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Embryogeny of Sarcandra glabra (Thunb.) Nakai (Chloranthaceae) and Phylogenetic Position of the Chloranthaceae PDF

4 Pages·1998·0.31 MB·English
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植物研究雑誌 J. Jpn. Bot. 73: 22-25 (1998) Embryogeny of Sarcandra glabra (Thunb.) Nakai (Chloranthaceae) and Phylogenetic Position of the Chloranthaceae Takasi YA MAZAKI akanoku,T okyo,1 65-"J APAN (Recei ved on September 1,1 997) The development of embryo of Sarcandra glabra was described. The formation of embryo had the T-shaped proembryo at the beginning and the initial cells of the root- cortex were derived from the lowr tier of the cell m,so the embryonic development was conformed to the Polygonad type. It seems that the Chloranthaceae has closer affinity to the Polygonales and the Caryophyllales than the Magnoliales. Engler and Plantl (1894) treated Piperales feature with bracts (the so-called perianths in (including Chloranthales) as one of the most Hedyosmum,Y amazaki 1992),s tamens and primitive order in the Dicotyledons. Rendle carpels are arranged in opposite order as in (1937) considered that the Piperales (includ- those of Amentiferae. Therefore,th e Piperales ing Chloranthaceae) were allied to the and the Chrolanthales have less phylogenetic Polygonales. Recently,m any authors consid- affinity with the Lactoridaceae or the ered that the Piperales and the Chloranthales Trimeniaceae. were derived from Magnoliales (Swamy 1953, From the study of rbcL DNA sequences, Takhtajian 1954,M elchior 1964,C arlquist Chase et al. (1993) treated with Piperales and 1964,Cr onquist 1981 andEndress 1987). Many the Chrolanthales in the Paleoherbs which of them have considered that,w ithin the were those of the primitive Angiosperms. Magnoliidae,t he floral morphology of Method of embryonic development reveal Lactridaceae or Trimeniaceae showed some an important data in the study of the similar construction with the Piperales and the Angiosperms (Yamazaki 1982). Embryonic Chloranthales. They were regarded as ag roup development in the Chloranthacea was stud- with reduced forms of the Magnoliales. ied in Chloranthus serratus Roem. et Schult. The flower of the Lactridaceae consists of (Yo shida 1959a) and Sarcandra irvingbaileyi three perianths,si x stamens in two rows and Swamy (Vijayaraghavan 1964),bu t these re- three carpels. These floral elements were ar- ports were insufficient to discuss the ranged with the alternate order and were fun- phylogenetic affinity based on embryonic de- damentall ya rranged in the reduced form of the velopmen .tThe present work of embryonic Magnoliaceous flower (Carlquist 1964). In the development on Sarcandra glabrαis an at- Trimeniaceae stamens are spiral1yi nserted on tempt to contribute to the phylogenetic rela- receptacle independent of perianths. On the tionships in the Chloranthales. other hand,th e Piperales and the Chrolanthales have various floral construction. Nevertheless Materials and Methods the flowers have the fundamentally common Flowers and fruits were collected from cul- -22ー February 1998 Journal of Japanese Botany Vo .l73 No. 1 23 tivated plants in Botanical Gardens,Fa culty of Science,Un iversity ofTokyo,in July to Octo- ber,1 994 to 1997. The materials were fixed in an alcohol-formalin -acetic acid,an d imbedded in paraffin in the usual way. Serial sections, 12-14μm thick,w ere cut,a nd stained by Heidenhain' sh ematoxylin and fast green. Results Ovule and embryo sac. Ovule is ortho- tro-pous,b itegmic,a nd crassinucellar,w ith the micropyle formed by both integuments (Fig. 1-1). The megaspore mother cells lie in al inear tetrad,a nd the chalazal one develops into an embryo sac. These embryosac forma- tions have been reported by Yo shida (1959b), and will not be described in the present paper. Embryogenesis. The first division of the fertilized egg is transverse resulting in the formation of the terminal cell ca and the basal cell cb (Fig. 2: 1). The cell ca divides verti- cally,wh ile cb undergoes at ransverse division producing cells m and ci. The four-celled proembryo is in T-shaped (Figs. 2: 2,3) . The terminal cells ca divide vertically to form four cells. Each of four cells undergoes repeatedly transverse and vertical divisions to form the vegetative apex,tw o cotyledons and the upper part of hypocotyl (Figs. 2: 6-12). In earlier stages the cell m undergoes vertical divisions repeatedly (Figs. 2: 4-12) and in many celled proembryo stages each of these cells under- Fig. 1. 1,tr ansverse section of ovule. e,fe rtilied goes transverse divisions to form two tiers egg. 2a nd 3,yo ung stages of embryo. Abbrevia- tions in text. Bar 1= 0. 2 mm. 2a nd 3= 5 0μm. (Fig. 2: 13). The upper titer of m repeatedly divides transversely and vertically to form the lower part of hypocotyl. The lower tier of m 。