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Karyotype Analysis of Eight Species and One Variety of Carpesium (Compositae) in Japan PDF

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植物研究雑誌 J. Jpn. Bot. 66: 26-34 (1991) Karyotype Analysis of Eight Species and One Variety of Carpesium (Compositae) in Japan Young-Ho CHO Makino Herbarium,Fa culty of Science,T okyo Metropolitan University, 2・1・1,Fukazawa,S etagaya,T okyo,1 58 JAPANl) 日本産ヤアタノぐコ属8種 1変種の核型分析 曹英鏑 東京都立大学理学部牧野標本館 158東京都世田谷区深沢2-1-11) (Received on February 23,19 90) Chromosome number and karyotype of eight species and one variety of Japanese Gα'fpesium were studied. Chromosome number 2n = 40 of C. koidzumii is reported for the first time. Karyotypes of the species studied fall into two groups on the basis of the number of the largest chromosomes. The genus Carpesium,c onsisting of about 20 Materials and methods species of a perennial or biennial herb,i s All of the plants examined in this study were distributed in eastern Asia and Europe,es pecially collected from natural habitats. They were in China,Ko rea and Jap an. Eight species and one transplanted to the garden of Makino Herbarium variety have been known from Ja pan (Kitamura (MAK), T okyo Metropolitan University. The et al. 1981). Concerning the Japanese species, original source and number of individuals karyological investigations are still insufficient, examined are shown in Table 1. Cytological though Shimakawa (1952) and Arano (1962) have preparation was made by the following procedure. studied. Arano (1962) reported 2n=40 chromo- Root tips were pretreated with 0.002M 8-hydroxy- somes from six speci回 andone variety,an d 2n =3 6 quinoline solution for about 3 hr at room from C.αbrotanoides,a nd then,cl assified them temperature and fixed with 45% acetic acid for into five types based on karyological analysis. But about 30 min. After being macerated in a 1:2 Arano gave no description of karyological mixture of 450/0 acetic acid and 1N hydrochloric differences of these five types,pa rticularly,th e acid for 20 sec. at 60oC,ro ot tips w町estained with centromeric position and length of chromosomes. 2% acetic orcein solution for over one night. Therefore,1 r e-examined the chromosome number Subsequently,t hey were squashed with conven- and karyotype of eight species and one variety of tional method. The chromosome numbers and Jap anese Carpesium including C. koidzumii. karyotypes were observed in well-spread metaph邸e plates of somatic root tip cells. The arm ratio (length of the long arm/length of the short arm) 司ゐぷU February 1991 Journal of Japanese Botany Vol. 66 No. 1 27 Table 1. Chromosome number,lo cality and number of individuals examined. Species Chromosome Locality No. of ind. no. (2n) examined c.αbrotαnoides 40 Tokyo Metropolitan Univ. campus 7 , 40+2B Namezawa,T agata,S hizuoka Pref. 2 Nippara,N ishitama,T okyo 3 M1. Hinodesan,N ishitama,T okyo 2 M1. Takaosan,H achioji,T okyo c. rosulatum 40 M1. Sanagisan,N umazu,S hizuoka Pref. 2 Kannami,T agata,S hizuoka Pref. Mt. Takaosan,H achioji,T okyo 3 C. glossophyllum 40 Mt. Sanagisan,N umazu,S hizuoka Pref. Tazima,M inamiaizu,F ukushima Pref. 2 Chonan,C hosei,C hiba Pref. 2 Mt. Mitakesan,N ishitama,T okyo 2 Mt. Takaosan,H achioji,T okyo 4 C. macrocephalum 40 Tatsuno,K amiina,N agano