芒在中国的分布及其遗传多样性研究
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摘要
芒(Miscanthus sinensis)系禾本科(Poaceae)芒属(Miscanthus Andersson)的一个重要种,因其具有分布广、适应性强、多年生、生物质产量和纤维素含量高等众多优点而被认为是极具开发潜力的新型能源植物资源。我国是芒的分布中心,但这一宝贵物种在我国目前仍处于鲜为人知的野生状态。为了探明这一重要植物资源在我国的蕴藏情况,提供其种质资源开发、利用与保护的理论依据,本文对我国芒进行了系统的分布调查、气候研究和资源收集,采集了500余份活体材料,建立了芒种质资源圃,并从中选取388份有代表性的材料进行了我国芒种质资源遗传多样性的研究,同时还分别采用形态标记和分子标记对广西和海南两个省份的芒进行了野生居群遗传结构分析。主要研究结果如下：
1.芒的分布和气候模式研究结果表明：(1)芒在我国分布广泛。其地理分布范围为东经100.45°～127.55°E,北纬18.34°～43.70°N,垂直分布范围为一12～1900m；行政地区的分布为：安徽、重庆、福建、甘肃、广东、广西壮族自治区、贵州、海南、黑龙江、河南、河北、湖北、湖南、江苏、江西、吉林、辽宁、陕西、山东、四川、台湾、云南和浙江等23个省(直辖市、自治区)；区系分布为蒙古草原地区、东北地区、华北地区、华东地区、华中地区、岭南山地地区、滇黔桂地区、台湾地区、南海地区、北部湾地区和滇缅泰地区等11个植物地区。(2)芒的生境多样。其自然生境包括道路边、林缘、草甸、山脚、山坡、山谷、山顶、海滩边、溪边、灌木丛中、竹林边。土壤类型包括红壤、黄壤、砂壤、棕壤、紫壤、腐殖土；(3)芒的气候适应性强。其气候模式为年积温2650.00-8500.00℃,年平均温度为3.40～24.40℃,为30.90～45.20℃,最冷月低温—45.00～8.00℃,1月份平均气温为-23.00～18.70℃,7月份平均气温为18.20～32.70℃,全年无霜期为128.00～365.00d,年累积日照时数为511.00～2700.00h,年平均降雨量为466.50～2444.00mm。
2.芒种质资源表型多样性研究结果表明：(1)芒种质资源表型变异丰富。15个分类学性状的变异范围16.21～61.39%,平均29.27%。15个农艺性状的变异范围20.78～89.75%,平均38.05%。(2)分类学性状变异主要来源于种子宽、种子厚、种子长和千粒重等4个性状,农艺性状的变异主要来源于单茎干重,叶片数,主茎长,最大叶片宽,第1节窄边宽,第1节宽边直径等6个性状。(3)不同区系间表型性状变异大小顺序为华北地区(34.95%)>华东地区(32.17%)>华中地区(27.08%)>东北地区(26.40%)>岭南山地地区(25.11%)>滇黔桂地区(24.69%)>北部湾地区(23.86%)>南海地区(23.65%)>滇缅泰地区(20.69%),说明华北地区芒种质资源多样性最丰富,而滇缅泰地区遗传多样性相对贫乏。(4)通过聚类分析可以将参试的388份种质资源成“低矮型”和“高大型”2类,低矮型主要来源于东北和华北地区,高大型主要来源于华东地区、华中地区、岭南山地地区、滇黔桂地区、南海地区、北部湾地区和滇缅泰地区。不同区系来源种质能聚成一类,说明芒种群内不同区系间可以产生杂交。(5)综合考虑产量和产量构成因子,根据隶属函数法对参试种质资源进行赋值(D),根据D值从小到大的顺序将所有种质资源分成“差”(32份,D介于0.05～0.18),“较差”(72份,D介于0.18～0.32),“中等”(129份,D介于0.32～0.45),“良”(131份,D介于0.45～0.58)和“优”(23份,D介于0.58～0.71)等5大类。
3.芒种质资源SSR标记遗传多样研究结果表明：(1)芒种质资源DNA水平遗传多样性丰富。利用33对SSR引物对来自中国16个省的46份野生芒种质进行遗传多样性分析。结果显示：33对SSR引物共扩增出87条DNA条带,75条为多态性条带,占86.21%,条带大小范围80-310bp。(2)遗传多样性参数分析结果：Shannon's信息指数(I)变幅为0.020～1.522,平均0.745,引物多态性信息含量(PIC)变幅为0.040～0.738,平均0.445,遗传相似系数(GS)的变幅为0.315～0.933,平均0.569,说明我国芒种质资源遗传基础宽,遗传多样性丰富。(3)相似系数UMPGA聚类结果与主成分聚类(PCA)结果一致,可将46份种质分为3大类群,类群Ⅰ主要由中部芒组成,类群Ⅱ主要由北方芒组成,类群Ⅲ主要由南方芒组成,西南芒在每个类群中均有渗透,这一结果说明芒种质资源的遗传分化与其种源的地理分布有一定的相关性,但与地理起源不能完全吻合。
4.芒野生居群遗传多样性研究结果表明：(1)芒野生居群表型变异丰富。各表型性状在居群内的和居群间差异均达到极显著水平,同一性状在不同居群间变异系数的变化范围为11.35～49.86%,同一居群内不同的性状变异系数的变化范围为17.07～27.65%。(2)芒居群内的表型变异大于居群间的表型变异,18个表型性状分化系数变化范围为17.99～47.21%,平均为32.05%,居群内表型分化率为67.95%,说明芒表型性状的变异主要来自居群内。(3)芒居群在DNA水平上也有着丰富的遗传多样性。SSR标记分析海南岛15个野生居群,显示其居群遗传相似性系数在0.8241～0.9647之间,遗传多样性指数H居群=0.0930,Shannon信息指数J居群=0.1371。(4)芒在DNA水平上变异也同样是居群内大于居群间。居群遗传变异结构分析表明芒的基因多样性指数Ht=0.1755,居群内的基因多样性指数Hs=0.0993,居群间的遗传分化系数Gst=0.4342<0.5,说明芒居群内的基因变异大于居群间。居群间的基因流Nm=0.5643<1.0,说明居群间基因流较弱,遗传漂变和近交可能是造成居群内遗传分化的主要因素。
Miscanthus sinensis is one of the species of Miscanthus Genus, Poaceae. Recently, it was regarded as a kind of energy crop for its wild distribution, large biomass, strong resistance and higher cellulose content. In this paper, we investigated the nature distribution of M. sinensis in China and collected a number of germplsams, from which we selected388collections to do genetic diversity research. At the same time, we chose Guangxi and Hainan provinces to analyze genetic diversity in wild populations of M. sinensis with morphological and molecular markers respectively. The results show that:
