珍珠黄杨遗传多样性及其资源保护与利用
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摘要
珍珠黄杨(Buxus sinica var. parvifolia)为中国特有的珍稀濒危植物,属半岩生植物。本研究通过对其分布区的广泛调查,选择了7个天然群体和1个栽培群体,从形态性状和DNA水平,分析了珍珠黄杨群体的表型多样性和遗传多样性。从分子生态学、种群生态学、更新复壮及扦插繁殖角度系统研究了珍珠黄杨资源的保护与利用。结合表型标记和分子标记分析,探讨了珍珠黄杨相关类群的等级地位及命名。主要结论如下:
①珍珠黄杨种内存在丰富的群体间、群体内表型变异,其8个群体的枝叶等17个表型性状差异显著,变异系数较大。
②利用21个RAPD引物对8个珍珠黄杨群体进行遗传多样性分析,共扩增出176个(84.21 %)多态位点;21个ISSR引物共扩增出397个(91.89 %)多态位点。经POPGENE分析,Shannon信息指数在群体内为0.3039,占总遗传变异51.39 %;Gst是0.2723;Nm是1.3933(RAPD);Shannon信息指数在群体内为0.3368,占总遗传变异82.51 %;Gst是0.1627;Nm是2.5741(ISSR)。经AMOVA分析,群体内变异为62.65 %,Φst 0.554(RAPD);群体内变异达到77.65 %,Φst 0.223(ISSR)。基于Nei遗传距离对各群体进行聚类分析,截取λ=0.1653(RAPD)/0.0503(ISSR),可将不同地区的珍珠黄杨分为两大类:神农架山区的为一类,而华东片珍珠黄杨归为另一类。
③表型标记聚类结果表明:栽培黄杨的形态稳定一致,形态分化较小;尖叶黄杨与雀舌黄杨在形态上比较一致;珍珠黄杨各种群表型性状变异大。分子标记的UPGMA、NJ和PCA分析均支持华东片珍珠黄杨为独特的进化枝,这与其仅根据表型特征而定的分类单位相冲突,其它分类单位的聚类结果基本与传统分类结果一致。
④珍珠黄杨天然种群有2个死亡高峰:第一个死亡高峰在幼苗幼树阶段,第二个死亡高峰在各种群出现的时间及强度不同。其存活曲线介于Deevey-II型与Deevey-III型之间。
⑤在愈伤组织形成及根突出表皮时期,初始世代IAA,ABA和GA4水平与后继世代有明显差异并呈现一定规律。Fv,Fm,Fv/Fm,Fv/Fo,Fm/Fo这5个参数从初始世代到后继世代呈现一个明显地逐渐增大规律。荧光诱导动力学随继代次数的增加,从P相到T相的时间逐渐缩短,振荡衰减逐渐加快。
⑥春季扦插中,在愈伤组织产生前后及根突出表皮时,IAA,ABA和GA4含量的变化曲线在不同品种处理之间存在明显差异并呈现一定规律。夏季扦插中,以蛭石珍珠岩为扦插基质的插穗经外源激素尤其是IBA+BA处理后,生根效果明显提高;以轻壤土为扦插基质的则以IBA+BA和ZZHY处理的效果为好。
Buxus sinica var. parvifolia was a rare, endangered and endemic tree species in China, which belonged to semi-rock plants. Through extensive investigation of its natural distribution, 7 natural populations and 1 cultivated population were selected to analyse phenotypic diversity and genetic diversity of B. sinica var. parvifolia at the morphology and molecular level. From aspects of molecular ecology, population ecology, rejuvenation and cutting propagation, the conservation and utilization of B. sinica var. parvifolia resources was systematically studied. Combining phenotypic marker and molecular marker, the taxonomic status and naming of B. sinica var. parvifolia relevant groups were researched. Based on the above study, the conclusions were as follows:
①The phenotypic variations within populations or between populations of B. sinica var. parvifolia were rich, the differences of 17 branch and leaf characters in 8 populations were significant, the coefficients of variation were comparatively big.
