利用SSR研究榛属种间亲缘关系及平榛居群遗传多样性
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摘要
为了探讨榛属(Corylus)植物的遗传变异规律,为榛子培育提供理论基础与技术支持,本试验选取了榛属7个(变)种及平榛(C.heterophylla Fisch) 14个野生居群为研究对象,通过SSR分子标记,对榛属种间亲缘关系及平榛居群遗传多样性进行了分析。研究结果如下:
(1)建立和优化了榛属植物的SSR反应体系;筛选了29对欧榛的SSR引物对榛属植物进行扩增,有效性达100 %。利用其中的17对引物对榛属种间亲缘关系进行了研究,在43份种质中共检测到137个等位基因,平均每个位点可检测到等位基因8.0588个;观测杂合度(Ho)和预期杂合度(He)平均值分别为0.5773和0.7855。基于遗传距离的聚类可初步将7种分为3组:平榛(C.heterophylla Fisch)、榛(C.avellana L.)及川榛(C.kweichowensis Hu)为1组;臧刺榛[(C.feroxWall.var.thibetica (Batal.Franch.)]、滇榛(C.yunnanensis A.Camus)及华榛(C.chinensis Franch)为1组;毛榛(C.mandshurica Maxim.et Rupr.)为1组,从分子水平上对榛属种间亲缘关系给予了鉴定。平榛和欧榛的遗传多样性较高,且亲缘关系较近,为已成功的种间远缘杂交提供了理论支持。
(2)利用Sanger末端终止法测序原理,在位点CAC-B014对榛属7种及虎榛属(Ostryopsis Decne)虎榛(O. davidiana Decne)的微卫星序列进行了测序,结果表明:欧榛微卫星在榛属种间及虎榛属间具有高度的同源性及多态性,可用于近缘种研究,为榛子种质资源的进一步研究提供了依据。
(3)利用样方调查和SSR分子标记相结合,初步对平榛克隆结构及个体进行了研究。平榛克隆繁殖比例较大,成集群分布,可分3种类型:株丛、株系及群系。研究结果显示同1株丛(系)的个体基因型相同,为1个独立的遗传个体;同时存在一定比例的种子繁殖使平榛具有较高的遗传多样性(Na=5.1250,Ne =2.4996,Ho=0.7148,He=0.5474)。居群水平取样时要考虑平榛兼性繁殖特性,根据居群大小、聚集程度、榛丛斑块大小及斑块距离进行适当调整,确定具体样本数。
(4) 25对引物共检测出239个等位基因变异,位点拥有的等位基因数量(Na)在2~19个之间,平均每个位点可检测出9.56个等位基因;有效等位基因数(Ne)1.209~10.889个,平均值为4.78个;观察杂合度(Ho)变化范围为0.191~0.921,平均值为0.552;期望杂合度(He)的变化范围为0.174~0.911,平均值为0.729。平榛物种水平具有较高的遗传多样性,表明其生态适应性有一定差异,这为平榛杂交育种、无性系良种选育提供了理论基础。
(5)平榛在居群水平上,各个居群平均等位基因数(Na)为3.57个,有效等位基因数(Ne)为2.68个,观察杂合度(Ho)和期望杂合度(He)分别0.553和0.582。河南辉县(HX)居群的遗传多样性水平最高(Na=4.76,Ne=3.41,Ho=0.608,He=0.685),河北宽城(KC)居群的遗传多样性水平最低(Na=1.44,Ne=1.44,Ho=0.44,He=0.251)。遗传分化系数(Fst)为0.2485,即有24.85 %的遗传变异存在居群之间,75.15 %的遗传变异存在于居群内。分子水平方差分析表明,居群间及居群内差异均达到极显著水平。
(6)利用算术加权平均数法(UPGMA)将14个居群分成4大类,地理位置较近的聚在一起;居群间的基因流较低,为0.756。经Mantel检测,居群间的遗传距离和地理距离之间没有显著的相关关系(r =0.5144,P=0.9989)。
7 species (variety) of Corylus about 43 accessions and 14 C. heterophylla Fisch populations were selected and investigated in this study. The phylogenetic relationship among 7 species of Corylus was explored and genetic diversity of C.heterophyllaa populations was evaluated using SSR markers. The obtained results and conclusions were as follows:
(1) The SSR reaction system was established and optimized; 29 pairs of SSR loci of C.avellana L. were selected. A total number of 137 alleles were amplified using 17 SSR primers. The average value of observed allele was 8.0588; The average value of effective allele was 5.0259; The mean of Ho and He was 0.5773 and 0.7855, respectively; UPGMA cluster analysis based on Nei’s genetic distance divided the 7 species (variety) into three main groups: C.heterophylla Fisch, C.avellana L. and C.kweichowensis Hu; C.feroxWall.var.thibetica (Batal.Franch), C.yunnanensis A.Camus and C.chinensis Franch; C. mandshurica Maxim.et Rupr. The genetic diversity of C. heterophylla Fisch and C.avellana L. were higher than others and they were phylogenetically closer.
