식물분류학회지 (Korean Journal of Plant Taxonomy)

  • 제46권3호
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  • Pages.314-325
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  • 2016
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  • 1225-8318(pISSN)
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  • 2466-1546(eISSN)
한국식물분류학회 (The Korean Society of Plant Taxonomists)
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한국산 방동사니족(사초과) 식물의 분자계통과 광합성경로의 분화

Molecular phylogeny and divergence of photosynthetic pathways of Korean Cypereae (Cyperaceae)

  • 정종덕 (동북아생물다양성연구소) ;
  • 류영일 (동북아생물다양성연구소) ;
  • 최홍근 (아주대학교 자연과학대학 생명과학과)
  • Jung, Jongduk (Northeastern Asia Biodiversity Institute) ;
  • Ryu, Youngil (Northeastern Asia Biodiversity Institute) ;
  • Choi, Hong-Keun (Department of Biological Science, College of Natural Sciences, Ajou University)
  • 투고 : 2016.09.19
  • 심사 : 2016.09.29
  • 발행 : 2016.09.30
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초록

광합성 경로의 전환은 현화식물의 계통에서 여러 번에 걸쳐서 독립적으로 일어난 진화적 사건으로서 사초과에서는 다섯 번 이상 발생한 것으로 추정되고 있다. 방동사니족에서 나타난 C4 광합성 경로로의 전환은 한 번 발생하였으며 이는 방동사니족 내 C4 식물의 공유파생형질로 여겨진다. 방동사니족에 포함된 속들의 형태학적 한계는 분자계통학적 유연관계와 일치하지 않으며, 특히 다계통군으로 여겨지는 방동사니속의 한계는 계통분류학적으로도 논란이 되고 있다. 본 연구에서는 한국산 방동사니족 식물의 광합성 경로와 분자계통을 비교하고자 하였다. 해부학적 관찰을 통해 우리나라 방동사니족 식물 20종(방동사니속 18종, 파대가리속 1종, 세대가리속 1종)의 광합성 경로를 확인하였다. 또한 nrITS, rbcL, trnL-F의 염기서열에 근거하여 각 분류군의 분자계통학적 위치를 파악하고자 하였으며, 분류군 전체의 계통을 파악하기 위하여 선행연구 결과와 함께 분석하였다. 엽록체가 밀집된 광합성 조직의 위치에 따라 우리나라 방동사니속 식물 중 병아리방동사니와 우산방동사니, 모기방동사니, 알방동사니의 네 종은 C3 식물로 확인되었고, 나머지 14종의 방동사니속 식물, 파대가리, 그리고 세대가리는 C4 식물로 결정되었다. 또한 분자계통학적 분석에서 방동사니족은 CYPERUS 분계군과 FICINIA 분계군으로 구분되었으며, 우리나라 방동사니족 식물은 모두 CYPERUS 분계군에 속하였다. CYPERUS 분계군내에서 C4 식물들은 단계통군을 형성하였지만 각 분류군 간의 유연관계는 명확하게 나타나지 않았다. 분자계통수에서 세대가리속과 파대가리속은 C4 식물인 방동사니속 식물들과 함께 단일 분계군을 형성함으로서 각각 독립된 속으로서 지지 되지 않는다. 이는 기존의 연구결과와 일치하며, CYPERUS 분계군에 속한 속들의 계통분류체계를 명확하게 하기 위해서는 면밀한 형태학적 연구와 더불어 높은 해상력을 갖춘 분자계통학적 연구가 이루어져야 할 것이다.

