川黔地区百合属植物资源表型多样性及孢粉学研究
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摘要
百合(Lilium spp)是单子叶植物亚纲百合科(Liliaceae)百合属(Lilium)植物的总称,为多年生露地鳞茎草本植物。我国是百合属植物的自然分布中心,以资源丰富、特有种多、分布广等特点著称,在世界百合品种改良中占有十分重要的地位。但目前很多野生百合资源没有得到充分利用。针对这一问题,作者对川黔地区百合分布集中的区域进行了野生百合种质资源调查、收集,并将采集到的野生百合进行了表型多样性分析和孢粉学研究。主要研究结果如下：
1.在调查中共收集到川黔地区野生百合7个种2个变种,即沪定百合(L.sargentiae)、卷丹(L. lancifolium)、野百合(L. brownii)、川百合(L. davidii)、宝兴百合(L. duchartrei)、岷江百合(L. regale)、紫脊百合(L. leucanthum)、黄绿花滇百合(L. bakerianum)和湖北百合(L. henryi)。采用样地调查的方法,了解其资源分布情况、生物学特性、观赏性状、生境条件和植物群落特点等。调查发现,川黔地区野生百合资源并没有相关文献记载的丰富,且破坏严重,亟待保护。
2.以收集到的7个种2个变种为研究对象,对其50个表型性状进行多样性分析和主成分分析,并对试验材料进行聚类分析。表型性状多样性分析结果表明:川黔地区野生百合种质资源的主要性状有明显差异。各形态性状的变异系数除了子房宽相对较小外,其他性状的变异系数都大于20%,其中下叶型的变异系数高达65.10%。同时,各性状遗传多样性指数也较高,植株高度的多样性指数达到2.02。
主成分分析表明,各性状间相关性强,提取了4个主成分,占信息总量的69.595%,其中第1主成分所占信息量就达43.656%。4个主成分共提取26个形态性状指标,其中第1主成分中花柱长、外轮长、花丝长、内轮长、子房长、有无花斑、内轮宽、花丝宽和柱头宽等性状均表现较高的正向载荷量,全部超过0.75,表明花性状是川黔地区野生百合形态分化的主要指标。
聚类分析结果将25份材料聚为2大类,准确地将各材料归入遗传基础、演化路线不同的百合组和卷瓣组。2大类群主要性状表现为植株高矮、花朵大小、叶片大小等性状的差异。
3.用扫描电镜观察所采集到的野生百合及课题组之前搜集的2个待定材料的花粉形态,百合花粉粒均呈两侧对称,极面观为长椭圆形,赤道面观为舟形,大小为68.87-120.65μm×36.02-53.70μm,种间变化较大。萌发沟为远极单沟,沟长几达两端,沟缘较整齐,沟裂两极较尖,种间差异很小。外壁表面具网状纹饰,网眼形状为不规则多边形至近圆形,在赤道面中部最大,向两极和边缘逐渐变小,网眼内有瘤状或疣状突起,网眼大小为12.11-50.78μm2,种间差异较大。网脊宽1.96-3.63μm,其颗粒形态为瘤状或盘珠状,单排或单双排基柱。
以花粉粒大小(赤道轴长、极轴长),P/E,网脊宽和纹饰网眼大小等5个形态指标对9个种和变种及2个待定材料进行聚类分析,结果表明：百合组的泸定百合、紫脊百合、野百合、岷江百合和待定1号聚为一类群,卷瓣组的卷丹、湖北百合、宝兴百合、川百合、待定2号和黄绿花滇百合为另一类群。这两个类群的聚类结果与形态性状分类相一致。泸定百合、紫脊百合和野百合的亲缘关系近,卷丹、湖北百合、宝兴百合和川百合亲缘关系近,而待定1号与岷江百合、待定2号与川百合亲缘关系近。
对不同产地泸定百合(BH1-BH13)的P值、E值及P／E值进行方差分析,结果表明其形态性状差异均达到极显著水平。
Lily is one of the world's most important cut flowers. China is the natural distribution center, rich in natural resources and endemic species. But at present, resources of many wild lilies are not fully utilized. To solve this problem, in this study, germplasm collection and studies on phenotype and pollen diversity were conducted for wild lily in Sichuan-Guizhou region, a lily concentrated region in China. The study was aimed at the collection and preservation of lily resources, evaluating the genetic relationships of some wild species. The main results were as follows.
1. In the survey and collection 7 species and two varieties of wild lily were collected, including L.sargentiae, L.lancifolium, L.brownii, L.davidii, L.duchartrei, L.regale, L.leucanthum, L.bakerianum and L. henryi. Plot survey method was used to investigate the distribution, biological characteristics, ornamental traits, habitat conditions and vegetation characteristics. The survey showed that wild lilies in Sichuan-Guizhou area were not as many as recorded in the relevant literature, seriously damaged and in urgent need of protection.
