물부추속($Iso{\ddot{e}}tes$ L.)은 물부추과($Iso{\ddot{e}}taceae$)에 속하는 이형포자성을 보이는 다년생 정수성 수생식물로, 전 세계에 200여종이 분포하는 것으로 알려져 있다. 물부추속 식물은 고생대 말기에 출현하여 오랜 진화적인 역사를 지닌다. 다양한 생육환경에서 광범위하게 분포하고, 분포 지역에서는 많은 종들이 높은 고유성을 보임으로서 멸종위기종으로 보호되고 있다. 오랜 종분화 과정에서 극도의 수렴진화와 자가배수체 형성과정을 거치면서 형태적으로 매우 단순화되었다. 이로 인하여 이 식물군의 형태적인 형질을 이용한 계통학적 연구와 유연관계의 규명에 많은 어려움을 보여주고 있다. 본 연구에서는 분자계통학적 마커를 이용하여 극동아시아에 분포하는 물부추속의 계통학적 유연관계를 파악하고, 분자시계를 이용하여 이들의 식물지리학적 기원 및 분화시기 등에 대해서 알아보고자 하였다. 분자마커로서 핵과엽록체 DNA의 염기서열을 이용한 분자계통학적 연구결과, 동아시아 물부추속은 크게 두 개의 분계군으로 구분된다: 일본 홋카이도에 분포하는 북방계분계군과 나머지 물부추속 식물을 포함하는 동아시아 분계군으로 구분이 된다. 북방계인 아시아물부추($Iso{\ddot{e}}tes$ asiatica)는 극동러시아와 북미의 북서부지역의 물부추속 식물과 깊은 유연관계를 보인다. 이 분계군은 북미의 알래스카 지역에서 베링육교(Bering land bridge)를 통해 중신세후기(late Miocene)에 시베리아로 전파된 것으로 분석되었다. 나머지 동아시아 물부추속 식물분계군($Iso{\ddot{e}}tes$ sinensis, I. yunguiensis, I. hypsophila, I. orientalis, I. japonica, I. coreana, I. taiwanensis, I. jejuensis, I. hallasanensis)은 파푸아뉴기니아와 호주의 물부추속 식물과 밀접한 유연관계를 보인다. 이들은, 점신세 후기(late Oligocene)에 호주 대륙의 동부 지역으로부터 원거리 산포과정(long-distance dispersal)을 통해 이동되어진 것으로 추론되었다. 향후에 차세대 염기서열 분석(next generation sequencing)과 같은 대규모 유전자 분석법을 이용하여 유용한 분자마커들을 개발하게 되면 전 세계에 분포하는 물부추속 식물에 대한 전반적인 계통지리학적 분석과 각 대륙에 고유종으로 분포하고 있는 이들의 진화적인 역사를 규명할 수 있을 것으로 보인다.
물부추속($Iso{\ddot{e}}tes$ L.)은 물부추과($Iso{\ddot{e}}taceae$)에 속하는 이형포자성을 보이는 다년생 정수성 수생식물로, 전 세계에 200여종이 분포하는 것으로 알려져 있다. 물부추속 식물은 고생대 말기에 출현하여 오랜 진화적인 역사를 지닌다. 다양한 생육환경에서 광범위하게 분포하고, 분포 지역에서는 많은 종들이 높은 고유성을 보임으로서 멸종위기종으로 보호되고 있다. 오랜 종분화 과정에서 극도의 수렴진화와 자가배수체 형성과정을 거치면서 형태적으로 매우 단순화되었다. 이로 인하여 이 식물군의 형태적인 형질을 이용한 계통학적 연구와 유연관계의 규명에 많은 어려움을 보여주고 있다. 본 연구에서는 분자계통학적 마커를 이용하여 극동아시아에 분포하는 물부추속의 계통학적 유연관계를 파악하고, 분자시계를 이용하여 이들의 식물지리학적 기원 및 분화시기 등에 대해서 알아보고자 하였다. 분자마커로서 핵과 엽록체 DNA의 염기서열을 이용한 분자계통학적 연구결과, 동아시아 물부추속은 크게 두 개의 분계군으로 구분된다: 일본 홋카이도에 분포하는 북방계분계군과 나머지 물부추속 식물을 포함하는 동아시아 분계군으로 구분이 된다. 북방계인 아시아물부추($Iso{\ddot{e}}tes$ asiatica)는 극동러시아와 북미의 북서부지역의 물부추속 식물과 깊은 유연관계를 보인다. 이 분계군은 북미의 알래스카 지역에서 베링육교(Bering land bridge)를 통해 중신세후기(late Miocene)에 시베리아로 전파된 것으로 분석되었다. 나머지 동아시아 물부추속 식물분계군($Iso{\ddot{e}}tes$ sinensis, I. yunguiensis, I. hypsophila, I. orientalis, I. japonica, I. coreana, I. taiwanensis, I. jejuensis, I. hallasanensis)은 파푸아뉴기니아와 호주의 물부추속 식물과 밀접한 유연관계를 보인다. 이들은, 점신세 후기(late Oligocene)에 호주 대륙의 동부 지역으로부터 원거리 산포과정(long-distance dispersal)을 통해 이동되어진 것으로 추론되었다. 향후에 차세대 염기서열 분석(next generation sequencing)과 같은 대규모 유전자 분석법을 이용하여 유용한 분자마커들을 개발하게 되면 전 세계에 분포하는 물부추속 식물에 대한 전반적인 계통지리학적 분석과 각 대륙에 고유종으로 분포하고 있는 이들의 진화적인 역사를 규명할 수 있을 것으로 보인다.
