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소구상체(Orbicules)의 계통분류학적 검토

The phylogenetic potential of orbicules in angiosperms

  • 문혜경 (경희대학교이과대학 생물학과)
  • 투고 : 2017.11.21
  • 심사 : 2018.03.21
  • 발행 : 2018.03.30

초록

Orbicule(소구상체)의 분포를 확인하기 위하여 꿀풀과에 속하는 6속 11분류군과 마편초과 3속 4분류군의 약벽을 주사전자현미경(scanning electron microscope)을 이용하여 관찰하였으며, 문헌조사를 통해 현화식물 내 소구상체의 분포가 갖는 계통분류학적 유용성 및 융단조직 유형과의 관련성에 대해 검토하였다. 그 결과 꿀풀과 분류군들에서는 모두 소구상체가 관찰되지 않았고, 마편초과 분류군들에서는 평균 $1{\mu}m$ 이하, 구형의 소구상체들이 조밀하게 분포하는 것을 살펴볼 수 있었다. 이와 같은 과 수준에서 일관성 있는 소구상체의 분포 특징은 현화식물 전체 orbicules에 대해 조사된 150과 중 123과와 일치하는 것으로 소구상체의 분포 유형 자체가 분류학적 예견적 가치를 지니는 것으로 나타났다. 또한, 융단조직 유형이 확인된 분류군들 중 분비형 융단조직을 갖는 분류군의 약 84%에서 소구상체가 나타나고, 아메바형 융단조직을 갖는 분류군의 약 80%은 소구상체가 나타나지 않는 것으로 조사되어 소구상체의 발달이 융단조직 유형과 밀접한 관련이 있는 것으로 조사되었다. 하지만, 현재까지 수행된 소구상체 연구는 현화식물 전체 416과들 중 150과에 제한되어 있고, 이들 분류군들 중 융단조직 유형은 92과에서만 알려져 있어 소구상체의 기능 및 융단조직 유형과의 유연관계를 규명하기 위해서는 보다 광범위한 분류군을 대상으로 한 연구가 필요한 것으로 나타났다.

The distribution of orbicules was investigated for eleven taxa of six genera in Lamiaceae and four taxa of three genera in Verbenaceae using scanning electron microscopy. A literature survey to evaluate the phylogenetic potential of the orbicules and their possible correlations with tapetum types was also conducted. The orbicules are consistently absent in all investigated taxa of Lamiaceae, while small orbicules of an average size of less than $1{\mu}m$ are densely distributed in Verbenaceae. In fact, orbicules appear consistently in 123 of 150 angiosperm families when investigated in at least one species. Thus, the distribution patterns of orbicules could be a useful diagnostic character in angiosperms. In addition, orbicules occur in 84% taxa of the secretory tapetum type, while they are commonly absent in the amoeboid tapetum type (ca. 80%). The presence of orbicules may be correlated with the secretory tapetum type. However, the study of orbicules is restricted in 150 families and the tapetum type within these families can be applied for 92 families out of a total of 416 angiosperm families. Thus, further investigation of orbicules is necessary in extended taxa to address the questions pertaining to orbicules.

