祁连圆柏的分子谱系地理学研究
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摘要
青藏高原作为全球温度最敏感的地区,在第四纪冰期存在气候反复变化的多次轮回,但是第四纪冰期气候如何影响其动植物的分布以及所导致的生态系统变化?环境的艰苦性导致相关资料和研究积累极少,因而存在各种假说和争论。本论文以DNA序列的群体遗传学标记为基础,重建了该地区一个特有、并且也是该地区残遗森林群落的建群种-祁连圆柏的谱系地理结构。
对20个种群392棵树的两套叶绿体DNA序列(trnT-trnF和trnS-trnG)进行了全部测序,序列变异组合成6种单倍型(A, B, C, D, E和F)。种群间遗传分化较大(GST = 0.772, NST = 0.834),有明显的谱系地理结构(Nst > Gst, P <0.05),同时也表明现存种群间的基因流较小。高原面高海拔9个种群的8个种群单倍型都十分一致,均为A;而最西边的一个种群除这种主要单倍型之外,还发现有少量的单倍型C和F。高原边缘低海拔地区的多数种群,单倍型多样性也低;但是单倍型类型不一样,或为A或为E。其中的一个种群具有固四种单倍型(A、B、D和E),表明该种群所在地或为冰期避难所或为冰期回迁的交汇地。6种单倍型的进化关系表明,单倍型A起源的时间较晚,不是广泛固定于高原面上的起源较早的原始类型。高原边缘丰富的单倍型表明此地区是祁连圆柏的冰期避难所;但由于瓶颈作用,种群内的多样性减少,不同的种群却随机固定了不同的单倍型。高原面上多数种群具有相同的单倍型,表明它们可能是冰期大面积回迁形成的;而这次回迁应该是连续的,而现代间断分布则可能是全新世内人类活动或者其它原因导致高寒草甸生态系统大面积发育形成的。高原最西边种群的独特单倍性,表明其度过了最后的几次冰期。总之,这些结果表明青藏高原东北部全新世内还有一次大面积的森林分布过程,而转换为高原草甸或荒漠生态系统是最近才发生的。
此外,采用最新的磁珠富集法对该种的微卫星位点进行分离,通过含有微卫星位点序列的获得,设计出了12对SSR多态性引物,为该物种遗传多样性的SSR分子标记实验研究奠定了基础。
The Qinghai-Tibetan Plateau (QTP) is the most sensitive to the climatic changes in the world. The Quaternary climatic oscillation must have resulted in changes of the distributional ranges of the organisms occurring there. However, due to the arid environments and fewer accumulations of the related data, it remains unknown how these species survived in situ, retreated and recolonized as well as how the ecological systems changed correspondingly. The contrasting hypotheses had been proposed and no independent and solid evidence has been provided. In this study, based the DNA sequence variation at the population level, phylogeographic patterns were constructed for Juniperus przewalskii Kom., an endemic species to the QTP and a dominant species in the scattered forest communities.
Two cpDNA fragments (trnT-trnF and trnS-trnG) were sequenced for 392 trees of 20 populations through the entire distribution range of the species. Six cpDNA haplotypes were identified (A, B, C, D, E and F). Significant population subdivision was detected (Gst = 0.772, Nst = 0.834), suggesting low levels of recurrent gene flow among populations and significant phylogeographic structure (Nst > Gst, P<0.05). Eight of the nine disjunct populations surveyed on the high-elevation northeast plateau were fixed for a single haplotype (A), while the remaining, more westerly population, contained the same haplotype at high frequency together with two low frequency haplotypes (C and F). In contrast, most populations that occurred at lower altitudes at the plateau edge were fixed or nearly fixed for one of two haplotypes, A or E. However, two plateau edge populations had haplotype compositions different from the rest. In one, four haplotypes (A, B, D and E) were present at approximately equivalent frequencies, which might reflect a larger refugium in the area of this population during the last glacial period. Phylogenetic analysis indicated that the most widely distributed haplotype A is not ancestral to other haplotypes. The edge regions remained as the refugia and these populations randomly fixed different haplotype due to bottleneck effects. Most platform populations probably derived commonly from the postglacial recolonization and their current disjunction occurred within Holocene because of the artificial or other factors. One unique haplotype of one population located in the central plateau suggested that this population survived the later glacial ages. These findings suggested that forests had appeared at a large scale in the northeastern plateau within the Holocene while the current alpine meadow or grasslands probably formed much later since then.