$Iso{\ddot{e}}tes$ L. ($Iso{\ddot{e}}taceae$) is a cosmopolitan genus of heterosporous lycopods containing ca. 200 species being found in lakes, streams, and wetlands of terrestrial habitats. Despite its ancient origin, worldwide distribution, and adaptation to diverse environm...
$Iso{\ddot{e}}tes$ L. ($Iso{\ddot{e}}taceae$) is a cosmopolitan genus of heterosporous lycopods containing ca. 200 species being found in lakes, streams, and wetlands of terrestrial habitats. Despite its ancient origin, worldwide distribution, and adaptation to diverse environment, species in $Iso{\ddot{e}}tes$ show remarkable morphological simplicity and convergence. Allopolyploidy appears to be a significant speciation process in the genus. These characteristics have made it difficult to assess the phylogenetic relationships and biogeographic history of $Iso{\ddot{e}}tes$ species. In recent years, these difficulties have somewhat been reduced by employing multiple molecular markers. Here, we reconstruct the phylogenetic relationships in East Asian $Iso{\ddot{e}}tes$ species. We also provide their divergence time and biogeographic origin using a fossil calibrated chronogram. East Asian $Iso{\ddot{e}}tes$ species are divided into two clades: I. asiatica and the remaining species. $Iso{\ddot{e}}tes$ asiatica from Hokkaido forms a clade with northeastern Russian and western North American $Iso{\ddot{e}}tes$ species. In clade I, western North America is the source area for the dispersal of $Iso{\ddot{e}}tes$ to Hokkaido and northeastern Russia via the Bering land bridge during the late Miocene. The remaining $Iso{\ddot{e}}tes$ species (I. sinensis, I. yunguiensis, I. hypsophila, I. orientalis, I. japonica, I. coreana, I. taiwanensis, I. jejuensis, I. hallasanensis) from East Asia form a sister group to Papua New Guinean and Australian species. The biogeographic reconstruction suggests an Australian origin for the East Asian species that arose through long-distance dispersal during the late Oligocene.
$Iso{\ddot{e}}tes$ L. ($Iso{\ddot{e}}taceae$) is a cosmopolitan genus of heterosporous lycopods containing ca. 200 species being found in lakes, streams, and wetlands of terrestrial habitats. Despite its ancient origin, worldwide distribution, and adaptation to diverse environment, species in $Iso{\ddot{e}}tes$ show remarkable morphological simplicity and convergence. Allopolyploidy appears to be a significant speciation process in the genus. These characteristics have made it difficult to assess the phylogenetic relationships and biogeographic history of $Iso{\ddot{e}}tes$ species. In recent years, these difficulties have somewhat been reduced by employing multiple molecular markers. Here, we reconstruct the phylogenetic relationships in East Asian $Iso{\ddot{e}}tes$ species. We also provide their divergence time and biogeographic origin using a fossil calibrated chronogram. East Asian $Iso{\ddot{e}}tes$ species are divided into two clades: I. asiatica and the remaining species. $Iso{\ddot{e}}tes$ asiatica from Hokkaido forms a clade with northeastern Russian and western North American $Iso{\ddot{e}}tes$ species. In clade I, western North America is the source area for the dispersal of $Iso{\ddot{e}}tes$ to Hokkaido and northeastern Russia via the Bering land bridge during the late Miocene. The remaining $Iso{\ddot{e}}tes$ species (I. sinensis, I. yunguiensis, I. hypsophila, I. orientalis, I. japonica, I. coreana, I. taiwanensis, I. jejuensis, I. hallasanensis) from East Asia form a sister group to Papua New Guinean and Australian species. The biogeographic reconstruction suggests an Australian origin for the East Asian species that arose through long-distance dispersal during the late Oligocene.
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가설 설정
In addition, in Isoëtes, speciation by polyploidization often occurs after hybridization. It is necessary to resolve the controversy as to whether the evolution of megaspores can be interpreted as a result of speciation by hybridization or whether it is associated with the habitat environment. Furthermore, it is necessary to estimate the timing of the evolution of spores.
제안 방법
In order to understand the phylogeny and evolutionary histories of Isoëtes plants distributed around the world, it is necessary to select representative taxa from each clade and analyze the structures and complete nucleotide sequences of the chloroplast genomes using the NGS technique. It is also necessary to understand the precise divergence time and biogeographic history of each clade through a molecular clock analysis (Fig. 5).
It will be necessary to confirm the dispersion mechanism of East Asian Isoëtes by comparing the results of biogeographical studies between Australia and East Asia with the migration routes of migratory birds (Fig. 5).
이론/모형
7). Ancestral states and distribution patterns were estimated by the Bayesian binary method of S-DIVA. The bar and pie graphs on each node indicate the distribution pattern and probability of the ancestral state, respectively.
성능/효과
2) and sister to the group consisting of the rest of the Isoëtes species. (2) Species distributed in South America (I. panamensis, I. cubana, I. jamaicensis) form a clade with Indian species (I. coromandelina var. coromandelina) and some species in Australia and Southeast Asia (I. australis, I. coromandelina var. macrotuberculata, I. laosiensis) (clade II). (3) The remaining Isoëtes species also form a clade (clade III).
However, this study not only performed a DNA analysis with limited samples but also had a very low resolution of the phylogenetic relationships among the lineages in the East Asia/Australia clades.
The first clade included I. taiwanensis, I. coreana (A, B), I. jejuensis, I. hallasanensis, I. coreana (A), and I. sinensis (A).
Therefore, the parental species of I. coreana (6 ×) in South Korea were I. taiwanensis (2 ×) and I. hallasanensis (4 ×), while those of I. japonica (6 ×) in Japan were I. taiwanensis and I. sinensis (4 ×) according to the tree (Fig. 4).
후속연구
In this paper, we intend to review the phylogenetic relationships among Isoëtes species in East Asia and present their biogeographical origins.
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