木本能源植物黄连木单株选择、类型划分与群落调查研究
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摘要
本研究面向生产,以黄连木为研究对象,以高产能源基地建设为目标,立足于充分挖掘和利用现存的遗传变异,从生物学、育种学、生态学和分子生物学多个角度,对黄连木进行个体、群体和群落多层次的研究,为黄连木进一步开发利用提供理论依据。主要研究结果如下:
(1)通过黄连木优树选择,初步制定了黄连木的选优方法和标准,在各集中分布区初选优树228株。在对主要性状分析的基础上,选择出含油率高、抗虫性强、产量高的各种类型优良单株,为进一步遗传改良奠定基础,也为将来建立黄连木种质资源库提供材料。
(2)根据黄连木作为能源树种的特点,遵照指标体系的制定原则,提出了由目标层、控制层和指标层组成的黄连木优良性状综合评价指标体系。同时应用层次分析法、特尔斐法、指数综合评价法确定了各评价指标的权重和评价方法,为黄连木优良性状的划分和评选提供理论依据。
(3)结合黄连木优良性状综合评价指标体系,进行多性状综合选优分析,选择出高产、高含油率、抗黄连小蜂综合性状优良的单株68株。在优树选择的基础上,进行多地点黄连木优树苗期测定,探讨苗期各生长性状之间的内在关系和遗传变异,进行遗传参数估算和优良家系的评选,并运用典范相关分析的方法对亲代和子代性状的关系进行分析,构造指数方程,对参试的黄连木优树进行分类复选。
(4)在全面调查分析的基础上,选用变异较为明显的七个性状,将黄连木划分为各种不同类型。通过系统分析,确定了三级分类依据,进行多性状综合分类,初步评选出一些品质好、产量高,抗性强的优良类型。同时提出成年树雌雄株形态鉴别的方法,以方便生产单位使用。
(5)对产量的主要影响因素的分析表明,黄连木果实产量是一个综合数量性状,受多因子的影响,其中虫果率、土壤肥力、果穗粒数对单株产量影响最为明显。加强病虫害防治,增加土壤肥力,进行树体管护,促进果穗结实是提高黄连木产量的关键环节。
(6)以11个黄连木天然群体为试材,对18个表型性状的变异与多样性的分析表明,黄连木表型性状存在极其丰富的变异,各性状变异系数相差较大,变幅在4.249%- 27.31%之间,果实性状的变异系数最小,受较高的遗传控制,群体间平均表型分化系数VST为22.27%,说明群体内变异是黄连木变异的主要来源,聚类分析把11个群体分为4类。不同生境黄连木表型性状变异研究表明,坡向和海拨对多数性状有明显的影响,而坡位的作用较不明显,同时用空间代时间的方法,分析了不同年龄黄连木表型性状的变异。
(7)采用核磁共振(NMR)技术,对黄连木分布区9个群体果实含油率进行取样测定,结果表明:果实、种子和果肉含油率分别为36.183%、26.594%、50.503%,不同群体间差异显著,群体平均表型分化系数达到92.08%。系统聚类分析将黄连木群体分为低变异中等含油率群体、低变异低含油率群体、高变异高含油群体3类。黄连木果实含油率呈东北-西南由低到高的地理变异走向,种子和果肉含油率与生态因子也表现相似的变异趋势,但对生态因子的反映更为迟钝。
(8)采用SSR分子标记对黄连木8个群体进行遗传多样性分析,系统地揭示了不同群体在遗传多样性水平上的差异及遗传分化状况,在建立良好的黄连木SSR反应体系基础上,从阿月浑子的SSR引物中筛选出的9对SSR引物,在8个黄连木群体中共检测到43条等位基因,位点平均等位基因数为4.78,总多态位点百分率达100.0%。按检测到的有效等位基因数(Ne)和期望杂合度(He)各群体的遗传多样性由高至低依次为安康>唐县>顺平>辉县>略阳>栾川>林州>涉县。群体间的遗传分化系数(FST)平均为0.319,说明群体内变异是黄连木遗传变异的主要来源,并根据遗传距离将8个群体分为三大类。
(9)通过对黄连木集中分布区群落结构特征研究表明:太行山区黄连木群落乔木树种构成简单,黄连木的重要值高达73.1931,形成黄连木为主的单优群落。植物生长型与群落多样性的关系为,灌木层与草本层物种丰富度相近且明显高于乔木层,物种多样性指数表现出灌木层>草本层>乔木层的规律,均匀度表现出与多样性指数相同的变化趋势。太行山北段、中段和南段3个区域各层物种多样性指数变化趋势各不相同,表现出明显的分异现象。太行山黄连木种群都是增长型,种群格局总体呈集群分布,而太行山中段地区却表现为均匀分布。
(10)以植物的重要值确定各层片的优势种,将建群种相同,林下片层优势种相同的群落划分为群丛,秦岭南坡黄连木次生林共分为3个植被型,5个群系,6个群丛。对不同群落物种多样性分析表明,秦岭南坡物种丰富度、多样性指数和均匀度,总体表现为灌木层>草本层>乔木层,灌木层和草本层种类丰富,乔木层组成相对简单。
In order to serve practical production, the research was carried out from different perspectives of biology, genetics and breeding science, ecology and molecular biology with Chinese Pistache (Pistachia chinensis Bunge) as research object, and the estabilishment of high-yielding energy base as the goal. Based on fully tap and utilization of existing genetic variations, the multi-level research for individual, population as well as community of Chinese Pistache were performed. It provided a theoretical basis for further development and utilization of Chinese pistache.The main results were as the following:
(1) The criterion and method of plus tree selection of Chinese Pistache were developed preliminarily through selection of plus trees with total 228 plus trees selected. On the basis of analysis of main characteristics, the individuals with such single superior traits as high-oil content, resistance to insect and high fruit yield were selected, which laid the foundation for further genetic improvement, and provided the material for the future establishment of Chinese Pistache germplasm resources as well.