1. The result of distribution and climate adaptation of Miscanthus sinensis show that1) The nature distribution field of M. sinensis in China was100.45°～127.55°E,18.34°～43.70°N, Alt.-12-1900m, the administrative areas distribute23provinces, which consist of Anhui, Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Heilongjiang, Henan, Hebei, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Shaanxi, Shandong, Sichuan, Taiwan, Yunnan and Zhejiang. The floristic distributions of M. sinensis were IB4, IIID7, IIID8, IIID9, IIID10, IIID11, IIID12, IVG19, IVG21, IVG22and IVG23.2) The habitat diversity of M. sinensis was rich:the habitat of M. sinensis included roadside, shrub of forest, foot of hill, hillside, top of mountain, seaside, riverside, bamboo forest, etc.3) The adaptation of M. sinensis was abundant. It's annual accumulated temperature of climate model was variable from2650.00℃to8500.00℃, yearly average temperature was variable from3.40℃to24.40℃, yearly average rain fall was varied from466.50from to2444.00mm, extremely high temperature was varied from-8.86℃to6.40℃,extremely low temperature was varied from -45.00℃to8.00℃average temperature of January was varied from-23.00℃to18.70℃, average temperature of July was varied from-23.00℃to18.70℃, frost-free period was varied from128.00d to365d and annual accumulated sun time was variable from511.00h to2700h.
2. The research results of genetic diversity of phenotypic traits of accessions show1) The germplasm phenotypic variation of M. sinensis was abundant. The average coefficients of variation of15taxonomic traits were29.27%with the variation from16.21%to61.39%and the average coefficients of variation of15agronomic traits were38.05%with the variation from20.78%to89.75%.2) The variation of taxonomic traits comes from4traits, which were seed width, seed thickness, seed length,1000-seed weight. The variation of agronomic traits come from6traits, which were dry weight per tiller, leaf number per tiller, stem height, largest leaf width, first node long axis and first node short axis.3) The average coefficient of phenotypic traits of9flora were IIID8(34.95%)> IIID9(32.17%)> IIID10(27.08%)> IIID7(26.40%)> IIID11(25.11%)> IIID12(24.69%)> IVG22(23.86%)>IVG21(23.65%)>IVG23(20.69%).4)388accessions can be divided into two categories by cluster analysis, which were lower type and topper type. The lower type mainly comes from IIID7and IIID8and the topper type were main consist of the accessions from other flora.5) Take yield and yield component factor into consideration,388accessions were comprehensively evaluated by subordination function method. According to the D index,388accessions can be divided into five categories:I type with the D value which was varied from0.05to0.18, II type with the D value which was varied from0.18to0.32, III type with the D value was varied from0.32to0.45, IV type with the D value which was varied from0.45to0.58and the V type with the D value which was varied from0.58to0.71.