Multiple changes of the photosynthesis pathway are independent evolutionary events occurring in the phylogeny of flowering plants, and such changes have occurred more than five times in Cyperaceae. In the tribe Cypereae, the C4 photosynthetic pathway appeared only once and is regarded as a synapomorphy of the C4 plants within this tribe. The morphological delimitation of genera within Cypereae does not correspond to their molecular phylogenetic relationships. In this study, the molecular phylogeny was compared with the photosynthetic pathways of Korean Cypereae (18 species of Cyperus, 1 species of Kyllinga, and 1 species of Lipocarpha). The photosynthetic pathways were determined by observing the leaf anatomy. The phylogenetic analysis was performed using three DNA regions (nrITS, rbcL, and trnL-F). According to the position of the photosynthetic tissue, 4 species (C. difformis, C. flaccidus, C. haspan, and C. tenuispica) and 16 species (14 Cyperus species, K. brevifolia var. leiolepis, and L. microcephala) were confirmed as C3 and C4 plants, respectively. Tribe Cypereae was divided into the CYPERUS and FICINIA clades, and all species of Korean Cypereae plants belonged to the CYPERUS clade in the phylogenetic analysis. Within the CYPERUS clade, C4 plants were monophyletic but their phylogenetic relationships were unclear. The genera Kyllinga and Lipocarpha were not supported as an independent genus in either case because they were nested by the Cyperus species in the molecular phylogenetic trees in the present and in previous studies. To determine the classification within the CYPERUS clade, a detailed morphological study and a molecular phylogenetic analysis at a high resolution will be necessary.

