点百分率为87.28%。种源水平上,每位点的平均等位基因数1.881,平均有效等位基因数1.568,Nei's基因多样性指数为0.331,Shannon信息指数为0.491。种源间遗传距离变幅为0.1029～0.3434,种源间遗传一致度范围为0.709～0.902。种源间的遗传分化系数Gst为0.4157,这表明58.4%的遗传多样性存在于二色胡枝子种源内。种源间基因流Nm为0.7028。基于Nei's遗传距离的聚类结果,二色胡枝子24个种源可以分为五大类。同时应用SRAP标记对10种胡枝子进行了亲缘关系研究,以遗传距离0.52为阀值,可以将10种胡枝子分为三大类,阴山胡枝子和牛枝子首先聚为一类,截叶胡枝子、绒毛胡枝子和多花胡枝子聚到一起,其余的五种胡枝子聚为一类。这一聚类结果并不完全符合传统形态学上的分类情况。
3.结合ISSR和染色体步移技术开发出SSR引物。依据ISSR引物扩增测序结果设计出6对巢式PCR引物,结合巢式PCR结果共开发出20对SSR引物,其中10对可用于胡枝子属内不同种间的转移扩增。同时应用所开发的SSR引物对10种胡枝子的亲缘关系进行了研究,结果表明,10种胡枝子可分为三大类,第一类包括截叶胡枝子和绒毛胡枝子,阴山胡枝子和牛枝子先聚到一起,然后再与多花胡枝子聚为一类,其余的五种胡枝子聚为一类。表明多花胡枝子与阴山胡枝子,以及牛枝子的亲缘关系相对较近。
4.依据测序结果构建了胡枝子属部分种的系统发育树。测序得到了9种胡枝子的ITS序列和10种胡枝子的叶绿体trnL-F序列,同时结合GenBank上近缘物种的相关序列开展胡枝子属系统发育研究。序列分析结果表明,ITS序列中包含61个信息位点,占比对长度的10%,相比之下,trnL-F序列中仅有不到1%的信息位点。对两个片段分别采用MP和NJ法构建了系统发育树,结果表明,对单个片段而言,基于不同方法构建的系统发育树大体上一致,仅是在自展支持率上略有不同,并且NJ树中小支的关系比MP树清楚。对比两个片段构建的系统发育树,显示二者的拓扑结构大体相似,但是多花胡枝子的聚类位置明显有不同。在基于ITS序列的系统发育树中,多花胡枝子和大胡枝子组的物种聚到一起,而在基于tmL-F序列的系统发育树中,它又独立于大胡枝子组和胡枝子组外,单独为一支。
Lespedeza species are an important ecological and economic-type native shrub species in China.In recent years,it has received special attention on its strong adaptability and wide use.Compared with developed countries,utilization of this genus is still in a low level in China.To carry out genetic improvement for Lespedeza genus,it is necessary to unfold detailed researching in genetic background and phylogenetic relationship between them.In this study,24 provenances in Lespedeza bicolor and 10 species in Lespedeza were collected and regarded as tested materials.Population genetic structure of L.bicolor and relationship of some Lespedeza species were investigated using the methods of pollination biology,molecular markers and DNA sequencing.This study firstly developed the SSR-specific primers of Lespedeza genus by combining ISSR markers and chromosomal walking technique.The results are as follows:
1) Pollen viability,stigma receptivity and breeding system for L.bicolor were carried out.The optimum liquid medium for pollen germination of L.bicolor was screened and rapid staining method was found.The changes of pollen viability stored in different conditions suggested that the pollen vitality can last 7 days under 4℃and dry conditions can last 7 days.Results of hybrid index(OCI), pollen-ovule ratio(P/O) and artificial pollination indicated that the breeding system of L.bicolor belonged to outcrossing,part self-compatibility and pollinator-dependence.As an entomophilous plant, the high frequency of visits can significantly increase the seed setting rate of L.bicolor.
2) The SRAP reaction system was established and optimized.Senventeen pairs of primer combinations were selected from 484 sets of SRAP primer combinations.Taken 248 individuals as materials,a total of 464 bands were amplified by using 17 pairs of primer combinations,among them 360 bands were polymorphic;the percentage of polymorphic bands was 87.28%.At provenance level, average number of alleles and effective alleles on per locus was 1.881 and 1.568.The Nei's gene diversity index was 0.331,and Shannon's information index was 0.491.Genetic distance between species ranged from 0.1029 to 0.3434,and genetic identity of inter-provenance ranged from 0.709 to 0.902.The coefficient of genetic differentiation was 0.4157,which indicated that 58.4%variabilities occurred in provenance of L.bicolor.The gene flow among provenances(Nm) was 0.7028.According to the UPMGA cluster analysis based on the genetic distance,24 provenances of L.bicolor were divided into five groups.In accordance with cluster figure,10 kinds of Lespedeza were divided into three categories,while L.fomosa showed a higher independence.
3) Chromosomal walking and ISSR technologies were used in developing SSR primers.According to sequencing results of ISSR PCR amplification,6 pairs of newly designed primers were successfully applied to chromosomal walking.Of the 20 obtained SSR primers,10 primers showed high polymorphisms.All of them can amplify in other species of Lespedeza.At the same time,new SSR primers were utilized to the research of the relationship between 10 Lespedeza species.UPGMA cluster analyses based on Nei's genetic distance dividedthe 10 species into three main groups.
4) Phylogenetic trees of Lespedeza genus were constracted based on sequencing results.Nine ITS sequences and 10 trnL-F sequences were amplified and sequenced.The results of sequence analysis showed that ITS sequence had 61 information sites,accounting for more than 10 percent of the sequencing length.Compared with ITS sequencing results,trnL-F sequence only provided less than 1% information sites.Phylogeny trees of Maximum Parsimony(MP) and Neighbor-Joining(NJ) were constructed based on two fragments respectively.For single fragment,phylogenetic trees constructed by different methods were largely consistent and there was only slightly different in bootstrap values.As a whole,NJ tree reflected more clear than the MP tree in the relationships between subgroups. Comparison analysis phylogenetic tree showed that the tapnology of trees based on two fragments are almost identical,but the cluster location of L.floribunda was obvious differences,which clustered with L.bicolor and other Lespedeza species in phylogenetic tree based on ITS sequence,as contrast,it is independent in trnL-F phylogenetic tree.
引文
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