3. The research result of genetic diversity by SSR markers show that1)33SSR primer pairs produced87bands with75polymorphic (86.21%) and the size of amplified DNA varied from80to310bp.2) The result of genetic diversity: Shannon diversity index (I) was0.020～1.522, the average is0.745. Polymorphic information content (PIC) was about0.040～0.738, the average is0.445. Genetic similarity coefficient is about0.315～0.933, the average is0.569. The genetic similarity coefficient of UPGMA and PCA Cluster were the same. The46collections of M. sinensis can be divided into3groups. Group I was mainly composed of the germplasm from center China, group II composed of those from north China and group III composed of those from south China. These results indicated that there were some relevance between M. sinensis varieties and geographical distributions, but M.sinensis varieties were not inconformity with geographic origin.
4. The research results of genetic diversity of nature populations show that (1) There were abundant genetic diversity in the nature populations of M. sinensis and the analysis of variance for all traits showed that there were significant differences among and within populations. The variation coefficients of the same trait in different populations were varied from11.35%to49.86%. The variation coefficients of different traits in the same population were varied from17.07%to27.65%.(2)The phenotypic differentiation coefficients (VST) within populations were bigger than among populations. The average is was32.05%.The variation range of18phenotypic differentiation coefficients was from17.99%to47.21%, phenotypic differentiation rate was67.95%. This means that the genetic variability of M. sinensis existed mainly within population.(3) DNA level of populations have high genetic diversity. SSR marks analyzed15wild populations showed that the genetic similarity coefficients from0.8241to0.9647, the gene diversity of Nei's (H) was0.0930and the average information index of Shannon (I) was0.1371.(4) The DNA level variation within populations were bigger than among populations.The total gene diversity of Nei's (Ht) was0.1755and the gene diversity of Nei's within population (Hs) was0.0993, the gene differentiation coefficient (Gst) was0.4342<0.50, which suggested genetic diversity existed within population. The index of gene flow (Nm) was0.5643<1.0suggested gene flow was small, genetic drift and inbreeding might be the main factors influencing population genetic structure of M. sinensis.
1.芒的分布和气候模式研究结果表明：(1)芒在我国分布广泛。其地理分布范围为东经100.45°～127.55°E,北纬18.34°～43.70°N,垂直分布范围为一12～1900m；行政地区的分布为：安徽、重庆、福建、甘肃、广东、广西壮族自治区、贵州、海南、黑龙江、河南、河北、湖北、湖南、江苏、江西、吉林、辽宁、陕西、山东、四川、台湾、云南和浙江等23个省(直辖市、自治区)；区系分布为蒙古草原地区、东北地区、华北地区、华东地区、华中地区、岭南山地地区、滇黔桂地区、台湾地区、南海地区、北部湾地区和滇缅泰地区等11个植物地区。(2)芒的生境多样。其自然生境包括道路边、林缘、草甸、山脚、山坡、山谷、山顶、海滩边、溪边、灌木丛中、竹林边。土壤类型包括红壤、黄壤、砂壤、棕壤、紫壤、腐殖土；(3)芒的气候适应性强。其气候模式为年积温2650.00-8500.00℃,年平均温度为3.40～24.40℃,为30.90～45.20℃,最冷月低温—45.00～8.00℃,1月份平均气温为-23.00～18.70℃,7月份平均气温为18.20～32.70℃,全年无霜期为128.00～365.00d,年累积日照时数为511.00～2700.00h,年平均降雨量为466.50～2444.00mm。