키워드

참고문헌

  1. Angiosperm Phylogeny Group. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20. https://doi.org/10.1111/boj.12385
  2. Chung, J.-D. and S.-R. Kuo. 2005. Reproductive cycles of Calocedrus formosana. Taiwan Journal of Forest Science 20: 315-329.
  3. Cole, T. C. H., H. H. Hilger and P. F. Stevens. 2017. Angiosperm phylogeny poster: flowering plant systematics. Retrieved October 1, 2017, available from http://www2.biologie.fu-berlin.de/sysbot/poster/poster1.pdf.
  4. Doria, M. G., N. Pabon-Mora and F. Gonzalez. 2012. Reassessing inflorescence and floral morphology and development in Hedyosmum (Chloranthaceae). International Journal of Plant Sciences 173: 735-750. https://doi.org/10.1086/666662
  5. El-Ghazaly, G. 1999. Tapetum and orbicules (Ubisch bodies): development, morphology and role of pollen grains and tapetal orbicules in allergenicity. In Fertilization in Higher Plants. Cresti, M., G. Cai and A. Moscatelli (eds.), Springer, Berlin. Pp. 157-173.
  6. El-Ghazaly, G. and S. Nilsson. 1991. Development of tapetum and orbicules of Catharanthus roseus (Apocynaceae). In Pollen Spores: Patterns of Diversity. Balckmore, S. and S. H. Barnes (eds.), Clarendon Press, Oxford. Pp. 317-329.
  7. Erdtman, G. 1960. The acetolysis method, a revised description. Svensk Botanisk Tidskrift 54: 561-564.
  8. Fernando, D. D. and D. D. Cass. 1994. Plasmodial tapetum and pollen wall development in Butomus umbellatus (Butomaceae). American Journal of Botany 81: 1592-1600.
  9. Furness, C. A. 2008. A review of the distribution of plasmodial and invasive tapeta in eudicots. International Journal of Plant Sciences 169: 207-223.
  10. Furness, C. A. 2011. Comparative structure and development of pollen and tapetum in Malpighiales, with a focus on the parietal clade. International Journal of Plant Sciences 172: 980-1011.
  11. Furness, C. A. 2014. Development of the remarkable pollen apertures of the jellyfish tree Medusagyne oppositifolia (Ochnaceae, Malpighiales). International Journal of Plant Sciences 175: 803-813. https://doi.org/10.1086/676984
  12. Furness, C. A. and P. J. Rudall. 2001. The tapetum in basal angiosperms: early diversity. International Journal of Plant Sciences 162: 375-392. https://doi.org/10.1086/319580
  13. Galati, B. G. 2003. Ubisch bodies in angiosperms. In Advances in Plant Reproductive Biology. Chauhann, M. R. and A. K. Pandey (eds.), Narendra Publishing House, Delhi. Pp. 1-20.
  14. Galati, B. G., F. Monacci, M. M. Gotelli and S. Rosenfeldt. 2007. Pollen, tapetum and orbicule development in Modiolastrum malvifolium (Malvaceae). Annals of Botany 99: 755-763.
  15. Garcia, M. T. A., B. G. Galati and A. M. Anton. 2002. Microsporogenesis, microgametogenesis and pollen morphology of Passiflora spp. (Passifloraceae). Botanical Journal of the Linnean Society 139: 383-394. https://doi.org/10.1046/j.1095-8339.2002.00072.x
  16. Gotelli, M. M., B. G. Galati and G. Zarlavsky. 2016. Pollen development and anther morphology in 14 species of Rhamnaceae. Plant Systematics and Evolution 302: 1433-1444. https://doi.org/10.1007/s00606-016-1342-2
  17. Hesse, M. 1986. Orbicules and the ektexine are homologous sporopollenin concretions in Spermatophyta. Plant Systematics and Evolution 153: 37-48. https://doi.org/10.1007/BF00989416
  18. Huysmans, S., G. El-Ghazaly and E. Smets. 1998. Orbicules in angiosperms: morphology, function, distribution, and relation with tapetum types. The Botanical Review 64: 240-272. https://doi.org/10.1007/BF02856566
  19. Huysmans, S., G. El-Ghazaly and E. Smets. 2000. Orbicules: still a well-hidden secret of the anther. In Plant Systematics for the 21st Century: Proceedings from a Symposium held at the Wenner-Gren Centre. Nordenstam, B., G. El-Ghazaly and M. Kassas (eds.), Portland Press, London. Pp. 201-212
  20. Huysmans, S., B. Verstraete, E. Smets and L. W. Chatrou. 2010. Distribution of orbicules in Annonaceae mirrors evolutionary trend in angiosperms. Plant Ecology and Evolution 143: 199-211. https://doi.org/10.5091/plecevo.2010.438
  21. Jeong, B.-K. 2009. Ultrastructural study of programmed cell death of tapetum in Panax ginseng. Journal of Life Science 19: 1016-1022. https://doi.org/10.5352/JLS.2009.19.8.1016
  22. Kosmath, L. 1927. Studien Uber Das Antherentapetum. Osterreichische Botanische Zeitschrift 76: 235-241. https://doi.org/10.1007/BF01246254