②21 RAPD primers were selected for genetic diversity analysis of 8 B. sinica var. parvifolia populations which amplified 176 (84.21 %) polymorphic loci; 21 ISSR primers were selected for genetic diversity analysis which amplified 397 (91.89 %) polymorphic loci. Using POPGENE, Shannon's information index within population was 0.3039, which accounted for 51.39 % of total genetic variation; Gst was 0.2723; Nm was 1.3933 (RAPD); Shannon's information index within population was 0.3368, which accounted for 82.51 % of total genetic variation; Gst was 0.1627; Nm was 2.5741 (ISSR). Applying AMOVA, the variation within populations was 62.65 %,Φst was 0.554 (RAPD); variation within populations reached 77.65 %,Φst was 0.223 (ISSR). The dendrogram based on Nei’s genetic distance with interceptionλ= 0.1653 (RAPD) / 0.0503 (ISSR), 8 populations of B. sinica var. parvifolia from different regions could be divided into 2 categories: one was composed of populations from Shennongjia Moutain, one was composed of populations from B. sinica var. parvifolia in East China.
③The clustering results of phenotypic markers showed that: the shape of cultivated B. sinica was stable and consistent, and the differentiation between populations was less; B. sinica subsp. aemulans and B. bodinieri were relatively consistent in morphology; the variations of phenotypic characters of B. sinica var. parvifolia were big. All UPGMA, NJ and PCA strongly supported the monophyly of distinct clade within B. sinica var. parvifolia in East China that conflicted with the existing taxonomic unit assignment only based on morphological characteristics, and clustering results of the other taxa were basically consistent with classical taxonomy.
④The natural populations of B. sinica var. parvifolia had two peaks in mortality curve: the first mortality peak of population was during the stage of seedling and sapling, and the second mortality peak in different populations varied in time and intensity. The survival curves of populations were between Deevey-II type and Deevey-III type.
⑤Around the time of callus forming and the root projected epidermis, the levels of IAA, ABA and GA4 between the initial generation and the subsequent generation had obvious differences, and presented certain rules. Fv, Fm, Fv / Fm, Fv / Fo, Fm / Fo these five parameters from the initial generation to the subsequent generation showed a gradually increasing tendency. The time from T phase to P phase of fluorescence induction kinetics was progressive shortening with the continuous generation cuttage proceeding, decrement of oscillation was gradually accelerated.
⑥In spring cutting rooting process, around the time of callus forming and the root projected epidermis, the levels of IAA, ABA and GA4 in different cultivars and treatments had obvious differences, and presented certain rules. In summer cutting rooting process, the rooting effects of cuttings in matrix composed of vermiculite and perlite treated by exogenous hormone especially IBA + BA were increased; in cutting matrix of light loam, hormone treatment effects of IBA + BA and ZZHY were comparative good.
①珍珠黄杨种内存在丰富的群体间、群体内表型变异,其8个群体的枝叶等17个表型性状差异显著,变异系数较大。
②利用21个RAPD引物对8个珍珠黄杨群体进行遗传多样性分析,共扩增出176个(84.21 %)多态位点;21个ISSR引物共扩增出397个(91.89 %)多态位点。经POPGENE分析,Shannon信息指数在群体内为0.3039,占总遗传变异51.39 %;Gst是0.2723;Nm是1.3933(RAPD);Shannon信息指数在群体内为0.3368,占总遗传变异82.51 %;Gst是0.1627;Nm是2.5741(ISSR)。经AMOVA分析,群体内变异为62.65 %,Φst 0.554(RAPD);群体内变异达到77.65 %,Φst 0.223(ISSR)。基于Nei遗传距离对各群体进行聚类分析,截取λ=0.1653(RAPD)/0.0503(ISSR),可将不同地区的珍珠黄杨分为两大类:神农架山区的为一类,而华东片珍珠黄杨归为另一类。
③表型标记聚类结果表明:栽培黄杨的形态稳定一致,形态分化较小;尖叶黄杨与雀舌黄杨在形态上比较一致;珍珠黄杨各种群表型性状变异大。分子标记的UPGMA、NJ和PCA分析均支持华东片珍珠黄杨为独特的进化枝,这与其仅根据表型特征而定的分类单位相冲突,其它分类单位的聚类结果基本与传统分类结果一致。
④珍珠黄杨天然种群有2个死亡高峰:第一个死亡高峰在幼苗幼树阶段,第二个死亡高峰在各种群出现的时间及强度不同。其存活曲线介于Deevey-II型与Deevey-III型之间。
⑤在愈伤组织形成及根突出表皮时期,初始世代IAA,ABA和GA4水平与后继世代有明显差异并呈现一定规律。Fv,Fm,Fv/Fm,Fv/Fo,Fm/Fo这5个参数从初始世代到后继世代呈现一个明显地逐渐增大规律。荧光诱导动力学随继代次数的增加,从P相到T相的时间逐渐缩短,振荡衰减逐渐加快。
⑥春季扦插中,在愈伤组织产生前后及根突出表皮时,IAA,ABA和GA4含量的变化曲线在不同品种处理之间存在明显差异并呈现一定规律。夏季扦插中,以蛭石珍珠岩为扦插基质的插穗经外源激素尤其是IBA+BA处理后,生根效果明显提高;以轻壤土为扦插基质的则以IBA+BA和ZZHY处理的效果为好。