2.芒种质资源表型多样性研究结果表明：(1)芒种质资源表型变异丰富。15个分类学性状的变异范围16.21～61.39%,平均29.27%。15个农艺性状的变异范围20.78～89.75%,平均38.05%。(2)分类学性状变异主要来源于种子宽、种子厚、种子长和千粒重等4个性状,农艺性状的变异主要来源于单茎干重,叶片数,主茎长,最大叶片宽,第1节窄边宽,第1节宽边直径等6个性状。(3)不同区系间表型性状变异大小顺序为华北地区(34.95%)>华东地区(32.17%)>华中地区(27.08%)>东北地区(26.40%)>岭南山地地区(25.11%)>滇黔桂地区(24.69%)>北部湾地区(23.86%)>南海地区(23.65%)>滇缅泰地区(20.69%),说明华北地区芒种质资源多样性最丰富,而滇缅泰地区遗传多样性相对贫乏。(4)通过聚类分析可以将参试的388份种质资源成“低矮型”和“高大型”2类,低矮型主要来源于东北和华北地区,高大型主要来源于华东地区、华中地区、岭南山地地区、滇黔桂地区、南海地区、北部湾地区和滇缅泰地区。不同区系来源种质能聚成一类,说明芒种群内不同区系间可以产生杂交。(5)综合考虑产量和产量构成因子,根据隶属函数法对参试种质资源进行赋值(D),根据D值从小到大的顺序将所有种质资源分成“差”(32份,D介于0.05～0.18),“较差”(72份,D介于0.18～0.32),“中等”(129份,D介于0.32～0.45),“良”(131份,D介于0.45～0.58)和“优”(23份,D介于0.58～0.71)等5大类。
3.芒种质资源SSR标记遗传多样研究结果表明：(1)芒种质资源DNA水平遗传多样性丰富。利用33对SSR引物对来自中国16个省的46份野生芒种质进行遗传多样性分析。结果显示：33对SSR引物共扩增出87条DNA条带,75条为多态性条带,占86.21%,条带大小范围80-310bp。(2)遗传多样性参数分析结果：Shannon's信息指数(I)变幅为0.020～1.522,平均0.745,引物多态性信息含量(PIC)变幅为0.040～0.738,平均0.445,遗传相似系数(GS)的变幅为0.315～0.933,平均0.569,说明我国芒种质资源遗传基础宽,遗传多样性丰富。(3)相似系数UMPGA聚类结果与主成分聚类(PCA)结果一致,可将46份种质分为3大类群,类群Ⅰ主要由中部芒组成,类群Ⅱ主要由北方芒组成,类群Ⅲ主要由南方芒组成,西南芒在每个类群中均有渗透,这一结果说明芒种质资源的遗传分化与其种源的地理分布有一定的相关性,但与地理起源不能完全吻合。
4.芒野生居群遗传多样性研究结果表明：(1)芒野生居群表型变异丰富。各表型性状在居群内的和居群间差异均达到极显著水平,同一性状在不同居群间变异系数的变化范围为11.35～49.86%,同一居群内不同的性状变异系数的变化范围为17.07～27.65%。(2)芒居群内的表型变异大于居群间的表型变异,18个表型性状分化系数变化范围为17.99～47.21%,平均为32.05%,居群内表型分化率为67.95%,说明芒表型性状的变异主要来自居群内。(3)芒居群在DNA水平上也有着丰富的遗传多样性。SSR标记分析海南岛15个野生居群,显示其居群遗传相似性系数在0.8241～0.9647之间,遗传多样性指数H居群=0.0930,Shannon信息指数J居群=0.1371。(4)芒在DNA水平上变异也同样是居群内大于居群间。居群遗传变异结构分析表明芒的基因多样性指数Ht=0.1755,居群内的基因多样性指数Hs=0.0993,居群间的遗传分化系数Gst=0.4342<0.5,说明芒居群内的基因变异大于居群间。居群间的基因流Nm=0.5643<1.0,说明居群间基因流较弱,遗传漂变和近交可能是造成居群内遗传分化的主要因素。
Miscanthus sinensis is one of the species of Miscanthus Genus, Poaceae. Recently, it was regarded as a kind of energy crop for its wild distribution, large biomass, strong resistance and higher cellulose content. In this paper, we investigated the nature distribution of M. sinensis in China and collected a number of germplsams, from which we selected388collections to do genetic diversity research. At the same time, we chose Guangxi and Hainan provinces to analyze genetic diversity in wild populations of M. sinensis with morphological and molecular markers respectively. The results show that:
1. The result of distribution and climate adaptation of Miscanthus sinensis show that1) The nature distribution field of M. sinensis in China was100.45°～127.55°E,18.34°～43.70°N, Alt.-12-1900m, the administrative areas distribute23provinces, which consist of Anhui, Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Heilongjiang, Henan, Hebei, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Shaanxi, Shandong, Sichuan, Taiwan, Yunnan and Zhejiang. The floristic distributions of M. sinensis were IB4, IIID7, IIID8, IIID9, IIID10, IIID11, IIID12, IVG19, IVG21, IVG22and IVG23.2) The habitat diversity of M. sinensis was rich:the habitat of M. sinensis included roadside, shrub of forest, foot of hill, hillside, top of mountain, seaside, riverside, bamboo forest, etc.3) The adaptation of M. sinensis was abundant. It's annual accumulated temperature of climate model was variable from2650.00℃to8500.00℃, yearly average temperature was variable from3.40℃to24.40℃, yearly average rain fall was varied from466.50from to2444.00mm, extremely high temperature was varied from-8.86℃to6.40℃,extremely low temperature was varied from -45.00℃to8.00℃average temperature of January was varied from-23.00℃to18.70℃, average temperature of July was varied from-23.00℃to18.70℃, frost-free period was varied from128.00d to365d and annual accumulated sun time was variable from511.00h to2700h.
