發(fā)布時間：2018-01-08 07:32

  本文關(guān)鍵詞：松嫩平原異質(zhì)生境蘆葦表型變異及其分子生態(tài)學(xué)研究　出處：《東北師范大學(xué)》2017年博士論文　論文類型：學(xué)位論文

  更多相關(guān)文章： 分子生態(tài)學(xué) 蘆葦 表型變異 遺傳多樣性 表觀遺傳分化 光合色素 異質(zhì)生境

【摘要】：物種形成雖然是一個漫長的時間過程,但經(jīng)常從占據(jù)不同生態(tài)環(huán)境的種群的適應(yīng)性分歧開始。遺傳分化是生物適應(yīng)異質(zhì)生境的重要機制之一。異質(zhì)生境間和生境內(nèi)植物的遺傳變異、表觀遺傳變異為自然選擇和適應(yīng)性進(jìn)化提供了原材料。探討植物在異質(zhì)生境間和生境內(nèi)表型差異的遺傳、表觀遺傳機理及其與適應(yīng)性進(jìn)化的關(guān)系是分子生態(tài)學(xué)的主流研究內(nèi)容之一。然而在小空間尺度上這方面的研究還不多見。蘆葦(Phragmites australis)是世界廣布種,在松嫩平原的濕地、沙地和草地廣泛生長,其營養(yǎng)生長期為各種家畜采食的優(yōu)良牧草,也是當(dāng)?shù)佧}堿化草甸生態(tài)恢復(fù)的主要植物。本研究以小空間尺度上松嫩平原異質(zhì)生境的蘆葦(Phragmites australis)、毛蘆葦(Phragmites hirsuta)為材料,通過野外調(diào)查、實驗室測定、AFLP(Amplified fragment length polymorphism)和MSAP(Methylation-sensitive amplified polymorphism)分子標(biāo)記檢測和同質(zhì)園實驗,運用分子生態(tài)學(xué)的統(tǒng)計分析方法,探究異質(zhì)生境基礎(chǔ)上的表型差異及其遺傳、表觀遺傳基礎(chǔ),以及適應(yīng)性進(jìn)化的意義。得到主要結(jié)果如下:(1)在土壤理化特征差異明顯的4種生境,蘆葦表型數(shù)量特征差異顯著。季節(jié)性積水的低洼堿化草甸土生境(H1)和沙地生境(H4)的株高、葉片等生長特征和生物量明顯大于典型堿化草甸土的羊草(Leymus chinensis)蘆葦混生群落的生境(H3)和明堿斑的生境(H2),大多數(shù)情況下H2最低。H1和H3的莖生物量分配較大,H4的葉生物量分配較大,H2的葉鞘生物量分配較大。土壤含水量、全磷、全氮均與蘆葦?shù)纳L呈正相關(guān),它們和有機質(zhì)含量與莖生物量分配呈正相關(guān),與葉生物量分配呈負(fù)相關(guān);土壤pH、電導(dǎo)率、硝態(tài)氮、Cl-、SO42-、Na+或K+與蘆葦生長呈負(fù)相關(guān),隨著土壤電導(dǎo)率、Cl-、Na+或K+的增加,莖生物量分配呈先增后減的趨勢,葉生物量分配呈先減后增、葉鞘生物量分配呈增加的趨勢;Mg2+和Ca2+與蘆葦生長呈正相關(guān)。葉生物量分配與莖生物量分配之間存在權(quán)衡現(xiàn)象。數(shù)量性狀之間、生物量分配之間的相關(guān)關(guān)系以及異速生長規(guī)律在生境間存在顯著差異。(2)在小空間尺度上,異質(zhì)生境內(nèi)和種群內(nèi)蘆葦存在遺傳多態(tài)。H1和H3蘆葦比H2和H4有更高的遺傳多樣性,這和生境幅度變異假說一致。生境基礎(chǔ)上堿地(H1、H2、H3)與沙地(H4)間遺傳分化更為明顯。應(yīng)用修飾的FDIST,Bayescan和Samβada3種方法探測到33個異常值位點,土壤Ca2+、全磷、Mg2+、pH等是自然選擇壓力主要環(huán)境因子。本章結(jié)果支持“帶有基因流的分歧”(“divergence-with-gene-flow”)模型,為進(jìn)一步研究局域適應(yīng)提供了基因組水平的數(shù)據(jù)。(3)在MSAP分析中約有2/3的位點是甲基化敏感位點(MSL)。異質(zhì)生境間4種類型甲基化模式所占比例不同。生境內(nèi)、種群內(nèi)表觀遺傳多樣性較高,H1、H2和H3明顯高于H4。4種生境間表觀遺傳分化顯著,堿地與沙地之間更明顯。H1與H3、H2與H3的表觀遺傳分化明顯大于遺傳分化。遺傳變異與表觀遺傳變異顯著相關(guān)。生境間表觀遺傳分化與適應(yīng)性遺傳分歧之間相關(guān)。植株表型特征與表觀遺傳多樣性的關(guān)系要比與遺傳多樣性的關(guān)系更為密切;遺傳多樣性參數(shù)與土壤全磷、Cl-、Ca2+含量呈顯著相關(guān)關(guān)系,而表觀遺傳多樣性參數(shù)與土壤含水量、全磷、全氮以及有機質(zhì)呈顯著相關(guān)關(guān)系,表觀遺傳多樣性受土壤營養(yǎng)成分的影響更大。(4)通過AFLP標(biāo)記,采用UPGMA、STRUCTURE、主坐標(biāo)分析方法對蘆葦、毛蘆葦和日本葦(Phragmites japonica)做分子水平分析,結(jié)果顯示,蘆葦、毛蘆葦組內(nèi)H3和H4明顯分開,長春市南湖公園蘆葦和H3、H4的蘆葦遺傳距離較近,意味蘆葦和毛蘆葦之間的分化是先于地理分化、生境分化的。MSAP標(biāo)記顯示蘆葦和毛蘆葦顯著的表觀遺傳分化,蘆葦、毛蘆葦組內(nèi)H3和H4也明顯分開,反映出蘆葦與毛蘆葦之間以DNA甲基化為基礎(chǔ)的表觀遺傳位點經(jīng)歷了比異質(zhì)生境引起的分化更為強烈的分化。推測毛蘆葦處在從蘆葦中分化出來的早中期。(5)野外調(diào)查發(fā)現(xiàn)H2(堿斑)中蘆葦?shù)娜~綠素a(Chla),葉綠素b(Chlb),總?cè)~綠素(Chl(a+b)),類胡蘿卜素(Car)濃度以及總?cè)~綠素/類胡蘿卜素(Chl(a+b)/Car ratio)均顯著低于H1。只有Chla/Chlb沒有顯著變化。同質(zhì)園實驗中小空間尺度上Chla,、Chlb、Chl(a+b)、和Car濃度存在局域適應(yīng)現(xiàn)象。應(yīng)用種群基因組方法FDIST,Bayescan和Bayenv探測到異常值位點(outlier loci)和形態(tài)分化系數(shù)-遺傳分化系數(shù)(QST-FST)比較分析均支持局域適應(yīng)的結(jié)論。土壤含水量、pH和電導(dǎo)率均可形成自然選擇壓力而引起光合色素的適應(yīng)性分歧。本研究有助于小尺度異質(zhì)生境不同表型蘆葦?shù)倪z傳和表觀遺傳基礎(chǔ)的理解,豐富了蘆葦分子生態(tài)學(xué)的研究內(nèi)容,可為生態(tài)適應(yīng)性、生物抗逆性、蘆葦屬分類的研究、以及包括生態(tài)恢復(fù)、遺傳資源保護(hù)、遺傳改良在內(nèi)的應(yīng)用實踐提供科學(xué)參考或理論依據(jù)。