2. Seven species and two varieties were collected for diversity analysis, principal component analysis and cluster analysis.50 phenotypic traits such as plant height, petal length, leaf length were studied. The results showed that the germplasm resources of wild lily in Sichuan-Guizhou area were significantly different in main characters. The coefficients of variation of morphological traits were more than 20%, except the width of the ovary, with coefficient of variation of bottom leaf sharp up to 276.39%.
Principal component analysis showed a strong correlation between the traits. Four extracted principal components accounted for 69.595% of the total information, principal component 1 accounting for up to 43.656% of the total information. Four principal components extracted a total of 26 morphological traits and principal component 1, including style length, outer petal length, filament length, inner petal length, ovary length, whether or piebald, inner petal width, filament width, stigma width, showed a highly positive load capacity, all more than 0.75, indicating that flower traits were the main indicators of morphological differentiation in wild lilies in Sichuan-Guizhou area.
25 materials were clustered into two major categories, in agreement with the genetic basis and evolution of different lily group. The characters of the two categories mainly differred in plant height, flower size and leaf size.
3. Scanning electron microscopy was used to observe the pollen morphology of the collected wild lilies and the two unidentified materials collected before. It was found that the pollen of lilium was single-form, bilaterally symmetrical, polar view oblong to elliptic, the edge with a depression (as germination groove), boat-shaped in equatorial view, with a single germination ditch, ditch extended to slender poles, trench edge was neat, polar crack tip groove. Surface ornamentation was reticulate, mesh for irregular polygon to nearly round, with different sizes. Net ridge morphology was in particle-and tumor-like shape or disk beads, arranged closely, a single row of pillars, ridge width 1.96-3.63μm, sometimes breakpoint. There were tumor-like or verrucous apophysises in the meshes, and the number varied with the species.
Cluster analysis was carried out using 5 morphology indexes including pollen size (equatorial axis, polar axis length), P/E, net ridge width, decoration mesh size. All materials were first clustered into two classes:L. sargentiae, L.leucanthum, L. brownii, L. regale and unidentified 1 were in group 1, L. lancifolium, L. henryi, L. duchartrei, L. davidii, unidentified 2 and L. bakerianum were in another group. The clustering result was consistent with the morphological traits. The results showed that L. sargentiae, L. leucanthum and L. brownii were closely related, while L. lancifolium, L. henryi, L. duchartrei and L. regale shareed close genetic relationships. Unidentified 1 was close to L. regale, while unidentified 2 was close to L. davidii.
Variance analysis according to P values, E values and P/E values of L. sargentiae from different places (BH1-BH13) showed that the morphologic differences were very significant.
1.在调查中共收集到川黔地区野生百合7个种2个变种,即沪定百合(L.sargentiae)、卷丹(L. lancifolium)、野百合(L. brownii)、川百合(L. davidii)、宝兴百合(L. duchartrei)、岷江百合(L. regale)、紫脊百合(L. leucanthum)、黄绿花滇百合(L. bakerianum)和湖北百合(L. henryi)。采用样地调查的方法,了解其资源分布情况、生物学特性、观赏性状、生境条件和植物群落特点等。调查发现,川黔地区野生百合资源并没有相关文献记载的丰富,且破坏严重,亟待保护。
2.以收集到的7个种2个变种为研究对象,对其50个表型性状进行多样性分析和主成分分析,并对试验材料进行聚类分析。表型性状多样性分析结果表明:川黔地区野生百合种质资源的主要性状有明显差异。各形态性状的变异系数除了子房宽相对较小外,其他性状的变异系数都大于20%,其中下叶型的变异系数高达65.10%。同时,各性状遗传多样性指数也较高,植株高度的多样性指数达到2.02。
主成分分析表明,各性状间相关性强,提取了4个主成分,占信息总量的69.595%,其中第1主成分所占信息量就达43.656%。4个主成分共提取26个形态性状指标,其中第1主成分中花柱长、外轮长、花丝长、内轮长、子房长、有无花斑、内轮宽、花丝宽和柱头宽等性状均表现较高的正向载荷量,全部超过0.75,表明花性状是川黔地区野生百合形态分化的主要指标。
聚类分析结果将25份材料聚为2大类,准确地将各材料归入遗传基础、演化路线不同的百合组和卷瓣组。2大类群主要性状表现为植株高矮、花朵大小、叶片大小等性状的差异。
3.用扫描电镜观察所采集到的野生百合及课题组之前搜集的2个待定材料的花粉形态,百合花粉粒均呈两侧对称,极面观为长椭圆形,赤道面观为舟形,大小为68.87-120.65μm×36.02-53.70μm,种间变化较大。萌发沟为远极单沟,沟长几达两端,沟缘较整齐,沟裂两极较尖,种间差异很小。外壁表面具网状纹饰,网眼形状为不规则多边形至近圆形,在赤道面中部最大,向两极和边缘逐渐变小,网眼内有瘤状或疣状突起,网眼大小为12.11-50.78μm2,种间差异较大。网脊宽1.96-3.63μm,其颗粒形态为瘤状或盘珠状,单排或单双排基柱。
以花粉粒大小(赤道轴长、极轴长),P/E,网脊宽和纹饰网眼大小等5个形态指标对9个种和变种及2个待定材料进行聚类分析,结果表明：百合组的泸定百合、紫脊百合、野百合、岷江百合和待定1号聚为一类群,卷瓣组的卷丹、湖北百合、宝兴百合、川百合、待定2号和黄绿花滇百合为另一类群。这两个类群的聚类结果与形态性状分类相一致。泸定百合、紫脊百合和野百合的亲缘关系近,卷丹、湖北百合、宝兴百合和川百合亲缘关系近,而待定1号与岷江百合、待定2号与川百合亲缘关系近。