Pref. Mt. Kumotorisan,N ishitama,T okyo 3 M1. Hinodesan,N ishitama,T okyo C. cernuum 40 M1. Misakasan,F ujiyoshida,Y amanashi 3 Pref. Um izawa,N ishitama,T okyo 4 C. triste 40 Nikko,T ochigi Pref. 3 Chitose,H okkaido 2 Nippara,N ishitama,T okyo 4 C. divaricαtum 40 Mt. Sanagisan,N umazu,S hizuoka Pref. 3 Mt. Mitakesan,N ishitama,T okyo 2 Mt. Hinodesan,N ishitama,T okyo 2 Mt. Takaosan,H achioji,T okyo 7 C. divaricatum 40 Minakami,T one,G umina Pref. var. matsuei 40+2B M1. Mitakesan,N ishitama,T okyo 2 40+3B Mt. Kumotorisan,N ishitama,T okyo M1 . Hinodesan,N ishitama,T okyo 4 M1. Takaosan,H achioji,T okyo C. koidzumii 40 Kadoma valley,C hiisagata,N agano Pref. M1. Misakasan,F ujiyoshida,Y amanashi 3 Pref. 28 植物研究雑誌第66巻第1号 平成3年2月 and the classification for the centromeric position 9& 10,2 5-30,3 3-36,3 9 & 40),2 2 submeta- on a chromosome used here are as follows; centric (nos. 1& 2,5 -8,1 1-24,37 & 38) and 4 1.0-1.7: metacentric,1 .7-3.0: submetacentric, subtelocentric chromosomes (nos. 3& 4 ,31 &3 2). 3.0-7.0: subtelocentric chromosome. The voucher C. glossophyllum Maxim. The chromosome specimens are deposited in Kyungpook National number was counted as 2n =4 0σig. lC). The University Herbarium. chromosomes of this species were 5.6-1.7μmin length and comparatively larger than those of the Results other taxa examined. They were somewhat sym- The chromosome numbers and karyotypes metrical. A pair of the largest submetacentric counted in this study are presented in Table 1a nd chromosomes (nos. 1& 2) w町eeasily distinguished Figs. 1& 2. The description of the karyotype of from the others in its karyotype σig. 2C). The each species is as follows: karyotype was composed of 16 metacentric (nos. C. abrotanoides L. The chromosome number 7& 8,15 & 16,21 -26,33 -38),16 submetacentric was counted as 2n =4 0 and 40 +2 B in somatic (nos. 1& 2,9 & 10,13 & 14,19 & 20,27 -32,39 cells (Fig. lA),u ncongruent with previous & 40) and 8s ubtelocentric chromosomes (nos. reports (Sh凶akawa1952,A rano 1962; 2n=36). 3-6,11 & 12,17 & 18). The chromosomes were 4.9-1.0μm in length and C. macro,c同phalumFr. et Sav. The chro- relatively smaller than those of the other species mosome number was counted as 2n =4 0 (Fig. examined and their shape was more or less lD). The chromosomes were 5.3-1.6μm in symmetrical. In its karyotype (Fig. 2A),a p air of length and relatively larger than those of the others the largest metacentric chromosome (nos. 1& 2) and also more or less symmetrical. In its karyotype were easily distinguished from the others. The σig. 2D),t wo pairs of the largest metacentric karyotype included 20 metacentric (nos. 1& chromosomes (nos. 1-4) were easily distinguished 2,9 -14,1 7-24,2 7-30),8 s ubmetacentric (nos. from the others. The karyotype was consisted of 3-6,15 & 16,25 & 26),2 s ubtelocentric (nos. 7 30 metacentric (nos. 1-18,2 1-28,3 5-38),8 & 8) and other 10 chromosomes. The ten chromo- submetacentric (nos. 19 & 20,2 9-32,3 9 & 40) somes were observed as meta-or submetacentric and 2s ubtelocentric chromosomes (nos. 33 & 34). (nos. 31-40),a lthough the centromeric position C. cernuum L. The chromosome number was not always visible. Two B-chromosomes were was counted as 2n =4 0 (Fig. lF). The chromo- observed in chromosome complements of the in- somes were 4.0-1.1μm long and more or less dividuals