Buxus sinica var. parvifolia was a rare, endangered and endemic tree species in China, which belonged to semi-rock plants. Through extensive investigation of its natural distribution, 7 natural populations and 1 cultivated population were selected to analyse phenotypic diversity and genetic diversity of B. sinica var. parvifolia at the morphology and molecular level. From aspects of molecular ecology, population ecology, rejuvenation and cutting propagation, the conservation and utilization of B. sinica var. parvifolia resources was systematically studied. Combining phenotypic marker and molecular marker, the taxonomic status and naming of B. sinica var. parvifolia relevant groups were researched. Based on the above study, the conclusions were as follows:
①The phenotypic variations within populations or between populations of B. sinica var. parvifolia were rich, the differences of 17 branch and leaf characters in 8 populations were significant, the coefficients of variation were comparatively big.
②21 RAPD primers were selected for genetic diversity analysis of 8 B. sinica var. parvifolia populations which amplified 176 (84.21 %) polymorphic loci; 21 ISSR primers were selected for genetic diversity analysis which amplified 397 (91.89 %) polymorphic loci. Using POPGENE, Shannon's information index within population was 0.3039, which accounted for 51.39 % of total genetic variation; Gst was 0.2723; Nm was 1.3933 (RAPD); Shannon's information index within population was 0.3368, which accounted for 82.51 % of total genetic variation; Gst was 0.1627; Nm was 2.5741 (ISSR). Applying AMOVA, the variation within populations was 62.65 %,Φst was 0.554 (RAPD); variation within populations reached 77.65 %,Φst was 0.223 (ISSR). The dendrogram based on Nei’s genetic distance with interceptionλ= 0.1653 (RAPD) / 0.0503 (ISSR), 8 populations of B. sinica var. parvifolia from different regions could be divided into 2 categories: one was composed of populations from Shennongjia Moutain, one was composed of populations from B. sinica var. parvifolia in East China.
③The clustering results of phenotypic markers showed that: the shape of cultivated B. sinica was stable and consistent, and the differentiation between populations was less; B. sinica subsp. aemulans and B. bodinieri were relatively consistent in morphology; the variations of phenotypic characters of B. sinica var. parvifolia were big. All UPGMA, NJ and PCA strongly supported the monophyly of distinct clade within B. sinica var. parvifolia in East China that conflicted with the existing taxonomic unit assignment only based on morphological characteristics, and clustering results of the other taxa were basically consistent with classical taxonomy.
④The natural populations of B. sinica var. parvifolia had two peaks in mortality curve: the first mortality peak of population was during the stage of seedling and sapling, and the second mortality peak in different populations varied in time and intensity. The survival curves of populations were between Deevey-II type and Deevey-III type.
⑤Around the time of callus forming and the root projected epidermis, the levels of IAA, ABA and GA4 between the initial generation and the subsequent generation had obvious differences, and presented certain rules. Fv, Fm, Fv / Fm, Fv / Fo, Fm / Fo these five parameters from the initial generation to the subsequent generation showed a gradually increasing tendency. The time from T phase to P phase of fluorescence induction kinetics was progressive shortening with the continuous generation cuttage proceeding, decrement of oscillation was gradually accelerated.
⑥In spring cutting rooting process, around the time of callus forming and the root projected epidermis, the levels of IAA, ABA and GA4 in different cultivars and treatments had obvious differences, and presented certain rules. In summer cutting rooting process, the rooting effects of cuttings in matrix composed of vermiculite and perlite treated by exogenous hormone especially IBA + BA were increased; in cutting matrix of light loam, hormone treatment effects of IBA + BA and ZZHY were comparative good.
引文
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