2008-Chromosome numbers of five species of the Marattiaceae in Taiwan

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2008-Chromosome numbers of five species of the Marattiaceae in Taiwan
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  335Taiwan J For Sci23(4):335-9,2008 Research note Chromosome Numbers of Five Species of the Marattiaceae in Taiwan Tsung-Hsin Hsieh, 1)  Yi-Han Chang, 2)  Wen-Liang Chiou, 2)  Yao-Moan Huang 2,3) 【 Summary 】 The chromosome numbers of root tips of all 5 species of the Marattiaceae in Taiwan were determined. Results showed that  Angiopteris lygodiifolia  (2 n  = 80),  Angiopteris palmiformis  (2 n  = 80), and  Marattia pellucida  (2 n  = 78) are diploid;  Archangiopteris itoi is triploid (2 n  = 120), and  Archangiopteris somai  is tetraploid (2 n  = 160). These chromosome numbers together with previ-ous evidence support the hypothesis that  Archangiopteris   itoi  is a hybrid between  Angiopteris ly- godiifolia and  Archangiopteris somai . Key words: Chromosome number, ploidy, Marattiaceae, Taiwan. Hsieh TH,   Chang   YH, Chiou WL, Huang YM. 2008.  Chromosome numbers of ve species of the Marattiaceae in Taiwan. Taiwan J For Sci 23(4):335-9. 研究簡報 台灣產五種觀音座蓮舅科植物之染色體數目 謝宗欣 1) 張藝翰 2) 邱文良 2) 黃曜謀 2,3) 摘 要 本文觀察台灣產 5 種觀音座蓮舅科植物之染色體數目。觀音座蓮 (2 n  = 80) 、蘭嶼觀音座蓮 (2 n  = 80) 及觀音座蓮舅 (2 n  =78) 為二倍體,伊藤氏原始觀音座蓮為三倍體 (2 n  = 120) ,台灣原始觀音座蓮為四倍體 (2 n  = 160) 。這些染色體數目與已往之研究證據均支持伊藤氏原始觀音座蓮為觀音座蓮與台灣原始觀音座蓮之雜交後代的假說。 關鍵詞 :染色體數、倍體數、觀音座蓮舅科、台灣。 謝宗欣、張藝翰、邱文良、黃曜謀 。 2008 。台灣產五種觀音座蓮舅科植物之染色體數目。台灣林業科學 23(4):335-9 。 1) Department of Biological Sciences and Technology, National University of Tainan, 33 Su-Lin St., Sec. 2, Tainan 70005, Taiwan.  國立台南大學生物科技學系, 70005 台南市樹林街二段 33 號。 2) Division of Forest Biology, Taiwan Forestry Research Institute, 53 Nanhai Rd., Taipei 10066, Taiwan.  林業試驗所森林生物組, 10066 臺北市南海路 53 號。 3) Corresponding author,e-mail:huangym@tfri.gov.tw  通訊作者。 Received July 2008, Accepted September 2008.2008 年 7 月送審 2008 年 9 月通過。    336Hsieh et al. ― Chromosome numbers of Marattiaceae There are 5 species of the family Marat-tiaceae native to Taiwan:  Angiopteris lygo-diifolia  Rosenst.,  Angiopteris palmiformis (Cav.) C. Chr.,  Archangiopteris itoi  Shieh,  Archangiopteris somai  Hayata, and  Marat-tia pellucida  Presl (DeVol and Shieh 1994). In Taiwan,  Angiopteris   lygodiifolia is a very common fern.  Angiopteris palmiformis is common on Lanyu (Orchid Island).  Marattia  pellucida  is also only distributed on Lanyu  but is rare.  Archangiopteris itoi  and  Arch-angiopteris somai are endemic species and sympatrically distributed in Wulai, northern Taiwan (DeVol 1975, DeVol and Shieh 1994).Two basic chromosome numbers of Marattiaceae (x = 40, 39) have been docu-mented. The basic chromosome number is 39 for  Marattia  and 40 for other genera (Braith-waite 1977, Walker 1985, Manickam and Irudayaraj 1988, Nakato 1988, Singh and Roy 1988, Weng and Qiu 1988, Li 1989, Camus 1990, Takamiya 1995). Of these,  Angiopteris lygodiifolia  was previously reported to be n  = 40 (Tsai and Shieh 1983), and  Archangiop-teris somai was reported to be 2 n  = 4x = 160 in Taiwan (Chou et al. 2007). Chromosome numbers of the root tips of all 5 species of the Marattiaceae in Taiwan were determined herein. Sixteen plants of 5 species were collect-ed from Taiwan and a nearby island, Lanyu. Those plants were transplanted to the Taipei Botanical Garden (Table 1). After about 1 month of cultivation, the root tips were pre-treated for 3~4 h with a