羌活與寬葉羌活的種間基因流和物種分化研究
發(fā)布時(shí)間：2019-03-14 20:17
【摘要】：羌活(Notopterygium incisum)和寬葉羌活(N.franchetii)是鄰域分布于我國(guó)青藏高原及其臨近高山地區(qū)的兩種多年生珍惜瀕危草本植物,兩種植物的物種形成和群體歷史進(jìn)化可能受到了過(guò)去高原隆升以及第四紀(jì)冰期氣候波動(dòng)的影響。本研究取樣兩個(gè)物種高原地區(qū)青海、甘肅、四川及陜西等地的21個(gè)群體共122個(gè)個(gè)體材料為研究對(duì)象,同時(shí)選取羌活屬近緣物種青海當(dāng)歸(Angelica nitida)、東當(dāng)歸(A.acutiloba)和白芷(A.dahurica)作為外類群,利用單親遺傳的葉綠體基因片段和線粒體基因片段以及雙親遺傳的多個(gè)核基因片段作為分子標(biāo)記,通過(guò)群體水平上的DNA測(cè)序檢測(cè)羌活和寬葉羌活自然群體遺傳變異的地理分布模式,分析兩物種的遺傳多樣性和群體結(jié)構(gòu),進(jìn)行探討兩物種間基因流和物種分化歷史,主要結(jié)果如下：(1)葉綠體(cpDNA)和線粒體基因(mtDNA)序列分析得出：選用的一個(gè)葉綠體片段OG28079長(zhǎng)261 bp,分離得到3個(gè)變異位點(diǎn),都集中在羌活中；鑒定到4種單倍型,其中有3種單倍型是羌活的特有單倍型,寬葉羌活只有1種mtDNA單倍型；兩個(gè)線粒體基因(OG537和OG917)長(zhǎng)度總和872 bp,序列中共有3個(gè)變異位點(diǎn),其中1個(gè)在羌活中,2個(gè)在寬葉羌活中,共統(tǒng)計(jì)出4種單倍型,其中有1個(gè)羌活的特有單倍型,2個(gè)寬葉羌活的特有單倍型,并且有1個(gè)單倍型為兩個(gè)物種所共有。在譜系分化和系統(tǒng)發(fā)育分析的研究中,發(fā)現(xiàn)羌活與寬葉羌活物種有緊密的親緣關(guān)系。單倍型地理分布圖顯示,兩個(gè)物種在葉綠體、線粒體和核基因序列上都存在有單倍型共享的現(xiàn)象,且這兩個(gè)物種在青藏高原東南邊緣有重疊的分布區(qū),可能是發(fā)生了二次接觸(Secondary contact)及基因漸滲(Introgression)事件。(2)由8個(gè)核DNA序列的研究發(fā)現(xiàn),羌活和寬葉羌活的基因多態(tài)性水平較低；但寬葉羌活(πT=0.00303;πs=0.00422)的核苷酸多態(tài)性水平略高于羌活(πT= 0.00367;πs=0.00408).AMOVA和Permut分析表明兩個(gè)物種的遺傳變異主要存在于群體間,兩物群體體間的平均分化比率為79.02%,其中羌活群體間的分化比率平均值達(dá)到了89.94%；寬葉羌活群體間的分化比率平均值為75.28%,而群體內(nèi)變異較少。兩物種在所研究的核DNA序列中單倍型種類較多,羌活及寬葉羌活的特有單倍型也較多,且在Network圖中有各自明顯的區(qū)分區(qū)域,兩物種共享單倍型的現(xiàn)象較少,說(shuō)明兩物種間的分化程度較大。(3)應(yīng)用隔離-遷移模型進(jìn)行基因流分析,發(fā)現(xiàn)在羌活和寬葉羌活之間檢測(cè)到了明顯不對(duì)等的雙向基因流,從寬葉羌活遷移至羌活的基因流大于從羌活遷移至寬葉羌活的基因流。分化時(shí)間T=4.635×105年,推測(cè)兩物種進(jìn)化歷史與青藏高原隆升相關(guān)。羌活的平均有效群體大小Ne1=7.824x106,寬葉羌活的有效群體大小Ne2=6.792×106,祖先群體的大小NeA=4.249x107,兩物種的有效群體大小有明顯的收縮；單個(gè)核DNA序列的進(jìn)化時(shí)間相近,由此推測(cè)兩物種的物種形成符合異域物種形成模式,并伴隨近期的二次接觸及基因漸滲事件。同時(shí)相對(duì)于祖先群體,兩物種的群體大小都有明顯的收縮；結(jié)合Mismatch的分析結(jié)果,暗示在羌活和寬葉羌活進(jìn)化歷史上有可能經(jīng)歷過(guò)分布區(qū)反復(fù)收縮及擴(kuò)張的事件。(4)根據(jù)三種基因片段的單倍型地理分布,發(fā)現(xiàn)羌活和寬葉羌活中單倍型多樣性較高的群體大多集中在青海西部阿壩自治州的東部河湟谷地,青藏高原與黃土高原接壤處的互助、大通地區(qū),四川西北部的高海拔山區(qū)及秦嶺山脈的太白山地區(qū)。據(jù)此推測(cè)羌活和寬葉羌活的冰期避難所可能在青藏高原東北部與黃土高原接壤處、青藏高原東部邊緣及秦嶺西部地區(qū)。
[Abstract]:Notopterygium incisum and Notopterygium root (N. franchetii) are two kinds of rare and endangered herb which are distributed in the Qinghai-Tibet plateau and adjacent high-mountain areas of China. The species formation and the evolution of the population history of the two plants may be affected by the uplift of the past plateau and the climatic fluctuation of the Quaternary. In this study,122 individual materials of 21 groups in Qinghai, Gansu, Sichuan and Shaanxi were taken as the research objects in the two species of Qinghai, Gansu, Sichuan and Shaanxi. the geographical distribution mode of the genetic variation of the notopterygium root and the broad-leaf notopterygium root natural population is detected by the DNA sequencing on the population level by using the chloroplast gene fragment and the mitochondrial gene fragment and the plurality of nuclear gene fragments of the parent to be used as the molecular marker, The genetic diversity and population structure of the two species were analyzed, and the history of the gene flow and the species differentiation between the two species was studied. The main results are as follows: (1) the sequence analysis of the chloroplast (cpDNA) and the mitochondrial gene (mtDNA): one of the selected chloroplast fragments OG28079 is 261bp, 3 mutation sites were isolated and concentrated in the notopterygium root;4 single-fold types were identified, among which three single-fold types were the