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참고문헌

  1. Bauters, K., I. Larridon, M. Reynders, P. Asselman, A. Vrijdaghs, A. M. Muasya, D. A. A. Simpson and P. Goetghebeur. 2014. A new classification for Lipocarpha and Volkiella as infrageneric taxa of Cyperus s.l. (Cypereae, Cyperoideae, Cyperaceae): insights from species tree reconstruction supplemented with morphological and floral developmental data. Phytotaxa 166: 1-32. https://doi.org/10.11646/phytotaxa.166.1.1
  2. Besnard, G., A. M. Muasya, F. Russier, E. H. Roalson, N. Salamin and P.-A. Christin. 2009. Phylogenomics of $C_4$ photosynthesis in sedges (Cyperaceae): multiple appearances and genetic convergence. Molecular Biology and Evolution 26: 1909-1919. https://doi.org/10.1093/molbev/msp103
  3. Borchsenius, F. 2009. FastGap 1.2. Department of Biosciences, Aarhus University, Denmark. Retrieved Sep. 19, 2016, from http://www.aubot.dk/FastGap_home.htm.
  4. Bruhl, J. J. 1995. Sedge genera of the world: relationships and a new classification of the Cyperaceae. Australian Systematic Botany 8: 125-305. https://doi.org/10.1071/SB9950125
  5. Chen, D.-H. and P. C. Ronald. 1999. A rapid DNA minipreparation method suitable for AFLP and other PCR applications. Plant Molecular Biology Reporter 17: 53-57. https://doi.org/10.1023/A:1007585532036
  6. Chung, T. 1957. Korean Flora. Shinjisa, Seoul, 1025 pp.
  7. Clarke, C. B. 1908. New genera and species of Cyperaceae. Bulletin of Miscellaneous Information (Royal Gardens, Kew) Additional Series 8: 1-196.
  8. Dengler, N. and T. Nelson. 1999. Leaf structure and development in C4 plants. In C4 Plant Biology. Sage, R. F. and R. K. Monson (eds.), Academic Press, San Diego, CA. Pp. 133-172.
  9. Edgar, R. C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792-1797. https://doi.org/10.1093/nar/gkh340
  10. Goetghebeur, P. 1998. Cyperaceae. In The Families and Genera of Vascular Plants. Vol. 4. Kubitzki, K. (ed.), Springer Verlag, Berlin. Pp. 141-190.
  11. Govaerts, R., D. A. Simpson, P. Goetghebeur, K. L. Wilson, T. Egorova and J. Bruhl. 2007. World Checklist of Cyperaceae: Sedges. Royal Botanical Garden, Kew, 765 pp.
  12. Jung, J. and H.-K. Choi. 2010. Systematic rearrangement of Korean Scirpus L. s.l. (Cyperaceae) as inferred from nuclear ITS and chloroplast rbcL sequences. Journal of Plant Biology 53: 222-232. https://doi.org/10.1007/s12374-010-9109-8
  13. Jung, J. and H.-K. Choi. 2011. Taxonomic study of Korean Scirpus L. s.l. (Cyperaceae) II: Pattern of phenotypic evolution inferred from molecular phylogeny. Journal of Plant Biology 54: 409-424. https://doi.org/10.1007/s12374-011-9181-8
  14. Jung, J. and H.-K. Choi. 2013. Recognition of two major clades and early diverged groups within the subfamily Cyperoideae (Cyperaceae) including Korean sedges. Journal of Plant Research 126: 335-349. https://doi.org/10.1007/s10265-012-0534-2
  15. Kim, I. S., J.-H. Pak, B.-B. Seo and S.-D. Song. 1999. Ultrastructure of leaves in $C_4$ Cyperus iria and $C_3$ Carex siderosticta. Journal of Plant Biology 42: 213-221. https://doi.org/10.1007/BF03030481
  16. Larridon, I. 2011. Exploring giant genera and their satellites, tales from the C3 Cyperus universe. PhD dissertation, Ghent University, Ghent, Belgium, 402 pp.
  17. Larridon, I., M. Reynders, W. Huygh, K. Bauters, A. Vrijdaghs, O. Leroux, A. M. Muasya, D. A. Simpson and P. Goetghebeur. 2011. Taxonomic changes in $C_3$ Cyperus (Cyperaceae) supported by molecular data, morphology, embryography, ontogeny and anatomy. Plant Ecology and Evolution 144: 327-356. https://doi.org/10.5091/plecevo.2011.653
  18. Larridon, I., K. Bauters, M. Reynders, W. Huygh, A. M. Muasya, D. A. Simpson and P. Goetghebeur. 2013. Towards a new classification of the giant paraphyletic genus Cyperus (Cyperaceae): phylogenetic relationships and generic delimitation in $C_4$ Cyperus. Botanical Journal of the Linnean Society 172: 106-126. https://doi.org/10.1111/boj.12020
  19. Lee, K. B. 2004. Plant Anatomy. Lifescience Publishing Co., Seoul, 140 pp.
  20. Lee, T. B. 2003. Coloured Flora of Korea. Hyang-Mun Publishing Co., Seoul, 914 pp.
  21. Lee, W. T. 1996. Lineamenta Florae Koreae. Academy Press, Seoul. Pp. 1695-2383.
  22. Lee, Y. N. 2006. New Flora of Korea. Kyo-Hak Publishing Co., Seoul, Vol. 1, 975 pp, Vol. 2, 885 pp.
  23. Linnaeus, C. 1753. Species Plantarum. Ray Society, London, 176 pp.
  24. Linnaeus, C. 1754. Genera Plantarum. Ray Society, Stockholm, 26 pp.