(2) The PCR products were directly seqnenced using Sanger method and DNA sequence analyzer (Beckman 2000). The result showed SSR has highly homology among Corylus, O.davidiana Decne and the primers of SSR could be used to analyze relatives.
(3) The C.heterophylla Fisch could reproduce both sexually and vegetativly. According to the methods of samLpes investigation and SSR marker, the vegetative reproduction was main. It clusters. The clusters have three main forms: single cluster, individual clone and mulriple clusters. Seed reproduction was beneficial to it’s high genetic diversity(Na=5.1250,Ne =2.4996,Ho=0.7148,He=0.5474). The appropriate sampling method should be designed on the basis of distribution on species level and the population size, the degree of the mass, cluster size and distance on population level.
(4) 25 pairs of primers were chosen for the analysis of 14 wild C. heterophylla Fisch Populationss. 239 alleles was presented at 25 SSR loci.Observed number of alleles per locus ranged from 2 to 19 and average value of allele (Na) was 9.56; Effective number of alleles (Ne) was from 1.209 to 10.889, averagely 4.78; The expected total heterozygosity (He) ranged from 0.174 to 0.911 and mean was 0.729; The observed total heterozygosity (Ho) ranged from 0.191 to 0.921, mean was 0.552.
(5) On the population level, Na =3.57, Ne=2.68, Ho =0.553, He=0.582. The genetic diversity of Henan province Hui county was most (Na=4.76,Ne=3.41,Ho=0.608,He=0.685) and Hebei province Kuancheng county’s was the least(Na=1.44,Ne=1.44,Ho=0.44,He=0.251). A low level of genetic differentiation among populations was detected based on Nei’s genetic diversity analysis (Fst=0.2485) and main variations was within population (75.15 %). A significantly high degree of genetic differentiation occurred among populations.
(6) UPGMA cluster analysis based on Nei’s genetic distance divided the 14 population into four main groups; Gene flow was low (0.756). Additionally, Nei’s unbiased genetic distance matrix compared with a corresponding geographic distance matrix showed the two matrices were not significantly correlated (r =0.5144, P=0.9989).
(1)建立和优化了榛属植物的SSR反应体系;筛选了29对欧榛的SSR引物对榛属植物进行扩增,有效性达100 %。利用其中的17对引物对榛属种间亲缘关系进行了研究,在43份种质中共检测到137个等位基因,平均每个位点可检测到等位基因8.0588个;观测杂合度(Ho)和预期杂合度(He)平均值分别为0.5773和0.7855。基于遗传距离的聚类可初步将7种分为3组:平榛(C.heterophylla Fisch)、榛(C.avellana L.)及川榛(C.kweichowensis Hu)为1组;臧刺榛[(C.feroxWall.var.thibetica (Batal.Franch.)]、滇榛(C.yunnanensis A.Camus)及华榛(C.chinensis Franch)为1组;毛榛(C.mandshurica Maxim.et Rupr.)为1组,从分子水平上对榛属种间亲缘关系给予了鉴定。平榛和欧榛的遗传多样性较高,且亲缘关系较近,为已成功的种间远缘杂交提供了理论支持。
(2)利用Sanger末端终止法测序原理,在位点CAC-B014对榛属7种及虎榛属(Ostryopsis Decne)虎榛(O. davidiana Decne)的微卫星序列进行了测序,结果表明:欧榛微卫星在榛属种间及虎榛属间具有高度的同源性及多态性,可用于近缘种研究,为榛子种质资源的进一步研究提供了依据。
(3)利用样方调查和SSR分子标记相结合,初步对平榛克隆结构及个体进行了研究。平榛克隆繁殖比例较大,成集群分布,可分3种类型:株丛、株系及群系。研究结果显示同1株丛(系)的个体基因型相同,为1个独立的遗传个体;同时存在一定比例的种子繁殖使平榛具有较高的遗传多样性(Na=5.1250,Ne =2.4996,Ho=0.7148,He=0.5474)。居群水平取样时要考虑平榛兼性繁殖特性,根据居群大小、聚集程度、榛丛斑块大小及斑块距离进行适当调整,确定具体样本数。
(4) 25对引物共检测出239个等位基因变异,位点拥有的等位基因数量(Na)在2~19个之间,平均每个位点可检测出9.56个等位基因;有效等位基因数(Ne)1.209~10.889个,平均值为4.78个;观察杂合度(Ho)变化范围为0.191~0.921,平均值为0.552;期望杂合度(He)的变化范围为0.174~0.911,平均值为0.729。平榛物种水平具有较高的遗传多样性,表明其生态适应性有一定差异,这为平榛杂交育种、无性系良种选育提供了理论基础。
(5)平榛在居群水平上,各个居群平均等位基因数(Na)为3.57个,有效等位基因数(Ne)为2.68个,观察杂合度(Ho)和期望杂合度(He)分别0.553和0.582。河南辉县(HX)居群的遗传多样性水平最高(Na=4.76,Ne=3.41,Ho=0.608,He=0.685),河北宽城(KC)居群的遗传多样性水平最低(Na=1.44,Ne=1.44,Ho=0.44,He=0.251)。遗传分化系数(Fst)为0.2485,即有24.85 %的遗传变异存在居群之间,75.15 %的遗传变异存在于居群内。分子水平方差分析表明,居群间及居群内差异均达到极显著水平。
