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Phytochemical variation of Quercus mongolica Fisch. ex Ledeb. and Quercus serrata Murray (Fagaceae) in Mt. Jiri, Korea - Their taxonomical and ecological implications -

지리산 신갈나무와 졸참나무의 식물화학적 변이 양상 - 분류학적, 생태학적 의미 -

  • Received : 2014.09.29
  • Accepted : 2014.10.21
  • Published : 2014.10.31

Abstract

In this study, vertical distribution patterns of Quercus mongolica Fisch. ex Ledeb. and Q. serrata Murray in Korea were recognized and possibility of introgressive hybridization and gene flow between Q. mongolica and Q. serrata in Mt. Jiri was inferred by flavonoid analyses. The most critical factor on distribution patterns was the altitude in accordance with temperature condition. A zonal distribution was recognized: Quercus mongolica zone in the upper area and Q. serrata zone in the lower area. In Central Korea, the range of vertical distribution of Q. mongolica was above alt. 100m, almost everywhere, whereas that of Q. serrata was from alt. 0 m to alt. 500(-700) m, and the species is rare above that altitude. But in Southern Korea, Q. serrata is found up to above alt. 1,000 m, whereas frequency of Q. mongolica reduces as elevation in decline and the species is rare below alt. 300 m, even though pure stands being formed on higher mountain slope. Altitudinal distribution of the two species, however, overlaps, where the two species occur together. Thirty-seven individuals of Q. mongolica and Q. serrata in Mt. Jiri and other area were examined for leaf flavonoid constituents. Twenty-three flavonoid compounds were isolated and identified; they were glycosylated derivatives of the flavonols kaempferol, quercetin, isorhamnetin, myricetin, and four compounds among the flavonoid compounds were acylated. Kaempferol 3-O-glucoside, quercetin 3-O-glucoside, quercetin 3-O-galactoside and its acylated compounds were major constituents and present in all individuals. Quercus mongolica is distinguished from Q. serrata by the presence of quercetin 3-O-arabinosylglucoside and by high concentration of three acylated compounds, acylated kaempferol 3-O-glucoside, quercetin 3-O-glucoside, quercetin 3-O-galactoside, and by relatively low concentration or lacking of rhamnosyl flavonol compounds. There are intraspecific variations in flavonoid profiles for Q. mongolica and Q. serrata, the flavonoid profiles for individuals of two species in hybrid zone (sympatric zone) tend to be similar to each other, qualitatively and quantitatively. These findings strongly suggest that gene exchange or gene flow occurs through the introgressive hybridization between Q. mongolica and Q. serrata in Mt. Jiri. Therefore, Quercus crispula, occupying morphologically intermediate position between Q. mongolica and Q. serrata, is suspected of being a hybrid taxon of two putative parental species.