unique haplotypes of the notopterygium, only one mtDNA haplotype was found in the broad-leaf notopterygium, and the total length of the two mitochondrial genes (OG537 and OG917) was 872 bp, and the total number of the two mitochondrial genes (OG537 and OG917) was 872 bp, and there were 3 mutation sites in the sequence. Among them,1 of notopterygium root and 2 were in the notopterygium root of broad leaf, four single-fold types were counted, one of which had the unique haplotype of notopterygium root, the unique haplotype of 2 broad-leaf notopterygium, and one haplotype was common to both species. In the study of lineage differentiation and phylogenetic analysis, it was found that notopterygium root was closely related to the notopterygium root species. The haplotype geographical distribution map shows that the two species have a haplotype-sharing phenomenon in the chloroplast, the mitochondria and the nuclear gene sequence, and the two species have overlapping distribution areas on the southeast edge of the Qinghai-Tibet Plateau. Secondary contact and the progression of the gene may be the case. (2) The genetic polymorphism of notopterygium root and notopterygium root is lower than that of notopterygium root (IGT = 0.00303;% s = 0.00422), which is slightly higher than that of notopterygium root (IGT = 0.00367;% s = 0.00408), according to the eight nuclear DNA sequences. AMOVA and Permut analysis showed that the genetic variation of the two species was mainly present in the group, the average differentiation rate among the two groups was 79.02%, the average value of the differentiation ratio among the notopterygium was 89.94%, and the average value of the differentiation ratio among the notopterygium root groups was 75.28%. And the intra-population variation is less. The single-fold type of the two species in the studied nuclear DNA sequence is more, the unique haplotype of the notopterygium root and the notopterygium root is more, and there is a distinct distinguishing region in the network graph, the phenomenon that the two species share the haplotype is less, and the differentiation degree between the two species is relatively large. And (3) carrying out gene flow analysis by using an isolation-migration model, and finding that a two-way gene flow is detected between the notopterygium root and the notopterygium root and the notopterygium root, and the gene flow from the notopterygium root to the notopterygium root is larger than the gene flow from the notopterygium root to the notopterygium root. The differentiation time T = 4.635-105 years, and the evolution history of the two species is presumed to be related to the uplift of the Qinghai-Tibet Plateau. The mean effective population size of Notopterygium root is Ne1 = 7.824x106, the effective population size of Notopterygium root is Ne2 = 6.792-106, the size of the ancestor group is NeA = 4.249x107, the effective population size of the two species is obviously contracted, and the evolution time of the single nuclear DNA sequence is similar, It is presumed that the species formation of two species is in conformity with the formation pattern of the alien species and is accompanied by the recent secondary contact and the gene gradual infiltration. At the same time, the population size of the two species is obviously contracted with respect to the group of the ancestors, and the results of the analysis of Mismatch suggest that it is possible to experience the repeated contraction and expansion of the distribution