  25. Muasya, A. M., D. A. Simpson, M. W. Chase and A. Culham. 1998. An assessment of suprageneric phylogeny in Cyperaceae using rbcL DNA sequences. Plant Systematics and Evolution 211: 257-271. https://doi.org/10.1007/BF00985363
  26. Muasya, A. M., D. A. Simpson and M. W. Chase. 2001. Generic relationships and character evolution in Cyperus s.l. (Cyperaceae). Systematics and Geography of Plants 71: 539-544. https://doi.org/10.2307/3668698
  27. Muasya, A. M., D. A. Simpson and M. W. Chase. 2002. Phylogenetic relationships in Cyperus L. s.l. (Cyperaceae) inferred from plastid DNA sequence data. Botanical Journal of the Linnean Society 138: 145-153. https://doi.org/10.1046/j.1095-8339.2002.138002145.x
  28. Muasya, A. M., D. Simpson, G. Verboom, P. Goetghebeur, R. F. C. Naczi, M. W. Chase and E. Smets. 2009a. Phylogeny of Cyperaceae based on DNA sequence data: current progress and future prospects. The Botanical Review 75: 2-21. https://doi.org/10.1007/s12229-008-9019-3
  29. Muasya, A. M., A. Vrijdaghs, D. A. Simpson, M. W. Chase, P. Goetghebeur and E. Smets. 2009b. What is a genus in Cypereae: phylogeny, character homology assessment and generic circumscription in Cypereae. The Botanical Review 75: 52-66. https://doi.org/10.1007/s12229-008-9018-4
  30. Muasya, A. M. and P. J. de Lange. 2010. Ficinia spiralis (Cyperaceae) a new genus and combination for Desmoschoenus spiralis. New Zealand Journal of Botany 48: 31-39. https://doi.org/10.1080/00288251003660703
  31. Nylander, J. A. A. 2004. MrModeltest 2.3. Evolutionary Biology Centre, Uppsala University, Uppsala.
  32. Oh, Y. C., C. S. Lee and K. M. Ko. 1999. Anatomical study of Korean Cyperus Linne. Korean Journal of Plant Taxonomy 29: 325-354. https://doi.org/10.11110/kjpt.1999.29.4.325
  33. Oh, Y. C. and Y. H. Lee. 1999. A systematic study on Cyperus L. of Korea (Cyperaceae). Korean Journal of Plant Taxonomy 29: 37-62. https://doi.org/10.11110/kjpt.1999.29.1.037
  34. Oh, Y. C. 2007. Cyperaceae. In The Genera of Vascular Plantd of Korea. Park, C.-W. (ed.), Academy Publishing Co., Seoul. Pp. 1113-1181.
  35. Ohwi, J. 1944. Cyperaceae Japonicae, II. A synopsis of the Rhynchosporoideae and Scirpoideae of Japan, including the Kuriles, Saghalin, Korea, and Formosa. Memoirs of the College of Science; Kyoto Imperial University Series B, (botany, zoology, geology) biology 18: 1-182.
  36. Ohwi, J. 1965. Flora of Japan. Shibundo, Tokyo, 1067 pp.
  37. Posada, D. and T. R. Buckley. 2004. Model selection and model averaging in phylogenetics: Advantages of Akaike Information Criterion and Bayesian approaches over likelihood ratio tests. Systematic Biology 53: 793-808. https://doi.org/10.1080/10635150490522304
  38. Ronquist, F. and J. P. Huelsenbeck. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572-1574. https://doi.org/10.1093/bioinformatics/btg180
  39. Sage, R. F. 2004. The evolution of $C_4$ photosynthesis. New Phytologist 161: 341-370. https://doi.org/10.1111/j.1469-8137.2004.00974.x
  40. Shobe, W. R. and N. R. Lersten. 1967. A technique for clearing and staining Gymnosperm leaves. Botanical Gazette 128: 150-152. https://doi.org/10.1086/336391
  41. Soros, C. L. and J. J. Bruhl. 2000. Multiple evolutionary origins of C4 photosynthesis in the Cyperaceae. In Monocots: Systematics and Evolution. Wilson, K. L. and D. A. Morrison (eds.), CSIRO Publishing, Melbourne. Pp. 629-635.
  42. Swofford, D. L. 2002. PAUP*: Phylogenetic analysis using parsimony (*and other methods). Ver. 4.0b10. Sinauer Associates Inc., Sunderland, MA.
  43. Taberlet, P., L. Gielly, G. Pautou and J. Bouvet. 1991. Universal primers for amplification of three non-coding regions of chloroplast DNA. Plant Molecular Biology 17: 1105-1109. https://doi.org/10.1007/BF00037152
  44. Tieszen, L. L., M. M. Senyimba, S. K. Imbamba and J. H. Troughton. 1979. The distribution of $C_3$ and $C_4$ grasses and carbon isotope discrimination along an altitudinal and moisture gradient in Kenya. Oecologia 37: 337-350. https://doi.org/10.1007/BF00347910
  45. Tucker, G. C., B. G. Marcks and J. R. Carter. 2002. Cyperus. In Flora of North America North of Mexico. Vol. 23. Flora of North America Editorial Committee (ed.), Oxford University Press, New York. Pp. 141-191.
  46. White, T. J., T. Bruns, S. Lee and J. Taylor. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In PCR Protocols: A Guide to Methods and Applications. Innis, M. A., D. H. Gelfand, J. J. Sninsky and T. J. White (eds.), Academic Press, San Diego, CA, Pp. 315-322.

피인용 문헌

  1. CONTRIBUTIONS TO THE KNOWLEDGE OF CAMBODIAN CYPERACEAE vol.76, pp.2, 2019, https://doi.org/10.1017/s0960428619000015