본 연구에서는 우리나라 신갈나무(Quercus mongolica Fisch. ex Ledeb.)와 졸참나무(Q. serrata Murray) 두 종의 수직분포 양상을 관찰하고, 지리산 지역을 중심으로 두 종간의 교잡이입 및 유전자 전달 가능성을 식물화학적 분석을 통해 추론하고자 하였다. 우리나라의 신갈나무와 졸참나무의 수직분포는 위도에 따라 지역 간 차이가 난다. 중부지방에서는 신갈나무가 해발 100~200m의 낮은 고도에서부터 고재대에 이르기까지 널리 분포하나 남부지방의 경우 일반적으로 해발 300m 이하 저지대에서는 거의 분포하지 않으며, 졸참나무는 중부지방의 경우 저지대에서 주로 관찰되며 해발 500~700m이상에서는 거의 발견되지 않으나 남부지방의 경우 해발 1,000m이상에서도 관찰된다. 두 종은 분포대가 달라 신갈나무는 주로 높은 해발고도에서 졸참나무는 주로 낮은 해발고도에서 생육하나, 상당한 범위의 고도 구간에서 두 종은 혼생한다. 지리산 지역을 위주로 설악산, 소백산, 마니산 등에서 채집된 신갈나무와 졸참나무의 잎 플라보노이드 성분을 분석한 결과, 2종 37개체로부터 총 23종류의 서로 다른 화합물이 분리, 동정되었다. 이들 플라보노이드 화합물은 flavonol인 kaempferol, quercetin, myricetin 및 isorhamnetin에 당이 결합된 flavonol glycoside이었으며, 4 종류의 acylated flavonoid compound가 동정되었다. 이들 중 kaempferol 3-O-glucoside, quercetin 3-O-glucoside와 quercetin 3-O-galactoside 및 이들의 acylated compounds가 주요 성분으로 두 종의 모든 개체에서 나타났다. 신갈나무의 플라보노이드 조성은 졸참나무에서는 나타나지 않는 diglycoside인 quercetin 3-O-arabinosylglucoside가 분포하며, acylated compound인 acylated kaempferol 3-O-glucoside, acylated quercetin 3-O-galactoside 및 acylated quercetin 3-O-glucoside가 다량 분포한다는 점에서 졸참나무의 flavonoid 조성과 구분된다. 졸참나무의 flavonoid 조성은 3개의 rhamnosyl flavonol compounds가 전체 졸참나무 개체에 걸쳐서 나타나며 또한 신갈나무에 비해 다량으로 나타나고, diglycoside인 kaempferol 3-O-rhamnosylglucoside를 함유하는 특징을 갖는다. 두 종 개체들의 flavonoid 조성은 고도에 따라 종내 개체 간 변이가 있었으며, 동소적으로 분포하는 두 종의 개체들은 대체로 상대 종의 플라보노이드 조성을 정량적으로 또는 정성적으로 닮는 경향이 있었다. 이러한 사실은 지리산 지역에서 두 종간에 교잡이입을 통한 유전자 교환이 일어나고 있음을 강하게 암시한다. 이와 같은 상호 교배 및 교잡이입 가능성으로 볼 때, 형태적으로 신갈나무와 졸참나무의 중간적인 특징을 나타내는 물참나무는 두 종을 부모종으로 하는 교잡에 의해 생긴 잡종분류군일 가능성이 높은 것으로 사료된다.

Keywords

References

  1. Alston, R.E. and B.L. Turner(1963) Natural hybridization among four species of Baptisia (Leguminosae). Amer. J. Bot. 50: 159-173. https://doi.org/10.2307/2439849
  2. Bacilieri, R., A. Ducousso, R.J. Petit and A. Kremer(1996) Mating system and asymmetric hybridization in a mixed stand of European oaks. Evolution 50: 900-908. https://doi.org/10.2307/2410861
  3. Camus, A.(1936-1954) Les chenes. Monographie du genre Quercus (et Lithocarpus). Encyclopedie Economique de Sylviculture. Vols. 6-8. Paul Lechevalier, Paris. (in French).
  4. Chang, C.S.(2007) Fagaceae. In: C.W. Park(ed.), The Genera of Vascular Plants of Korea, Academy Publishing Co., Seoul, pp. 268-274.
  5. Chang, J.S. and T. Lee(1984) A biosystematic study on natural populations of Quercus mongolica Fisch. in Korea and Japan. Korean J. Pl. Taxon. 14(2): 71-85. https://doi.org/10.11110/kjpt.1984.14.2.071
  6. Choung, Y.S.(1998) Vegetation in the Paekdoo Great Mountain Chain. Preservation Nature 103: 48-54. (in Korean with English abstract).
  7. Chung, T.H.(1957) Korean Flora Vol. 1. Woody Plants. Sinjisa, Seoul, pp. 87-106. (in Korean).
  8. Chung, T.H. and W.C. Lee(1965) A study of the Korean woody plant zone and favorable region for the growth and proper species. Thesis Collect. Sungkyunkwan Univ. 10: 329-435. (in Korean).