area in the history of notopterygium root and notopterygium root. (4) according to the single-type geographical distribution of the three gene fragments, most of the groups of the notopterygium root and the notopterygium root in the broad-leaved notopterygium root are found to be mostly concentrated in the eastern river, the valley of the east river of the Abba autonomous prefecture of the western Qinghai, the mutual aid of the Qinghai-Tibet plateau and the loess plateau of the Loess Plateau, and the Datong area, The high-altitude mountainous area in the northwest of Sichuan and the Taibai Mountain area of the Qinling Mountains. Based on this, it is estimated that the ice age shelters of notopterygium root and notopterygium root may be located in the northeast of the Qinghai-Tibet Plateau and the Loess Plateau, the eastern edge of the Qinghai-Tibet Plateau and the western part of the Qinling Mountains.
【學(xué)位授予單位】：西北大學(xué)
【學(xué)位級(jí)別】：碩士
【學(xué)位授予年份】：2016
【分類號(hào)】：Q943
本文編號(hào)：2440334
[Abstract]:Notopterygium incisum and Notopterygium root (N. franchetii) are two kinds of rare and endangered herb which are distributed in the Qinghai-Tibet plateau and adjacent high-mountain areas of China. The species formation and the evolution of the population history of the two plants may be affected by the uplift of the past plateau and the climatic fluctuation of the Quaternary. In this study,122 individual materials of 21 groups in Qinghai, Gansu, Sichuan and Shaanxi were taken as the research objects in the two species of Qinghai, Gansu, Sichuan and Shaanxi. the geographical distribution mode of the genetic variation of the notopterygium root and the broad-leaf notopterygium root natural population is detected by the DNA sequencing on the population level by using the chloroplast gene fragment and the mitochondrial gene fragment and the plurality of nuclear gene fragments of the parent to be used as the molecular marker, The genetic diversity and population structure of the two species were analyzed, and the history of the gene flow and the species differentiation between the two species was studied. The main results are as follows: (1) the sequence analysis of the chloroplast (cpDNA) and the mitochondrial gene (mtDNA): one of the selected chloroplast fragments OG28079 is 261bp, 3 mutation sites were isolated and concentrated in the notopterygium root;4 single-fold types were identified, among which three single-fold types were the unique haplotypes of the notopterygium, only one mtDNA haplotype was found in the broad-leaf notopterygium, and the total length of the two mitochondrial genes (OG537 and OG917) was 872 bp, and the total number of the two mitochondrial genes (OG537 and OG917) was 872 bp, and there were 3 mutation sites in the sequence. Among them,1 of notopterygium root and 2 were in the notopterygium root of broad leaf, four single-fold types were counted, one of which had the unique haplotype of notopterygium root, the unique haplotype of 2 broad-leaf notopterygium, and one haplotype was common to both species. In the study of lineage differentiation and phylogenetic analysis, it was found that notopterygium root was closely related to the notopterygium root species. The haplotype geographical distribution map shows that the two species have a haplotype-sharing phenomenon in the chloroplast, the mitochondria and the nuclear gene sequence, and the two species have overlapping distribution areas on the southeast edge of the Qinghai-Tibet Plateau. Secondary