(6)利用算术加权平均数法(UPGMA)将14个居群分成4大类,地理位置较近的聚在一起;居群间的基因流较低,为0.756。经Mantel检测,居群间的遗传距离和地理距离之间没有显著的相关关系(r =0.5144,P=0.9989)。
7 species (variety) of Corylus about 43 accessions and 14 C. heterophylla Fisch populations were selected and investigated in this study. The phylogenetic relationship among 7 species of Corylus was explored and genetic diversity of C.heterophyllaa populations was evaluated using SSR markers. The obtained results and conclusions were as follows:
(1) The SSR reaction system was established and optimized; 29 pairs of SSR loci of C.avellana L. were selected. A total number of 137 alleles were amplified using 17 SSR primers. The average value of observed allele was 8.0588; The average value of effective allele was 5.0259; The mean of Ho and He was 0.5773 and 0.7855, respectively; UPGMA cluster analysis based on Nei’s genetic distance divided the 7 species (variety) into three main groups: C.heterophylla Fisch, C.avellana L. and C.kweichowensis Hu; C.feroxWall.var.thibetica (Batal.Franch), C.yunnanensis A.Camus and C.chinensis Franch; C. mandshurica Maxim.et Rupr. The genetic diversity of C. heterophylla Fisch and C.avellana L. were higher than others and they were phylogenetically closer.
(2) The PCR products were directly seqnenced using Sanger method and DNA sequence analyzer (Beckman 2000). The result showed SSR has highly homology among Corylus, O.davidiana Decne and the primers of SSR could be used to analyze relatives.
(3) The C.heterophylla Fisch could reproduce both sexually and vegetativly. According to the methods of samLpes investigation and SSR marker, the vegetative reproduction was main. It clusters. The clusters have three main forms: single cluster, individual clone and mulriple clusters. Seed reproduction was beneficial to it’s high genetic diversity(Na=5.1250,Ne =2.4996,Ho=0.7148,He=0.5474). The appropriate sampling method should be designed on the basis of distribution on species level and the population size, the degree of the mass, cluster size and distance on population level.
(4) 25 pairs of primers were chosen for the analysis of 14 wild C. heterophylla Fisch Populationss. 239 alleles was presented at 25 SSR loci.Observed number of alleles per locus ranged from 2 to 19 and average value of allele (Na) was 9.56; Effective number of alleles (Ne) was from 1.209 to 10.889, averagely 4.78; The expected total heterozygosity (He) ranged from 0.174 to 0.911 and mean was 0.729; The observed total heterozygosity (Ho) ranged from 0.191 to 0.921, mean was 0.552.
(5) On the population level, Na =3.57, Ne=2.68, Ho =0.553, He=0.582. The genetic diversity of Henan province Hui county was most (Na=4.76,Ne=3.41,Ho=0.608,He=0.685) and Hebei province Kuancheng county’s was the least(Na=1.44,Ne=1.44,Ho=0.44,He=0.251). A low level of genetic differentiation among populations was detected based on Nei’s genetic diversity analysis (Fst=0.2485) and main variations was within population (75.15 %). A significantly high degree of genetic differentiation occurred among populations.
(6) UPGMA cluster analysis based on Nei’s genetic distance divided the 14 population into four main groups; Gene flow was low (0.756). Additionally, Nei’s unbiased genetic distance matrix compared with a corresponding geographic distance matrix showed the two matrices were not significantly correlated (r =0.5144, P=0.9989).
引文
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