  23. Lattar, E. C., B. G. Galati and M. S. Ferrucci. 2012. Ultrastructural study of pollen and anther development in Luehea divaricata (Malvaceae, Grewioideae) and its systematic implications: role of tapetal transfer cells, orbicules and male germ unit. Flora: Morphology, Distribution, Functional Ecology of Plants 207: 888-894. https://doi.org/10.1016/j.flora.2012.10.005
  24. Lattar, E., B. G. Galati and M. S. Ferrucci. 2014. Comparative study of anther development, microsporogenesis and microgametogenesis in species of Corchorus, Heliocarpus, Luehea and Triumfetta (Malvaceae: Grewioideae) from South America. New Zealand Journal of Botany 52: 429-445. https://doi.org/10.1080/0028825X.2014.961490
  25. Lombardo, G. and L. Carraro. 1976. Tapetal ultrastructural changes during pollen development. III. Studies on Gentiana acaulis. Caryologia 29: 345-349. https://doi.org/10.1080/00087114.1976.10796674
  26. Lopez, A. and S. Rosenfeldt. 2015. Oxalis sect. Palmatifoliae (Oxalidaceae): pollen grains morphology and orbicules diversity. Boletin de la Sociedad Argentina de Botanica 50: 349-352.
  27. Lopez, A. and S. Rosenfeldt. 2016. Oxalis section Alpinae (Oxalidaceae): orbicule diversity and pollen grain morphology. Turkish Journal of Botany 40: 637-644. https://doi.org/10.3906/bot-1603-23
  28. Lovisolo, M. R. and B. G. Galati. 2012. Orbicules diversity in Poaceae. Boletin de la Sociedad Argentina de Botanica 47: 87-96.
  29. Marx, H. E., N. O’Leary, Y.-W. Yuan, P. Lu-Irving, D. C. Tank, M. E. Mulgura and R. G. Olmstead. 2010. A molecular phylogeny and classification of Verbenaceae. American Journal of Botany 97: 1647-1663. https://doi.org/10.3732/ajb.1000144
  30. Merckx, V., P. Schols, K. Geuten, S. Huysmans and E. Smets. 2008. Phylogenetic relationships in Nartheciaceae (Dioscoreales), with focus on pollen and orbicule morphology. Belgian Journal of Botany 141: 64-77.
  31. Moon, H.-K. 2008. Systematic studies in Mentheae (Lamiaceae): morphology, evolutionary aspects, phylogeny. PhD dissertation, Katholieke Universiteti Leuven, Leuven, Belgium, 196 pp.
  32. Moon, H.-K., S. Vinckier, E. Smets and S. Huysmans. 2008a. Comparative pollen morphology and ultrastructure of Mentheae subtribe Nepetinae (Lamiaceae). Review of Palaeobotany and Palynology 149: 174-186.
  33. Moon, H.-K., S. Vinckier, E. Smets and S. Huysmans. 2008b. Palynological evolutionary trends within the tribe Mentheae with special emphasis on subtribe Menthinae (Nepetoideae: Lamiaceae). Plant Systematics and Evolution 275: 93-108. https://doi.org/10.1007/s00606-008-0042-y
  34. Moon, H. K., S. Vinckier, J. B. Walker, E. Smets and S. Huysmans. 2008c. A search for phylogenetically informative pollen characters in the subtribe Salviinae (Mentheae: Lamiaceae). International Journal of Plant Sciences 169: 455-471. https://doi.org/10.1086/526463
  35. Nilsson, S. and A. Robyns. 1974. Pollen morphology and taxonomy of the genus Quararibea s. l. (Bombacaceae). Bulletin du Jardin Botanique National de Belgique/Bulletin van de Nationale Plantentuin van Belgie 44: 77-99. https://doi.org/10.2307/3667429
  36. Perez-Gutierrez, M. A., M. C. Fernandez, M. J. Salinas-Bonillo, V. N. Suarez-Santiago, S. Ben-Menni Schuler and A. T. Romero-Garcia. 2016. Comparative exine development from the posttetrad stage in the early-divergent lineages of Ranunculales: the genera Euptelea and Pteridophyllum. Journal of Plant Research 129: 1085-1096. https://doi.org/10.1007/s10265-016-0862-8
  37. Raj, B. and G. El-Ghazaly. 1987. Morphology and taxonomic application of orbicules (Ubisch bodies) in Chloanthaceae. Pollen Spores 29: 151-166.
  38. Rao, C. 1954. Embryological studies in Malvaceae I. Proceedings of the National Institute of Sciences in India Part B 20: 127-150.
  39. Rosanoff, S. 1865. Zur kenntniss des baues und der entwickelungsgeschichte des pollens der Mimosaceae. Jahrbuch fur Wissenschaftliche Botanik 4: 441-450.
  40. Rowley, J. R. 1962. Nonhomogeneous sporopollenin in microspores of Poa annua L. Grana 3: 5-19.
  41. Rowley, J. R. and N. I. Gabarayeva. 2004. Microspore development in Quercus robur (Fagaceae). Review of Palaeobotany and Palynology 132: 115-132. https://doi.org/10.1016/j.revpalbo.2004.05.003
  42. Rowley, J. R., K. Muhlethaler and A. Frey-Wyssling. 1959. A route for the transfer of materials through the pollen grain wall. The Journal of Biophysical and Biochemical Cytology 6: 537-538. https://doi.org/10.1083/jcb.6.3.537