(2) Based on investigation and analysis,a comprehensive evaluation system of superior traits of Chinese Pistache was set up with the method of analytic hierarchy process (AHP),which were composed of the grade system, the grade of subsystem and the grade of indicator, according to the energy tree’s characteristics as well as principle of indicator system for assessment. The weights of indicators and evaluation equavation were determined with the method of AHP, Delphi method and Index Comprehensive Evaluation Method.
(3) Combining with the comprehensive evaluation system of superior traits of Chinese Pistache, 68 high-yield, high-oil content and fine-resistant superior individual tree were chosen by means of analysis of comprehensive selection of multi-characters.Meanwhile, based on the selection of plus tree, seedling testing was carried out to study the intrinsic relationship and genetic variation of the seedling growth traits, as such,the genetic parameters were also calculated and the excellent families were selected. These selected families, requiring further observation in progeny forestry, could be used for seed orchard or vegetative propagation. At the same time, the relationship between parent and progeny characters of Chinese Pistache was studied by the means of Analysis of Canonical Correlation. The selection index model is put forward, and the Chinese Pistache plus tree tested is classified and selected repeatly.
(4) Based on the investigation and analysis, Chinese Pistache was divided into sorts of types according to 7 obvious variation traits. Through system analysis, and three-level classification indicator, Multi-traits comprehensive classification was carried out to divide Chinese Pistache into different types,and a number of superior types with good quality, high yield, strong resistance were preliminarily selected. Meanwhile,the identification methods of adult sexualization were presented to facilitate the use of production units.
(5) The analysis on main factors affecting individual yield showed that soil fertility, insect-fruit rate , crown area, fruit number per-panicle, proportion of fertile branches, hundred fruit weight have significant correlations with individual yield. Among these traits, insect-fruit rate, soil fertility, fruit quantity are the key factors contributing to individual yield, The other factors, such as tree height , DBH, elevation, crown also had some influence on the individual yield, which showed individual yield of Chinese Pistache was a comprehensive quantity character.
(6) Based on investigation and analysis of 11 natural pupulations of Chinese Pistache in China,morphological variation and diversity of pupulations were studied, variation of all 18 characteristics was significantly different among and within populations.CV among phenotypic characteristics was markably different, varied from 4.249%-27.31%. Average phenotypic differentiation coefficient (Vst) was 22.27%, which indicated that the genetic variation within population was the main sources of genetic diversity. The 11 populations were classified, according to cluster analysis, into 4 groups. The study on variation of Chinese Pistache phenotypic characters in different habitats showed that slope position and altitude have clear effectiveness on most traits, while aspect had less effective. With the method of spatial series substituting for time, the variations of phenotypic traits at different age were also analysed.