[Abstract]:The formation of species although it is a long time, but often from the adaptability differences occupy different ecological environment. Population genetic differentiation is one of the important mechanisms of heterogeneous habitat biological adaptation. The genetic variation between heterogeneous habitats and habitat within plants, epigenetic variation in natural selection and adaptive evolution provides the raw material of. In plant genetic heterogeneity among habitats and habitat phenotypic differences, the epigenetic mechanism and its relationship with adaptive evolution is one of the main research contents of molecular ecology. However, research on this aspect at small spatial scales are rare. Reed (Phragmites australis) is a world wide distributed species in the Songnen Plain Wetland sand and grass, widely grown, the vegetative growth stage for various animal feed forage for food, the main plant is a local salinization meadow ecological restoration. In this study, a small space On the scale of heterogeneous habitats in Songnen Plain reed (Phragmites australis), Mao Luwei (Phragmites hirsuta) as material, through field investigation, laboratory test, AFLP (Amplified fragment length polymorphism) and MSAP (Methylation-sensitive amplified polymorphism) molecular markers and common garden experiments, using statistical analysis method to explore the molecular ecology, phenotypic differences in heterogeneous habitat based and on the genetic, epigenetic basis and significance of adaptive evolution. The main results are as follows: (1) on the physicochemical characteristics of the difference between the 4 habitats significantly soil, quantitative differences significant. Phenotypic characteristics of Phragmites low-lying meadow alkali soil habitat seasonal water (H1) and sandy habitat (H4) plant height leaves, growth characteristics and biomass was significantly higher than that of typical alkalized meadow soil of Leymus chinensis (Leymus chinensis) habitats, mixed communities (H3) and alkali spot The habitat (H2), stem biomass H2 minimum.H1 and H3 distribution in most cases is large, leaf biomass distribution of H4 large amount of leaf sheath distribution of H2 greatly. Biological soil moisture, total phosphorus, total nitrogen were positively related to growth of reed, they positively with organic matter content and stem biomass allocation correlation was negatively correlated with leaf biomass allocation; soil pH, conductivity, nitrate nitrogen, Cl-, SO42-, Na+ or K+ and reed growth was negatively correlated with soil conductivity, Cl-, Na+ or K+ increased the stem biomass allocation increased firstly and then decreased, the leaf biomass allocation was first decreased and then increased, leaf biomass allocation increased; Mg2+ and Ca2+ and reed growth was positively correlated. There is a trade-off phenomenon between leaf biomass allocation and stem biomass allocation. Quantitative traits between the relationship between biomass allocation and allometry have significant differences in habitat (2) in between. The small spatial scale, heterogeneous habitat and population genetic polymorphism of.H1 and H3 in reed reed than H2 and H4 have higher genetic diversity, and the favorable habitat. Habitat Based on amplitude variation hypothesis (H1, H2, alkaline H3) and sand (H4) genetic differentiation among the applications of modified is more obvious. FDIST, Bayescan and Sam beta ADA3 method to detect 33 outlier loci, soil Ca2+, total phosphorus, Mg2+, pH are the main environmental factors of natural selection pressure. The results of this chapter support "with gene flow differences" ("divergence-with-gene-flow") model, provides the genome level data for further study on local adaptation. (3) in the MSAP analysis about 2/3 sites