  43. Santos, R. P. and J. F. A. Mariath. 1999. Ultrastructure of the orbicules (Ubisch bodies) in Ilex paraguariensis St.Hil. (Aquifoliaceae). Acta Microscopica 8(Supplement C): 773-774.
  44. Sajo, M. G., C. A. Furness, C. J. Prychid and P. J. Rudall. 2005. Microsporogenesis and anther development in Bromeliaceae. Grana 44: 65-74.
  45. Schols, P., C. A. Furness, P. Wilkin, S. Huysmans and E. Smets. 2001. Morphology of pollen and orbicules in some Dioscorea species and its systematic implications. Botanical Journal of the Linnean Society 136: 295-311. https://doi.org/10.1111/j.1095-8339.2001.tb00574.x
  46. Schols, P., C. A. Furness, P. Wilkin, E. Smets, V. Cielen and S. Huysmans. 2003. Pollen morphology of Dioscorea (Dioscoreaceae) and its relation to systematics. Botanical Journal of the Linnean Society 143: 375-390. https://doi.org/10.1111/j.1095-8339.2003.00227.x
  47. Song, J.-H., H.-K. Moon and S.-P. Hong. 2016. Pollen morphology of the tribe Sorbarieae (Rosaceae). Plant Systematics and Evolution 302: 853-869. https://doi.org/10.1007/s00606-016-1303-9
  48. Song, J.-H., H.-K. Moon, M.-K. Oak and S.-P. Hong. 2017a. Phylogenetic evaluation of pollen and orbicule morphology in Rosaceae tribe Neillieae (subfamily Amygdaloideae). Botanical Journal of the Linnean Society 183: 439-453. https://doi.org/10.1093/botlinnean/bow019
  49. Song, J.-H., M.-K. Oak, H.-S. Roh and S.-P. Hong. 2017b. Morphology of pollen and orbicules in the tribe Spiraeeae (Rosaceae) and its systematic implications. Grana 56: 351-367. https://doi.org/10.1080/00173134.2016.1274334
  50. Strittmatter, L. and B. Galati. 2000. Embryological study in Oziroe acaulis (Baker) Speta (Hyacinthaceae). Phytomorphology 50: 161-171.
  51. Tang, Y., H. Gao, C.-M. Wang and J.-Z. Chen. 2006. Microsporogenesis and microgametogenesis of Excentrodendron hsienmu (Malvaceae s.l.) and their systematic implications. Botanical Journal of the Linnean Society 150: 447-457. https://doi.org/10.1111/j.1095-8339.2006.00476.x
  52. Tang, Y., H. Gao and J.-S. Xie. 2009. An embryological study of Eriolaena candollei Wallich (Malvaceae) and its systematic implications. Flora-Morphology, Distribution, Functional Ecology of Plants 204: 569-580. https://doi.org/10.1016/j.flora.2008.09.002
  53. Taylor, M. L., B. L. Gutman, N. A. Melrose, A. M. Ingraham, J. A. Schwartz and J. M. Osborn. 2008. Pollen and anther ontogeny in Cabomba caroliniana (Cabombaceae, Nymphaeales). American Journal of Botany 95: 399-413. https://doi.org/10.3732/ajb.95.4.399
  54. Ubisch, G. 1927. Zur Entwicklungsgeschichte der Antheren. Planta 3: 490-495. https://doi.org/10.1007/BF01916485
  55. Vardar, F., I. Ismailo?lu and M. Unal. 2013. Anther development and cytochemistry in Asphodelus aestivus (Asphodelaceae). Turkish Journal of Botany 37: 306-315.
  56. Verellen, J., E. Smets and S. Huysmans. 2004. The remarkable genus Coptosapelta (Rubiaceae): pollen and orbicule morphology and systematic implications. Journal of Plant Research 117: 57-68. https://doi.org/10.1007/s10265-003-0128-0
  57. Verstraete, B., I. Groeninckx, E. Smets and S. Huysmans. 2011. Phylogenetic signal of orbicules at family level: Rubiaceae as case study. Taxon 60: 742-757.
  58. Verstraete, B., H.-K. Moon, E. Smets and S. Huysmans. 2014. Orbicules in flowering plants: a phylogenetic perspective on their form and function. The Botanical Review 80: 107-134. https://doi.org/10.1007/s12229-014-9135-1
  59. Vinckier, S. and E. Smets. 2003. Morphological and ultrastructural diversity of orbicules in Gentianaceae. Annals of Botany 92: 657-672. https://doi.org/10.1093/aob/mcg187
  60. Yamada, Y. 1987. Pollen grains in the anther locule by scanning electron microscopy. Japanese Journal of Palynology 33: 59-76.
  61. Zini, L. M., B. G. Galati, G. Zarlavsky and M. S. Ferrucci. 2017. Developmental and ultrastructural characters of the pollen grains and tapetum in species of Nymphaea subgenus Hydrocallis. Protoplasma 254: 1777-1790. https://doi.org/10.1007/s00709-016-1074-8

피인용 문헌

  1. Phylogenetic and morphologic survey of orbicules in angiosperms vol.69, pp.3, 2018, https://doi.org/10.1002/tax.12281
  2. A Comprehensive Study of the Genus Sanguisorba (Rosaceae) Based on the Floral Micromorphology, Palynology, and Plastome Analysis vol.12, pp.11, 2018, https://doi.org/10.3390/genes12111764