(7) Fruit oil content of 9 populations of Chinese Pistache was investigated by Nuclear Magnetic Resonance( NMR) method. The result showed:contents of oil of fruit, seed and pulp were 36.183%、26.594%、50.503% respectively. There were significant variance among different populations, The populations were classified into 3 groups by clustering analysis,including lower variation and middle oil content population, lower variation and low content of oil population, high variation and high content of oil population. Fruit oil content of Chinese Pistache had the trend of southwest- northeast variation, the oil content of seed and pulp have same trend with ecological factors as fruits’, which reflected more slowly to ecological factors.
(8) Simple sequence repeat molecular marker systems were used to analyse the genetic diversity and genetic structure of 8 Chinese Pistache populations, on the basis of establishment of optimized PCR reaction system, nine pairs of SSR primers were selected from the primers which were designed for Pistacia vera, the total number of alleles 43 and average 4.78 for per locus was observed in eight populations, The percentage of polymorphic loci(P) was 100%. According to the numbers of effective alleles and expected heterozygosity, genetic diversity of populations from high to low was AK>TX>SP>HX>LY>LC >LZ>SX. The proportion of genetic differentiation among populations accounted for 0.319,which indicated that the main source of variation of Chinese Pistache was variation among populations,the eight population were divided into three groups with genetic distance.
(9) By the plots investigation, the structural characteristics of Chinese Pistache community was studied, the results showed that: In Taihang Mountains regions, Chinese Pistache community had a simple tree stratum component, the important value of Chinese pistache is up to 73.1931, as form Chinese Pistache-gifted community. The species richness and evenness of shrub stratum and herb stratum were similar but obviously higher than tree stratum. The diversity index for different strata was in the following order: shrub stratum> herb stratum> tree stratum, evenness had the same trend as species diversity index. The species diversity index of different regions had different variable trend, showing clear differentiation phenomenon. Chinese Pistache in Taihang Mountains was all growing type of population structure, distribution pattern was aggregate type in general, while the middle region was even distribution pattern.
(10) The dominant species of all levels was determined with important values, the community with same dominant species classified into community association, Chinese Pistache secondary forest in southern of Qinling Mountains could be divided into three vegetation types, five formations, six associations. The species diversity analysis on different communities showed that species richness, diversity index and evenness for different strata was in the following order: shrub stratum> herb stratum> tree stratum, the species of herbaceous and shrubs strata were rich, while species component of trees stratum relatively simple.
(1)通过黄连木优树选择,初步制定了黄连木的选优方法和标准,在各集中分布区初选优树228株。在对主要性状分析的基础上,选择出含油率高、抗虫性强、产量高的各种类型优良单株,为进一步遗传改良奠定基础,也为将来建立黄连木种质资源库提供材料。