methylation sensitive sites (MSL). Among the 4 types of heterogeneous habitat methylation patterns in different proportion. Chinese population apparent higher genetic diversity, H1, H2 and H3 significantly higher than the H4.4 table between habitats view Significant genetic differentiation between soil and sand is more obvious,.H1 and H3, H2 and H3 on genetic differentiation table is significantly greater than the genetic variation and genetic differentiation. The apparent significant genetic variation among habitats. The apparent genetic differentiation and genetic differences between related plant adaptability. The phenotypic characteristics and apparent genetic diversity than genetic diversity and relationship more closely; the genetic diversity parameters and soil total phosphorus, Cl-, there was a significant correlation between the content of Ca2+, and the apparent genetic diversity parameters and soil moisture content, total phosphorus, total nitrogen and organic matter were significant correlation between the effect of apparent genetic diversity by soil nutrients more. (4) by AFLP markers, using UPGMA, STRUCTURE, principal coordinate analysis method of reed, reed and reed hair Japan (Phragmites japonica) to do the molecular analysis, results showed that reed, reed hair groups within H3 and H4 significantly separated, Changchun City South Lake Park reed and reed H3, H4's genetic distance is short, means that the differentiation between the reed and reed hairs are prior to geographical differentiation,.MSAP marker showed habitat differentiation of reed and reed hair significant epigenetic differentiation, reed, reed group H3 and H4 in hair was also separated based epigenetic site experience the differentiation caused by heterogeneous habitat is more intense than the differentiation between reed and reed of hair reflected by DNA methyl. In the early stage of differentiation that Mao reed from reed. (5) field investigation found that H2 (alkali spot) in reed chlorophyll a (Chla), chlorophyll b (Chlb), total the chlorophyll (Chl (a+b)), carotenoid (Car) concentration and total chlorophyll / carotenoid (Chl (a+b) /Car ratio) were significantly lower than that of H1. only Chla/Chlb did not change significantly. The small spatial scale common garden experiment, Chla, Chlb, Chl (a+b), and the existence of local concentration of Car To adapt to the phenomenon. The application of FDIST method Bayescan and Bayenv genome species, to detect outlier loci (outlier loci) and the morphological differentiation coefficient of genetic differentiation coefficient (QST-FST) comparative analysis to support local adaptation. The conclusion of soil moisture, pH and conductivity can be formed by natural selection pressure adaptability differences of photosynthetic pigments in this research. There are genetic help to small scale heterogeneous habitats with different phenotypes of reed and superficial understanding of the genetic basis, enrich the research content of reed for molecular ecology, ecological adaptability, biological resistance, study on genus and reed, including ecological restoration, protection of genetic resources, and provide scientific reference and theoretical basis of the application of practice, genetic improvement.

【學(xué)位授予單位】：東北師范大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2017
【分類號】：Q948

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