  9. Chybicki, I. J. and J. Burczyk(2010) Realized gene flow within mixed stands of Quercus robur L. and Q. petraea (Matt.) L. revealed at the stage of naturally established seedling. Mol. Ecol. 19: 2137-2151. https://doi.org/10.1111/j.1365-294X.2010.04632.x
  10. Ducousso, A., H. Michaud and R. Lumaret(1993) Repro-duction and gene flow in the genus Quercus L. Ann. For. Sci. 50: 91-106. https://doi.org/10.1051/forest:19930107
  11. Gornall, R.J. and B.A. Bohm(1978) Angiosperm flavonoid evolution: A reappraisal. Syst. Bot. 3: 353-368. https://doi.org/10.2307/2418744
  12. Grant, V.(1981) Plant Speciation. Ed. 2. Columbia Univ. Press, New York.
  13. Hardin, J.W.(1975) Hybridization and introgression in Quercus alba. J. Arnold Arbor. 56: 336-363.
  14. Hattori, S.(1962) Glycosides of flavones and flavonols glycosides. In: T.A. Geissman(ed.), The Chemistry of Flavonoid Compounds, MacMillan, New York, pp. 317-352.
  15. Hong, Y.S., Y.H. You and H.B. Yi(2010) Seasonal change of macro nutrients concentration in acorns of six oak species in Korea. Korean J. Environ. Ecol. 24: 286-292. (in Korean).
  16. Huang, C., Y. Zhang and B. Bartholomew(1999) Fagaceae. In: Z.Y. Wu and P.H. Raven(eds.), Flora of China. Vol. 4. Cycadaceae through Fagaceae, Missouri Botanical Garden Press, St. Louis, pp. 314-400.
  17. Hutchinson, J.(1967) The Genera of Flowering Plants. Vol. 2. Clarendon press, Oxford, pp. 126-132.
  18. Ishida, T., K. Hattori, H. Sato and M. Kimura(2003) Differentiation and hybridization between Quercus crispula and Q. dentata (Fagaceae): insights from morphological traits, amplified fragment length polymorphism markers, and leafminer composition. Amer. J. Bot. 90(5): 769-776. https://doi.org/10.3732/ajb.90.5.769
  19. Isobe, T., N. Ito and Y. Noda(1980) Minor flavonoids of Polygonum nodosum. Phytochemistry 19: 1987.
  20. Jeong, B.K. and C.H. Oh(2013) Analysis on the community structure of Quercus mongolica Fisch. ex Ledeb. in the Baekdudaegan Mountains by elevation - between Hyangnobong and Gitdaebaggybong -. Korean J. Environ. Ecol. 27(4): 449-461. (in Korean).
  21. Kim, J.U. and B.S. Kil(2000) The Mongolian Oak Forest in Korea. Wonkwang National University Press, Iksan, Korea. (in Korean).
  22. Kim, J.W.(1992). Vegetation of Northeast Asia: On the syntaxonomy and syngeography of the oak and beech forests. Ph. D. Thesis, University of Vienna, Vienna. (in English with Korean, Japanese, Chinese, and German summary).
  23. Kim J.W. and Lee Y.K.(2006) Classification and Assessment of Plant Communities. World science, Seoul.
  24. Kim, M.H., J.H. Park and C.W. Park(2000a) Flavonoid chemistry of Fallopia sect. Fallopia (Polygonaceae). Biochem. Syst. Ecol. 28: 433-441. https://doi.org/10.1016/S0305-1978(99)00084-8
  25. Kim, M.H., H.S. Won, J.H. Park and C.W. Park(2000b) Flavonoid chemistry and chromosome number of Fallopia section Pleuropterus (Polygonaceae). Can. J. Bot. 78: 1136-1143.
  26. Kim, Y.H.(2013) Spatial distribution patterns of seven summer-green Quercus species in Korea. M.S. Thesis, Keimyung Univ., Daegu, Korea. (in Korean with the English abstract).
  27. Kim, Y.S., S.C. Ko and B.U. Oh(1981) Distribution Atlas of Plants in Korea(V). Atlas of Fagaceae in Korea. Korea University Press, Seoul, pp. 93-133. (in Korean with English abstract).
  28. Korea Forest Research Institute(1988) The research about use development of the oak resource. Ministry of Science and Technology, pp. 226. (in Korean).
  29. Korea Forest Service(2012) Natural Resources Change Survey and Management Practice Study of the Baekdudaegan Mountains. pp. 52-55. (in Korean)
  30. Kubitzki, K.(1993) Fagaceae. In: K. Kubitzki, J.G. Rohwer and V. Bittrich(eds.), The Families and Genera of Vascular Plants. Vol. II. Flowering plants. Dicotyledons: Mongoliid, Hamamelid and Caryophyllid families, Springer-Verlag, Berlin, Heidelberg, Germany, pp. 301-309.