(2)根据黄连木作为能源树种的特点,遵照指标体系的制定原则,提出了由目标层、控制层和指标层组成的黄连木优良性状综合评价指标体系。同时应用层次分析法、特尔斐法、指数综合评价法确定了各评价指标的权重和评价方法,为黄连木优良性状的划分和评选提供理论依据。
(3)结合黄连木优良性状综合评价指标体系,进行多性状综合选优分析,选择出高产、高含油率、抗黄连小蜂综合性状优良的单株68株。在优树选择的基础上,进行多地点黄连木优树苗期测定,探讨苗期各生长性状之间的内在关系和遗传变异,进行遗传参数估算和优良家系的评选,并运用典范相关分析的方法对亲代和子代性状的关系进行分析,构造指数方程,对参试的黄连木优树进行分类复选。
(4)在全面调查分析的基础上,选用变异较为明显的七个性状,将黄连木划分为各种不同类型。通过系统分析,确定了三级分类依据,进行多性状综合分类,初步评选出一些品质好、产量高,抗性强的优良类型。同时提出成年树雌雄株形态鉴别的方法,以方便生产单位使用。
(5)对产量的主要影响因素的分析表明,黄连木果实产量是一个综合数量性状,受多因子的影响,其中虫果率、土壤肥力、果穗粒数对单株产量影响最为明显。加强病虫害防治,增加土壤肥力,进行树体管护,促进果穗结实是提高黄连木产量的关键环节。
(6)以11个黄连木天然群体为试材,对18个表型性状的变异与多样性的分析表明,黄连木表型性状存在极其丰富的变异,各性状变异系数相差较大,变幅在4.249%- 27.31%之间,果实性状的变异系数最小,受较高的遗传控制,群体间平均表型分化系数VST为22.27%,说明群体内变异是黄连木变异的主要来源,聚类分析把11个群体分为4类。不同生境黄连木表型性状变异研究表明,坡向和海拨对多数性状有明显的影响,而坡位的作用较不明显,同时用空间代时间的方法,分析了不同年龄黄连木表型性状的变异。
(7)采用核磁共振(NMR)技术,对黄连木分布区9个群体果实含油率进行取样测定,结果表明:果实、种子和果肉含油率分别为36.183%、26.594%、50.503%,不同群体间差异显著,群体平均表型分化系数达到92.08%。系统聚类分析将黄连木群体分为低变异中等含油率群体、低变异低含油率群体、高变异高含油群体3类。黄连木果实含油率呈东北-西南由低到高的地理变异走向,种子和果肉含油率与生态因子也表现相似的变异趋势,但对生态因子的反映更为迟钝。
(8)采用SSR分子标记对黄连木8个群体进行遗传多样性分析,系统地揭示了不同群体在遗传多样性水平上的差异及遗传分化状况,在建立良好的黄连木SSR反应体系基础上,从阿月浑子的SSR引物中筛选出的9对SSR引物,在8个黄连木群体中共检测到43条等位基因,位点平均等位基因数为4.78,总多态位点百分率达100.0%。按检测到的有效等位基因数(Ne)和期望杂合度(He)各群体的遗传多样性由高至低依次为安康>唐县>顺平>辉县>略阳>栾川>林州>涉县。群体间的遗传分化系数(FST)平均为0.319,说明群体内变异是黄连木遗传变异的主要来源,并根据遗传距离将8个群体分为三大类。
(9)通过对黄连木集中分布区群落结构特征研究表明:太行山区黄连木群落乔木树种构成简单,黄连木的重要值高达73.1931,形成黄连木为主的单优群落。植物生长型与群落多样性的关系为,灌木层与草本层物种丰富度相近且明显高于乔木层,物种多样性指数表现出灌木层>草本层>乔木层的规律,均匀度表现出与多样性指数相同的变化趋势。太行山北段、中段和南段3个区域各层物种多样性指数变化趋势各不相同,表现出明显的分异现象。太行山黄连木种群都是增长型,种群格局总体呈集群分布,而太行山中段地区却表现为均匀分布。
(10)以植物的重要值确定各层片的优势种,将建群种相同,林下片层优势种相同的群落划分为群丛,秦岭南坡黄连木次生林共分为3个植被型,5个群系,6个群丛。对不同群落物种多样性分析表明,秦岭南坡物种丰富度、多样性指数和均匀度,总体表现为灌木层>草本层>乔木层,灌木层和草本层种类丰富,乔木层组成相对简单。
In order to serve practical production, the research was carried out from different perspectives of biology, genetics and breeding science, ecology and molecular biology with Chinese Pistache (Pistachia chinensis Bunge) as research object, and the estabilishment of high-yielding energy base as the goal. Based on fully tap and utilization of existing genetic variations, the multi-level research for individual, population as well as community of Chinese Pistache were performed. It provided a theoretical basis for further development and utilization of Chinese pistache.The main results were as the following:
(1) The criterion and method of plus tree selection of Chinese Pistache were developed preliminarily through selection of plus trees with total 228 plus trees selected. On the basis of analysis of main characteristics, the individuals with such single superior traits as high-oil content, resistance to insect and high fruit yield were selected, which laid the foundation for further genetic improvement, and provided the material for the future establishment of Chinese Pistache germplasm resources as well.
(2) Based on investigation and analysis,a comprehensive evaluation system of superior traits of Chinese Pistache was set up with the method of analytic hierarchy process (AHP),which were composed of the grade system, the grade of subsystem and the grade of indicator, according to the energy tree’s characteristics as well as principle of indicator system for assessment. The weights of indicators and evaluation equavation were determined with the method of AHP, Delphi method and Index Comprehensive Evaluation Method.