对不同产地泸定百合(BH1-BH13)的P值、E值及P／E值进行方差分析,结果表明其形态性状差异均达到极显著水平。
Lily is one of the world's most important cut flowers. China is the natural distribution center, rich in natural resources and endemic species. But at present, resources of many wild lilies are not fully utilized. To solve this problem, in this study, germplasm collection and studies on phenotype and pollen diversity were conducted for wild lily in Sichuan-Guizhou region, a lily concentrated region in China. The study was aimed at the collection and preservation of lily resources, evaluating the genetic relationships of some wild species. The main results were as follows.
1. In the survey and collection 7 species and two varieties of wild lily were collected, including L.sargentiae, L.lancifolium, L.brownii, L.davidii, L.duchartrei, L.regale, L.leucanthum, L.bakerianum and L. henryi. Plot survey method was used to investigate the distribution, biological characteristics, ornamental traits, habitat conditions and vegetation characteristics. The survey showed that wild lilies in Sichuan-Guizhou area were not as many as recorded in the relevant literature, seriously damaged and in urgent need of protection.
2. Seven species and two varieties were collected for diversity analysis, principal component analysis and cluster analysis.50 phenotypic traits such as plant height, petal length, leaf length were studied. The results showed that the germplasm resources of wild lily in Sichuan-Guizhou area were significantly different in main characters. The coefficients of variation of morphological traits were more than 20%, except the width of the ovary, with coefficient of variation of bottom leaf sharp up to 276.39%.
Principal component analysis showed a strong correlation between the traits. Four extracted principal components accounted for 69.595% of the total information, principal component 1 accounting for up to 43.656% of the total information. Four principal components extracted a total of 26 morphological traits and principal component 1, including style length, outer petal length, filament length, inner petal length, ovary length, whether or piebald, inner petal width, filament width, stigma width, showed a highly positive load capacity, all more than 0.75, indicating that flower traits were the main indicators of morphological differentiation in wild lilies in Sichuan-Guizhou area.
25 materials were clustered into two major categories, in agreement with the genetic basis and evolution of different lily group. The characters of the two categories mainly differred in plant height, flower size and leaf size.
3. Scanning electron microscopy was used to observe the pollen morphology of the collected wild lilies and the two unidentified materials collected before. It was found that the pollen of lilium was single-form, bilaterally symmetrical, polar view oblong to elliptic, the edge with a depression (as germination groove), boat-shaped in equatorial view, with a single germination ditch, ditch extended to slender poles, trench edge was neat, polar crack tip groove. Surface ornamentation was reticulate, mesh for irregular polygon to nearly round, with different sizes. Net ridge morphology was in particle-and tumor-like shape or disk beads, arranged closely, a single row of pillars, ridge width 1.96-3.63μm, sometimes breakpoint. There were tumor-like or verrucous apophysises in the meshes, and the number varied with the species.
Cluster analysis was carried out using 5 morphology indexes including pollen size (equatorial axis, polar axis length), P/E, net ridge width, decoration mesh size. All materials were first clustered into two classes:L. sargentiae, L.leucanthum, L. brownii, L. regale and unidentified 1 were in group 1, L. lancifolium, L. henryi, L. duchartrei, L. davidii, unidentified 2 and L. bakerianum were in another group. The clustering result was consistent with the morphological traits. The results showed that L. sargentiae, L. leucanthum and L. brownii were closely related, while L. lancifolium, L. henryi, L. duchartrei and L. regale shareed close genetic relationships. Unidentified 1 was close to L. regale, while unidentified 2 was close to L. davidii.
Variance analysis according to P values, E values and P/E values of L. sargentiae from different places (BH1-BH13) showed that the morphologic differences were very significant.
引文
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