2. The research results of genetic diversity of phenotypic traits of accessions show1) The germplasm phenotypic variation of M. sinensis was abundant. The average coefficients of variation of15taxonomic traits were29.27%with the variation from16.21%to61.39%and the average coefficients of variation of15agronomic traits were38.05%with the variation from20.78%to89.75%.2) The variation of taxonomic traits comes from4traits, which were seed width, seed thickness, seed length,1000-seed weight. The variation of agronomic traits come from6traits, which were dry weight per tiller, leaf number per tiller, stem height, largest leaf width, first node long axis and first node short axis.3) The average coefficient of phenotypic traits of9flora were IIID8(34.95%)> IIID9(32.17%)> IIID10(27.08%)> IIID7(26.40%)> IIID11(25.11%)> IIID12(24.69%)> IVG22(23.86%)>IVG21(23.65%)>IVG23(20.69%).4)388accessions can be divided into two categories by cluster analysis, which were lower type and topper type. The lower type mainly comes from IIID7and IIID8and the topper type were main consist of the accessions from other flora.5) Take yield and yield component factor into consideration,388accessions were comprehensively evaluated by subordination function method. According to the D index,388accessions can be divided into five categories:I type with the D value which was varied from0.05to0.18, II type with the D value which was varied from0.18to0.32, III type with the D value was varied from0.32to0.45, IV type with the D value which was varied from0.45to0.58and the V type with the D value which was varied from0.58to0.71.
3. The research result of genetic diversity by SSR markers show that1)33SSR primer pairs produced87bands with75polymorphic (86.21%) and the size of amplified DNA varied from80to310bp.2) The result of genetic diversity: Shannon diversity index (I) was0.020～1.522, the average is0.745. Polymorphic information content (PIC) was about0.040～0.738, the average is0.445. Genetic similarity coefficient is about0.315～0.933, the average is0.569. The genetic similarity coefficient of UPGMA and PCA Cluster were the same. The46collections of M. sinensis can be divided into3groups. Group I was mainly composed of the germplasm from center China, group II composed of those from north China and group III composed of those from south China. These results indicated that there were some relevance between M. sinensis varieties and geographical distributions, but M.sinensis varieties were not inconformity with geographic origin.
4. The research results of genetic diversity of nature populations show that (1) There were abundant genetic diversity in the nature populations of M. sinensis and the analysis of variance for all traits showed that there were significant differences among and within populations. The variation coefficients of the same trait in different populations were varied from11.35%to49.86%. The variation coefficients of different traits in the same population were varied from17.07%to27.65%.(2)The phenotypic differentiation coefficients (VST) within populations were bigger than among populations. The average is was32.05%.The variation range of18phenotypic differentiation coefficients was from17.99%to47.21%, phenotypic differentiation rate was67.95%. This means that the genetic variability of M. sinensis existed mainly within population.(3) DNA level of populations have high genetic diversity. SSR marks analyzed15wild populations showed that the genetic similarity coefficients from0.8241to0.9647, the gene diversity of Nei's (H) was0.0930and the average information index of Shannon (I) was0.1371.(4) The DNA level variation within populations were bigger than among populations.The total gene diversity of Nei's (Ht) was0.1755and the gene diversity of Nei's within population (Hs) was0.0993, the gene differentiation coefficient (Gst) was0.4342<0.50, which suggested genetic diversity existed within population. The index of gene flow (Nm) was0.5643<1.0suggested gene flow was small, genetic drift and inbreeding might be the main factors influencing population genetic structure of M. sinensis.
引文
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