  31. Lee, C.S., W.K. Lee, J.H. Yoon and C.C. Song(2006) Distribution pattern of Pinus densiflora and Quercus spp. stand in Korea using Spatial Statistics and GIS. J. Korean Forest. Soc. 95(6): 663-671. (in Korean).
  32. Lee, J.H. and H. Hashizume(2004) Currence of Trees of the Quercus fabri type in open-pollinated families of Quercus dentata. J. Korean Forest. Soc. 93: 471-476. (in Korean with the English abstract).
  33. Lee, M.J. 2007. Community structure analysis and ecological planting model subject of the principal Quercus community in Korea. Ph. D. Thesis, Chungnam National University, Daejeon, Korea. (in Korean).
  34. Lee, S.H. and Y.H. You(2012) Measurement of ecological niche of Quercus aliena and Q. serrata under environmental factors treatments and its meaning to ecological distribution. J. Ecol. Field Biol. 35(3): 227-234. https://doi.org/10.5141/JEFB.2012.027
  35. Lee, T.B.(1956) Chromosome numbers in Quercus. Commemoration Theses, Fiftieth Anniversary, College of Agriculture, Seoul Nat. Univ., pp. 35-41.
  36. Lee, T.B.(1961a) Phylogenetic study of the subgenus Lepidobalanus of the genus Quercus in Korea (1). Res. Bull. Korean Agr. Soc. 7: 87-108. (in Korean).
  37. Lee, T.B.(1961b) Phylogenetic study of the subgenus Lepidobalanus of the genus Quercus in Korea (2). Seoul Univ. J., Biol. Ser. 10: 97-141. (in Korean).
  38. Lee, T.B.(1961c) Triple hybird of the oak in Korea. Korean J. Bot. 4: 16-20. (in Korean).
  39. Lee, T.B.(1964) Trichome types of Q. acutissima$\times$ Q. variabilis Nakai. Journal of the Suwon Forestry Society 5: 12-14. (in Korean).
  40. Lee, T.B.(1966) Illustrated Woody Plants of Korea. Forest Experiment Station, Seoul, pp. 31-37, 250-253. (in Korean).
  41. Lee, T.B.(1980) Illustrated Flora of Korea. Hyangmunsa, Seoul, pp. 274-280. (in Korean).
  42. Lee, W.T.(1996) Lineamenta Florae Koreae. Vol. 1. Academy Publishing Co., Seoul, pp. 172-181. (in Korean).
  43. Lee, Y.N.(1996) Flora of Korea. Kyohak Publishing Co., Seoul, pp. 66-70. (in Korean).
  44. Levy, M. and D.A. Levin(1971) The origin of novel flavonoids in Phlox allotetraploids. Proc. Natl. Acad. U. S. A. 68: 1627-1630. https://doi.org/10.1073/pnas.68.7.1627
  45. Mabberley, D.J.(1987) The Plant Book. Cambridge Univ. Press, Cambridge.
  46. Mabry, T.J., K.R. Markham and M.B. Thomas(1970) The Systematic Iidentification of Flavonoids. Springer-Verlag, New York.
  47. MacDougal, D.T.(1907) Hybridization of wild plants. Bot. Gaz. 43: 45-58. https://doi.org/10.1086/329077
  48. Maleev, V. P.(1985) Genus 367. Quercus L. In: V. L. Komarov (ed.), Flora of the U.S.S.R. Vol. 5, Koeltz Scientific Books, Koenigstein, Germany, pp. 254-279.
  49. Markham, K.R.(1982) Techniques of Flavonoid Identification. Academic Press, New York.
  50. Melchior, H.(1964) Reihe Fagales. In: H. Melchior(ed.), A. Engler's Syllabus der Pflanzenfamilien. Vol. 2. Angiospermen, 12th ed., Gebruder Borntraeger, Berlin, pp. 44-51. (in German).