In addition, based on the biotin capture method to enrich microsatellites of J. przewalskii, a total of 12 pairs of SSR primers with polymorphisms were developed.
对20个种群392棵树的两套叶绿体DNA序列(trnT-trnF和trnS-trnG)进行了全部测序,序列变异组合成6种单倍型(A, B, C, D, E和F)。种群间遗传分化较大(GST = 0.772, NST = 0.834),有明显的谱系地理结构(Nst > Gst, P <0.05),同时也表明现存种群间的基因流较小。高原面高海拔9个种群的8个种群单倍型都十分一致,均为A;而最西边的一个种群除这种主要单倍型之外,还发现有少量的单倍型C和F。高原边缘低海拔地区的多数种群,单倍型多样性也低;但是单倍型类型不一样,或为A或为E。其中的一个种群具有固四种单倍型(A、B、D和E),表明该种群所在地或为冰期避难所或为冰期回迁的交汇地。6种单倍型的进化关系表明,单倍型A起源的时间较晚,不是广泛固定于高原面上的起源较早的原始类型。高原边缘丰富的单倍型表明此地区是祁连圆柏的冰期避难所;但由于瓶颈作用,种群内的多样性减少,不同的种群却随机固定了不同的单倍型。高原面上多数种群具有相同的单倍型,表明它们可能是冰期大面积回迁形成的;而这次回迁应该是连续的,而现代间断分布则可能是全新世内人类活动或者其它原因导致高寒草甸生态系统大面积发育形成的。高原最西边种群的独特单倍性,表明其度过了最后的几次冰期。总之,这些结果表明青藏高原东北部全新世内还有一次大面积的森林分布过程,而转换为高原草甸或荒漠生态系统是最近才发生的。
此外,采用最新的磁珠富集法对该种的微卫星位点进行分离,通过含有微卫星位点序列的获得,设计出了12对SSR多态性引物,为该物种遗传多样性的SSR分子标记实验研究奠定了基础。
The Qinghai-Tibetan Plateau (QTP) is the most sensitive to the climatic changes in the world. The Quaternary climatic oscillation must have resulted in changes of the distributional ranges of the organisms occurring there. However, due to the arid environments and fewer accumulations of the related data, it remains unknown how these species survived in situ, retreated and recolonized as well as how the ecological systems changed correspondingly. The contrasting hypotheses had been proposed and no independent and solid evidence has been provided. In this study, based the DNA sequence variation at the population level, phylogeographic patterns were constructed for Juniperus przewalskii Kom., an endemic species to the QTP and a dominant species in the scattered forest communities.
Two cpDNA fragments (trnT-trnF and trnS-trnG) were sequenced for 392 trees of 20 populations through the entire distribution range of the species. Six cpDNA haplotypes were identified (A, B, C, D, E and F). Significant population subdivision was detected (Gst = 0.772, Nst = 0.834), suggesting low levels of recurrent gene flow among populations and significant phylogeographic structure (Nst > Gst, P<0.05). Eight of the nine disjunct populations surveyed on the high-elevation northeast plateau were fixed for a single haplotype (A), while the remaining, more westerly population, contained the same haplotype at high frequency together with two low frequency haplotypes (C and F). In contrast, most populations that occurred at lower altitudes at the plateau edge were fixed or nearly fixed for one of two haplotypes, A or E. However, two plateau edge populations had haplotype compositions different from the rest. In one, four haplotypes (A, B, D and E) were present at approximately equivalent frequencies, which might reflect a larger refugium in the area of this population during the last glacial period. Phylogenetic analysis indicated that the most widely distributed haplotype A is not ancestral to other haplotypes. The edge regions remained as the refugia and these populations randomly fixed different haplotype due to bottleneck effects. Most platform populations probably derived commonly from the postglacial recolonization and their current disjunction occurred within Holocene because of the artificial or other factors. One unique haplotype of one population located in the central plateau suggested that this population survived the later glacial ages. These findings suggested that forests had appeared at a large scale in the northeastern plateau within the Holocene while the current alpine meadow or grasslands probably formed much later since then.
In addition, based on the biotin capture method to enrich microsatellites of J. przewalskii, a total of 12 pairs of SSR primers with polymorphisms were developed.
引文
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