from Nippara,T okyo and the popula- symmetrical. Two pairs of the largest metacentric tions in the campus,T okyo Metropolitan chromosomes (nos. 1-4) w eeasily distinguished 町 University,Se tagaya,T okyo (Fig. lA). from the others in its karyotype σig. 2E). The C. rosulatum Miq. The chromosome number karyotype was composed of 28 metacentric (nos. was counted as 2n =4 0 (Fig. lB). Chromosome 1-16,23 -26,31 -38),8 s ubmetacentric (nos. 17 & length was 4.3-1.5μm and more or less sym- 18,2 7-30,3 9 & 40) and 4 subtelocentric metrical. In its karyotype σig. 2B),a p air of the chromosomes (nos. 19-22). largest submetacentric chromosomes (nos. 1& 2) C. triste Maxim. The chromosome number were easily distinguished from the others. The was counted as 2n =4 0σig. lE). The chromo- karyotype was composed of 14 metacentric (nos. somes were 3.5-1.1μm long and relatively February 1991 Journal of Japanese Botany Vol. 66 No. 1 29 輔醐鵬 Fig. 1. Somatic metaphase chromosomes. A: Carpesium abrotanoides (2n =4 0 +2 B),B: C. rosulatum (2n=40),C : C. glossophyllum (2n=40),D : C. macrocephalum (2n=40),E: C. triste (2n=40),F: C. cernuum (2n=40),G: C. divαricatum (2n =4 0),H : C. divaricatum var. matsuei (2n =4 0 +3 B),1: C. koidzumii (2n =4 0). In A and H,ar rows indicate the B-chromosomes,wh ile in 1,th e small metacentric SATchromosomes.Bar indicates 5μm. 剛 30 植物研究雑誌第66巻第1号 平成3年2月 Fig. 1. (Continued) symmetrical. In its karyotype (Fig. 2F),tw o pairs C. divaricatum Sieb. et Zucc. The chromo- of the largest submetacentric chromosomes (nos. some number was counted as 2n=40 (Fig. 10). 1-4) were easily distinguished from the others. The chromosomes were 3.6-1.1μm long and The karyotype was composed of 22 metacentric more or less symmetrical. In its karyotype (Fig. (nos. 7-10,13 & 14,1 7 & 18,23 & 24,2 9-40), 20),t wo pairs of the largest submetacentric 16 submetacentric (nos. 1-4,11 & 12,15 & 16, chromosomes (nos. 1-4) wぽeeasily distinguished 19-22,25 -28) and 2s ubtelocentric chromosomes from the others. The karyotype was composed of (nos. 5& 6). 26 metacentric (nos. 5-8,1 3-16,2 1-32,35 -40), February 1991 Journal of Japanese Botany Vo1. 66 No. 1 31 轟 事 A 畿聾轟 轟轟購麟醐 織購韓鵬 官事 襲撃事襲亀襲撃襲撃事 響 嚇 轡 襲撃議響護肇審議轟轟轟轟轟轟轟轟轟轟轟曹移譲 B 轟 議官 鑓 C 襲 撃 襲撃 轟暢 議畿間帯 犠審 事 車 D i 事顧 襲 E 議議 鑑醐帯 織臨轍 轟饗ザ 構轍麟欄糊轟鱒叩噛 嘩輔韓嚇織輔機 鵜輔韓議臨嚇 鶴難輔嚇轟雛轡 融構韓轍轡轟鵬線 轟轟糊嚇 嘩購醐珊 轟轟嚇 畿欝織物 轟轟柵嚇 轟輯帯 議輯盟穆 事襲撃事霊事襲撃襲撃襲撃襲撃亀襲 F 10 20 襲譲事事 事 事 審事轟轟 21 30 40 Fig. 2. Serial arrangement of the somatic metaphase chromosomes. A: Carpesium abrotanoides (2n =4 0 +2 B),B: C. rosulatum (2n =4 0),C: C. glossophyllum (2n =4 0),D: C. macrocψhalum (2n =4 0), E: C. cernuum (2n=40),F: C. triste (2n=40),G : C. divaricatum (2n=40),H : C. divaricαtum var. , matsuei (2n =4 0 +3 B),1: C. koidzumiiοn= 4 0). Arrows in 1i ndicate the small metacentric SAT - chromosomes. Bar indicates 5μm. 10 submetacentric (nos. 1-4,9 & 1 0,17 -20) and C. divaricatum var. mαtsuei Kitam. The 4s ubtelocentric chromosomes (nos. 11 & 12,33 chromosome number was counted as 2n = 40, & 34). 