て produces the initial of root-cortex (iec) in the ca 山 川2hyp center (Figs. 2: 14,1 5). The lower cell ci divides transversely to form na nd s(F igs. 1: 2, C71/2hyp+iec 2: 4). )'he cell ns lowly divides transversely and vertically to form the [oot cap (Figs. 2: 14, \ 15). The cell ci divides to form the suspensor (Figs. 2: 13,14 ). s-一一-susp The recapitulatory table is as follows: 24 植物研究雑誌第73巻第1号 平成10年2月 与:;官専 , j、1似合Ir.ii_m_ \ψbl門 l トーγ- IJ 一円、 「t同Lト1ヱGLー?b今÷JθーθよィLィ斗--一ce問En-a一、(cid:631)一(cid:631)(cid:631)一一トlヤ@(@?4@町r9rfe1-H@ θ:;j 1 2 3 4 5 6 -ca ----寸ー『、 ----四--.-..._ 7 8 9 14 50~m 13 15 Fig. 2. Stages of development of embryo. Abbreviations in text. Abbreviations. pvt,p rovegetative apex. pco, zaki 1982). The embryonic deve10pment of procotyledon. phy,p rohypocoty .1iec,in itial cells Houttuynia cordata Thunb. in Saururaceae is of root-cortex. co,ro ot-cap. susp,s uspensor. the same as that of Sarcandra spp (Crete 1957). Crete stated that the embryonic devel- Discussion opment of Houttuynia cordαta resemb1es that The development of embryo of Sarcandra of the Polygonaceae and the Nyctangiacea. glabra is fundamentally the same as that of The Ch10ranthales and the Piperales seem to S. irvingbailleyi reported by Vijaraghavan have closer affinity to the Polygonales and the (1964). He determined that the embryonic Caryophy llales than the恥1agnoliales,in development of S. irvingbailleyi confirmed which the embryonic development is the to the Onagrad type by the T- shaped pro- Geraniad type or its variation. embryo. However,i n the Onagrad type the initial cells of root-cortex directly derives The author is grateful to Prof.孔1itsuko from one cell originated from the upper part of Sugiyama,Sa itama Women's Junior College, tier m. In Sarcandra spp. the initial cells of for her help to this study. root-cortex were derived from af ew cells References originated from the lower part of the tier m, Car1quist S. 1964. Morpho1ogy and re1ationships of and the embryonic development of Sarcandra Lactoridaceae. A1iso 4: 421-435. spp. is conformed with the Polygonad type as Cronquist A. 1981. An integrated system of c1assification of tlowering p1ants. 80-89. Co1umbia Univ. PressNew York. previously described by the author (Yama- , February 1998 Journal of Japanese Botany Vo .173 No. 1 25 Chase M. W. et a .11993. Phylogenetics of seed plants: an 一一一一一1996.Piperales,C hloranthales. 1η: Takh司janA. L. analysis of nucleotide sequences from the plastid gene rbcL. (ed.),Di versity and classification of flowering plants. pp. Ann. Miss. Bot. Gard. 80: 528-580. 50-53,63-65 Crete M. P. 1957. Embryogenie Vegetale. Saururaceae. Devel- Vijayaraghavan M. R. 1964. Morphology and embryology of a opment de l'embryon chez l'Houttuynia cordata Thunb. vesselless Dicotyledon. Sarcandra irvingbaileyi Swamy Compt. Rend. Acad. Sci. Paris 245: 1331-1334. and systematic position of the Chloranthaceae. Endres P. K. 1987. The Chloranthaceae: Reproductive structure Phytomorphology 14: 429-441. and phylogenetic position. Bo .tJahrb. Sys .t109: 153-226. Yamazaki T. 1982. Recognized types in early development of Engler A. und Prantl K. 1894. Chloranthaceae. In: Engler A. und the embryo and the phylogenetic significance in the Dicoty- Prantl K. (eds.),N a turlich Pflanzenfamilien 111. 1: pp. 12- ledon. Act. Phytotax. Geobot. 33: 400-409. 14. Wilhelm Engelmann,L eipzig. 一一一一一1992.Floral morphology of Hedyosmum orientale Melchior H. 1964. Chloranthaceae. In: Melchior H. (edよ Merr. et Chun (Chloranthaceae) and phylogenetic signifi- Engler's Syllabus der Pflanzenfamilien. Auf. 12. Band 11: cance of its perianth. J. Jpn. Bot. 67: 257-269. pp. 147-151. Borntnaegar,Be rlin. Yoshida O. 1959a. Embryologische Studien uber die Ordung Rendle A. B. 1938. Chloranthaceae. In: (ed.),Th e classification Piperales 11. Embryologie von Chloranthus serratus Roem. of flowering plants 11: 88-95. et Schult. 1. Coll. Art. Sci. Chiba Univ. 2: 295-303. Swamy B. G. L. 1953. The morphology and relationships ofthe 一一一一1959b.Embryologische Studien uber die Ordung Chloranthaceae. J. Arn. Arb. 34: 375-408. Piperales 111. Embryologie von Sarcandra glabra Nakai. 1. Takhtaian A. L. 1954. Origins of Angiospermous plants. Trans- Coll. Ar .tSci. Chiba Univ. 3: 55-60. lated and published by Amer. Inst. Bio .1Sc.,p. 45 (1957). 山崎敬:センリョウ(センリョウ科)の匪発生と センリョウ科の系統学上の位置 センリョウ科はドクダミ科と共に双子葉植物の タデ目やナデシコ目と関係があるものと考えられ 原始的な群として扱われたり モクレン目群から る.近縁とされるLactoridaceaeやTrimeniaceaeでは の特殊化した群として扱われたりしている.最近 脹形成の詳細な研究はされていないが,花の構造 は主に後者の見解が支持されている.匹形成は科 はセンリョウ科とは基本的に異なるから,モクレ や目によってほぼ一定していて,系統関係の研究 ン目との関連は考えられない.DNAからの結果と には重要な手掛かりとなる.今回の観察によりセ は,センリョウ目が原始的であることは一致する ンリョウの脹形成はPolygonadtypeで, ドクダミ科 が,他の目や科との類縁関係は一致しない.この のドクダミと同じであることが明らかとなった. 点については今後の研究が必要である. このことからするとセンリョウ科やドクダミ科は (東京都中野区・・・・・・・)

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