mixture of 70 ppm cycloheximide and 250 ppm 8-hydroxy-quinoline (1:1) at about 18~20 ℃ . The root tips were xed for 1~3 h in a mixture of 45 %  acetic acid and absolute ethanol (1:3) at about 20 ℃ , and then preserved in 70 %  ethanol at 4 ℃ . Then they were macerated for 1~3 min in 1 N HCl at 60 ℃ , washed for 10 s and di-gested for 1~2 h in 4 %  pectinase. Finally, the root tips were squashed in modied Sharma’s solution (Sharma 1982). The chromosome number of root tip cells was observed with a light microscope (Leitz, Dialux 20) and  photographed with a digital camera (Nikon, Coolpix 995). The basic chromosome number of more than 22 species of  Angiopteris  has been documented to be 40 (Lin et al. 1996). Table 2 and Figures 1~5 show that  Angiopteris ly- godiifolia and  Angiopteris palmiformis  are diploid (2 n  = 2x = 80).  Archangiopteris   itoi is triploid (2 n  = 3x = 120), assuming that the  basic number is 40.  Archangiopteris somai  is tetraploid (2 n = 4x = 160), as previous Table 1. Materials of marattoid species in this study Species Locality Sample size Voucher  1)  Angiopteris lygodiifolia  Wulai, Taipei Co. 2 CM Chen s.n.  Toucheng, Ilan Co. 1 MH Huang 387  Angiopteris palmiformis  Lanyu, Taitung Co. 3 CW Chen s.n. (2 plants) SW Chung s.n.  Archangiopteris itoi  Wulai, Taipei Co. 3 YM Huang 1314 ,  1315 ,  1316  Archangiopteris somai  Wulai, Taipei Co. 2 YM Huang 1317  ,  1318  Lienhuachih, Nantou Co. 3 YM Huang 1319, 1320, 1321  Marattia pellucida  Lanyu, Taitung Co. 2 CW Chun s.n.  PF Lu s.n. 1) Living plants cultured at Taipei Botanical Garden.  337Taiwan J For Sci23(4):335-9,2008 Figs. 1~5. Chromosomes of the root tips of members of the Marattiaceae   in Taiwan. 1.  Angiopteris lygodiifolia (2 n  = 80); 2.  Angiopteris palmiformis (2 n  = 80); 3.  Marattia pellucida (2 n  = 78); 4.  Archangiopteris itoi   (2 n  = 120); 5.  Archangiopteris somai   (2 n  = 160). Scale bars = 10 µm.Table 2. Chromosome numbers of Marattiaceae species in Taiwan Chromosome number Species Present count Previous count Ploidy level Reference  2n n 2n  Angiopteris lygodiifolia 80 40 2x Tsai and Shieh (1983) 80 2x Nakato (1988)  Angiopteris palmiformis 80 2x  Archangiopteris   itoi  120 3x  Archangiopteris somai 160 160 4x Chou et al. (2007)  Marattia pellucida 78 2x  338Hsieh et al. ― Chromosome numbers of Marattiaceae determination (Chou et al. 2007).  Marattia  pellucida  is diploid (2 n  = 2x = 78), based on the basic chromosome number of 39, as pre-viously documented in  Marattia fraxinea  and  Marattia smithii  (Manickam and Irudayaraj 1988, Takamiya 1995).The morphology of  Archangiopteris itoi is intermediate between  Angiopteris ly- godiifolia and  Archangiopteris somai. These 3 species are sympatrically distributed in Wulai . Archangiopteris itoi  shares common  patterns of RAPD markers with  Angiopteris lygodiifolia and  Archangiopteris somai  and was hypothesized to be a hybrid of the lat-ter 2 species (Wu 2002). The chromosome numbers shown in this study also support this hypothesis. The family Marattiaceae has previously  been considered to have 6 genera,  Archangi-opteris ,  Angiopteris ,  Christensenia ,  Danaea ,  Marattia , and  Macroglossum  (Holttum 1954, Pichi Sermolli 1977). However, the Marattia-ceae was recently treated as having 4 genera:  Danaea , Christensenia ,  Marattia , and  Angi-opteris  (Camus 1990, Smith et al. 2006, Mur-dock 2008). The genera  Archangiopteris and  Macroglossum  were nested within the  Angi-opteris clade based on molecular phylogeny.  