contact and the progression of the gene may be the case. (2) The genetic polymorphism of notopterygium root and notopterygium root is lower than that of notopterygium root (IGT = 0.00303;% s = 0.00422), which is slightly higher than that of notopterygium root (IGT = 0.00367;% s = 0.00408), according to the eight nuclear DNA sequences. AMOVA and Permut analysis showed that the genetic variation of the two species was mainly present in the group, the average differentiation rate among the two groups was 79.02%, the average value of the differentiation ratio among the notopterygium was 89.94%, and the average value of the differentiation ratio among the notopterygium root groups was 75.28%. And the intra-population variation is less. The single-fold type of the two species in the studied nuclear DNA sequence is more, the unique haplotype of the notopterygium root and the notopterygium root is more, and there is a distinct distinguishing region in the network graph, the phenomenon that the two species share the haplotype is less, and the differentiation degree between the two species is relatively large. And (3) carrying out gene flow analysis by using an isolation-migration model, and finding that a two-way gene flow is detected between the notopterygium root and the notopterygium root and the notopterygium root, and the gene flow from the notopterygium root to the notopterygium root is larger than the gene flow from the notopterygium root to the notopterygium root. The differentiation time T = 4.635-105 years, and the evolution history of the two species is presumed to be related to the uplift of the Qinghai-Tibet Plateau. The mean effective population size of Notopterygium root is Ne1 = 7.824x106, the effective population size of Notopterygium root is Ne2 = 6.792-106, the size of the ancestor group is NeA = 4.249x107, the effective population size of the two species is obviously contracted, and the evolution time of the single nuclear DNA sequence is similar, It is presumed that the species formation of two species is in conformity with the formation pattern of the alien species and is accompanied by the recent secondary contact and the gene gradual infiltration. At the same time, the population size of the two species is obviously contracted with respect to the group of the ancestors, and the results of the analysis of Mismatch suggest that it is possible to experience the repeated contraction and expansion of the distribution area in the history of notopterygium root and notopterygium root. (4) according to the single-type geographical distribution of the three gene fragments, most of the groups of the notopterygium root and the notopterygium root in the broad-leaved notopterygium root are found to be mostly concentrated in the eastern river, the valley of the east river of the Abba autonomous prefecture of the western Qinghai, the mutual aid of the Qinghai-Tibet plateau and the loess plateau of the Loess Plateau, and the Datong area, The high-altitude mountainous area in the northwest of Sichuan and the Taibai Mountain area of the Qinling Mountains. Based on this, it is estimated that the ice age shelters of notopterygium root and notopterygium root may be located in the northeast of the Qinghai-Tibet Plateau and the Loess Plateau, the eastern edge of the Qinghai-Tibet Plateau and the western part of the Qinling Mountains.
【學(xué)位授予單位】：西北大學(xué)
【學(xué)位級(jí)別】：碩士
【學(xué)位授予年份】：2016
【分類號(hào)】：Q943
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