(3) Combining with the comprehensive evaluation system of superior traits of Chinese Pistache, 68 high-yield, high-oil content and fine-resistant superior individual tree were chosen by means of analysis of comprehensive selection of multi-characters.Meanwhile, based on the selection of plus tree, seedling testing was carried out to study the intrinsic relationship and genetic variation of the seedling growth traits, as such,the genetic parameters were also calculated and the excellent families were selected. These selected families, requiring further observation in progeny forestry, could be used for seed orchard or vegetative propagation. At the same time, the relationship between parent and progeny characters of Chinese Pistache was studied by the means of Analysis of Canonical Correlation. The selection index model is put forward, and the Chinese Pistache plus tree tested is classified and selected repeatly.
(4) Based on the investigation and analysis, Chinese Pistache was divided into sorts of types according to 7 obvious variation traits. Through system analysis, and three-level classification indicator, Multi-traits comprehensive classification was carried out to divide Chinese Pistache into different types,and a number of superior types with good quality, high yield, strong resistance were preliminarily selected. Meanwhile,the identification methods of adult sexualization were presented to facilitate the use of production units.
(5) The analysis on main factors affecting individual yield showed that soil fertility, insect-fruit rate , crown area, fruit number per-panicle, proportion of fertile branches, hundred fruit weight have significant correlations with individual yield. Among these traits, insect-fruit rate, soil fertility, fruit quantity are the key factors contributing to individual yield, The other factors, such as tree height , DBH, elevation, crown also had some influence on the individual yield, which showed individual yield of Chinese Pistache was a comprehensive quantity character.
(6) Based on investigation and analysis of 11 natural pupulations of Chinese Pistache in China,morphological variation and diversity of pupulations were studied, variation of all 18 characteristics was significantly different among and within populations.CV among phenotypic characteristics was markably different, varied from 4.249%-27.31%. Average phenotypic differentiation coefficient (Vst) was 22.27%, which indicated that the genetic variation within population was the main sources of genetic diversity. The 11 populations were classified, according to cluster analysis, into 4 groups. The study on variation of Chinese Pistache phenotypic characters in different habitats showed that slope position and altitude have clear effectiveness on most traits, while aspect had less effective. With the method of spatial series substituting for time, the variations of phenotypic traits at different age were also analysed.
(7) Fruit oil content of 9 populations of Chinese Pistache was investigated by Nuclear Magnetic Resonance( NMR) method. The result showed:contents of oil of fruit, seed and pulp were 36.183%、26.594%、50.503% respectively. There were significant variance among different populations, The populations were classified into 3 groups by clustering analysis,including lower variation and middle oil content population, lower variation and low content of oil population, high variation and high content of oil population. Fruit oil content of Chinese Pistache had the trend of southwest- northeast variation, the oil content of seed and pulp have same trend with ecological factors as fruits’, which reflected more slowly to ecological factors.
(8) Simple sequence repeat molecular marker systems were used to analyse the genetic diversity and genetic structure of 8 Chinese Pistache populations, on the basis of establishment of optimized PCR reaction system, nine pairs of SSR primers were selected from the primers which were designed for Pistacia vera, the total number of alleles 43 and average 4.78 for per locus was observed in eight populations, The percentage of polymorphic loci(P) was 100%. According to the numbers of effective alleles and expected heterozygosity, genetic diversity of populations from high to low was AK>TX>SP>HX>LY>LC >LZ>SX. The proportion of genetic differentiation among populations accounted for 0.319,which indicated that the main source of variation of Chinese Pistache was variation among populations,the eight population were divided into three groups with genetic distance.
(9) By the plots investigation, the structural characteristics of Chinese Pistache community was studied, the results showed that: In Taihang Mountains regions, Chinese Pistache community had a simple tree stratum component, the important value of Chinese pistache is up to 73.1931, as form Chinese Pistache-gifted community. The species richness and evenness of shrub stratum and herb stratum were similar but obviously higher than tree stratum. The diversity index for different strata was in the following order: shrub stratum> herb stratum> tree stratum, evenness had the same trend as species diversity index. The species diversity index of different regions had different variable trend, showing clear differentiation phenomenon. Chinese Pistache in Taihang Mountains was all growing type of population structure, distribution pattern was aggregate type in general, while the middle region was even distribution pattern.
(10) The dominant species of all levels was determined with important values, the community with same dominant species classified into community association, Chinese Pistache secondary forest in southern of Qinling Mountains could be divided into three vegetation types, five formations, six associations. The species diversity analysis on different communities showed that species richness, diversity index and evenness for different strata was in the following order: shrub stratum> herb stratum> tree stratum, the species of herbaceous and shrubs strata were rich, while species component of trees stratum relatively simple.
引文
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