  51. Mun, J.H. and C.W. Park(1995) Flavonoid chemistry of Polygonum sect. Tovara (Polygonaceae): A systematic survey. Pl. Syst. Evol. 196: 153-159. https://doi.org/10.1007/BF00982956
  52. Nakai, T.(1926) Notulae ad plantas Japoniae et Coreae. 31. Bot. Mag. (Tokyo) 40: 161-171. https://doi.org/10.15281/jplantres1887.40.161
  53. Nixon, K.C.(1993) Infrageneric classification of Quercus (Fagaceae) and typification of sectional names. Ann. Sci. Forest. 50(Suppl. 1): 25s-34s. https://doi.org/10.1051/forest:19930701
  54. Nixon, K.C.(1997) Fagaceae. In: Flora of North America Editorial Committee(ed.), Flora of North America. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae, Oxford University Press, New York, pp. 436-506.
  55. Palmer, E.J.(1948) Hybrid oaks of North America. J. Arnold Arbor. 29: 1-48.
  56. Park, C.W.(1987) Flavonoid chemistry of Polygonum sect. Echinocaulon: A systematic survey. Syst. Bot. 12: 167-179. https://doi.org/10.2307/2419228
  57. Park, H.J.(1984) Studies on the distribution and ecological characteristics of the major Quercus spp. in the middle temperate zone in Korea. M.S. Thesis, Chungnam National University, Daejeon, Korea. (in Korean with the English abstract).
  58. Park, J.H.(2009) A systematic study of Quercus L. subgenus Quercus (Fagaceae) in Korea: Analyses of morphological variation and flavonoid chemistry. Ph. D. Thesis, Seoul National Univ. Seoul, pp. 282. (in Korean with the English abstract).
  59. Park, J.H., M.G. Chung, B.Y. Sun, K.J. Kim, J.H. Pak and C.W. Park(2005) Numerical analysis of Quercus L. subgenus Quercus (Fagaceae) in Korea. Korean J. Pl. Taxon. 35: 57-80. (in Korean with the English abstract). https://doi.org/10.11110/kjpt.2005.35.1.057
  60. Prantl, K.(1889) Fagaceae. In: A. Engler and K. Prantl(eds.), Die natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten insbesondere den Nutzpflanzen, bearbeitet unter Mitwirkung zahlreicher hervorragender Fachgelehrten. III. Teil. 1. Abteilung, Verlag von Wilhelm Engelmann, Leipzig, pp. 47-58. (in German).
  61. Schneider, C.K.(1906) Illustriertes Handbuch der Laubholzkunde. Verlag von Gustav Fischer, Jena, Germany, pp. 161-211. (in German).
  62. Sim, J.S. and S.S. Han(2003) Ecophysiological characteristics of deciduous oak species (3): photosynthetic responses of leaves to change of light intensity. J. Korean Forest. Soc. 92: 208-214. (in Korean).
  63. Song, M.S.(2007) Analysis of distribution and association structure on the Sawtooth oak (Quercus acutissima) forest in Korea. Ph. D. Thesis, Changwon National University, Changwon, Korea. (in Korean).
  64. Swain, T.(1975) Evolution of flavonoid compounds. In: J.B. Harborne, T.J. Mabry and H. Mabry(eds.), The Flavonoids Part 2, Academic Press, New York, pp. 1096-1129.
  65. Trelease, W.(1917) Naming American hybrid oaks. Proc. Amer. Philos. Soc. 56: 44-52.
  66. Uyeki, H.(1932) Cupuliferae novae Koreanae. Acta Phytotax. Geobot. 1: 253-257.
  67. Whittemore, A.T. and B.A. Schaal(1991) Interspecific gene flow in sympatric oaks. Proc. Natl. Acad. Sci. USA 88: 2540-2544. https://doi.org/10.1073/pnas.88.6.2540
  68. Williams, C.A. and J.B. Harborne(1994) Flavone and flavonol glycosides. In: J.B. Harborne(ed.), The Flavonoids. Advances in Research since 1986, Chapman & Hall, London, pp. 337-385.
  69. Wyatt, R. and D.M. Hunt(1991) Hybridization in north American Asclepias. II. Flavonoid evidence. Syst. Bot. 16: 132-142. https://doi.org/10.2307/2418978