Angiopteris  is monophyletic only   if  Archangi-opteris  and  Macroglossum  are included (Mur-dock 2008). The same basic chromosome numbers of  Angiopteris and  Archangiopteris , and the possibility of hybridization between species of these 2 genera indicate no signi -cant boundary between these 2 genera. ACKNOWLEDGEMENTS The authors wish to thank Mei-Hsueh Huang, Chien-Wen Chen, Pi-Fong Lu, and Shih-Wen Chung for collecting live materials. LITERATURE CITED Braithwaite AF. 1977.  A chromosome count and range extension for Christensenia (Marat-tiaceae). Am Fern J 67:49-50. Camus JM. 1990.  Marattiaceae. In: Kubitzki K, editor. The families and genera of vascular  plants. Volume I. Berlin, Germany: Springer-Verlag. p 174-80. Chou HM, Huang YM, Wong SL, Hsieh TH, Hsu SY, Chiou WL. 2007.  Observations on gametophytes and juvenile sporophytes of  Archangiopteris somai  Hayata (Marattia-ceae), an endangered fern in Taiwan. Bot Stud 48:205-13. DeVol CE. 1975.  Marattiaceae. In: Li HL, Liu TS, Huang TC, Koyama T, DeVol CE, editors. Flora of Taiwan. Volume I. Taipei, Taiwan: Ep-och Publishing Co. p 71-8. DeVol CE, Shieh WC. 1994. Marattiaceae. In: Editorial Committee of The Flora of Taiwan, editors. Flora of Taiwan. Volume I. Taipei, Taiwan: Editorial Committee of The Flora of Taiwan. p 74-9. Holttum RE.   1954. A revised ora of Malaya. Vol. 2, Ferns of Malaya. Singapore: Govt Print Off. 643 p. Li JW. 1989.  Chromosome numbers of some species in the genus  Angiopteris  Hoffm. from Yunnan, China. In: Shing KH, Kramer KU, editors. Proc Int Symp Systematic Pteridology. Beijing, China: Science and Technology Press.  p 109-10. Lin SJ, Iwatsuki K, Kato M. 1996.  Cytotaxo-nomic study of ferns from China I. Species of Yunnan. J Jpn Bot 71:214-22. Manickam VS, Irudayaraj V. 1988.  Cytology of ferns of the western Ghats (South India).  New Delhi, India: Today and Tomorrow Print-ers and Publishers. 81 p. Murdock A. 2008.  Phylogeny of marattoid ferns (Marattiaceae): inferring a root in the ab-sence of a closely related outgroup. Am J Bot 95:626-41. Nakato N. 1988.  Notes on chromosomes of  339Taiwan J For Sci23(4):335-9,2008 Japanese pteridophytes (2). J Jpn Bot 63:214-8. Pichi Sermolli REG. 1977. Tentamen pteri-dophytorum genera in taxonomicum ordinem redigendi. Webbia   31:313-512. Sharma HC. 1982.  A technique for somatic counts from root tips of cereal seedlings raised  by embryo culture. Curr Sci 51:143-4. Singh VP, Roy SK. 1988.  Cytology of forty four species from Sikkim, Himalaya. Indian Fern J 5:162-9. Smith AR, Rryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG. 2006. A classica -tion for extant ferns. Taxon 55:705-31. Takamiya M. 1995. Chromosomal studies of ferns and fern-allies in the Republics of Fiji and Vanuatu, South Pacific. I. Psilotaceae, Ophioglossaceae, Marattiaceae and Schizaea-ceae. Acta Phytotax Geobot 46:137-45. Tsai JL, Shieh WC. 1983.  A cytotaxonomic survey of the Pteridophytes in Taiwan. J Sci Engin 20:137-58. Walker TG. 1985.  Cytotaxonomic studies of the ferns of Trinidad 2. The cytology and taxo-nomic implications. Bull Br Mus (Nat Hist) Bot 13:149-249. Weng RF, Qiu SP. 1988.  Chromosome counts of some ferns from Zhejiang. Invest Stud Nat 8:43-52. Wu WH.   2002. A study on the natural fern hybrid –  Archangiopteris itoi Shieh [Master thesis]. Taipei, Taiwan: Department of Botany,  National Taiwan University.
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