40+2B and 40+3B (Fig. 1H) coincident with 32 植物研究雑誌第66巻第1号 平成3年2月 事 襲撃 G 事B豊島品購 護事 韓融事亀鷺 轟習も寵撃事襲 H 1 聾 襲舗も轟轟轟,君臨轟轟書官襲龍 電島署島争事 轟覇欝謀 議事兵器亀 轟毒事 事A 10 20 〆 -舗も轟 事事富島麓 事軍 費量 導監 21 30 40 Fig. 2. (Continued) previous report (Arano,19 62) except existence of composed of 26 metacentric (nos. 5-12,17 & 18, B-chromosome. The length of the chromosomes 25-40),10 submetacentric (nos. 1-4,1 3-16,23 & was 3.6-1.1μm and their shape was relatively 24) and 4 subtelocentric chromosomes (nos. symmetrical. Two pairs of the largest submeta- 19-22). A SAT-chromosome was observed on centric chromosomes (nos. 1-4) were easily the short arm of two metacentric chromosomes distinguished from the others in its karyotype σig. (nos. 29 & 30),a lthough it was not always 2H). The karyotype was consisted of 24 meta- confirmed. centric (nos. 5 & 6,9 & 10,1 3-18,2 5-32, 35-40),12 submetacentric (nos. 1-4,7 & 8,11 & Discussion 12,23 & 24,33 & 34) and 4s ubtelocentric chromo- Among the eight species and one variety of somes (nos. 19-22). Two or three B-chromo- Carpesium,n o great difference was observed in somes were observed in one individual from Mt. their karyotype; all having 2n =4 0 chromosomes Takaosan population,T okyo σig. lH). that are more or less symmetrical,al though slight C. koidzumii Makino The chromosome differences are recognized among their chromo- number was counted as 2n =4 0 (Fig. 11) for the somal complements (Fig. 2). According to first time. The chromosomes of this species were Shimakawa (1952) and Arano (1962), C. 3.4-1.1p mi n length and comparatively smaller abrotanoides was reported as 2n =3 6 in chromo- than those of the others,a nd more or less some number,b ut there is ap ossibility of mis- symmetrical. 1n its karyotype (Fig. 21),t wo pairs counting because the chromosomes are too small. of the largest submetacentric chromosomes (nos. 1n Carpesium,t wo or four largest chromosomes 1-4) were recognized. The karyotype was with median or submedian centromeres are February 1991 Journal of Japanese Botany Vol. 66 No. 1 33 recognized in general in ak aryotype. radical leaves persist at the flowering time. Therefore,th e karyotypes can be classified into The second group including C. macroo句phalum, two groups. The first group including C. C. cernuum, C. triste, C. divaricatum, C. αbro的noides,C. rosulatum and C. glossophyllum divaricatum var. mαtsuei and C. koidzumii has two has ap air of the largest chromosomes (nos. 1& pairs of the largest chromosomes which are meta- 2). This group can be divided into two subgroups or submetacentric (nos. 1-4). This group can be by the centromeric position of the largest chromo- divided into three subgroups based on their some pair. One of these subgroups includes C. karyotypes. The first subgroup includes C. αbrotαnoides; the largest pair are metacentric and mαcrocephalum and C. cernuum; the largest pairs , all of the chromosomes are relatively discontinuous are metacentric (nos. 1-4) and the others are in size. This species is ab iennial herb and the only mostly metacentric. These two species have winged species having sessile heads. among the species petioles and semi-globose heads which are rela- examined. The other subgroup (C. rosulatum and tively large in size. The second subgroup including C. glossophyllum) can be distinguished from the C. triste is characterized by four large former in having as ubmetacentric pair of the submetacentric (nos. 1-4) and two large largest chromosomes and the chromosomes are subtelocentric chromosomes (nos. 5 & 6). In more or less continuous in length. In addition, addition,th is species is distributed in relatively high these two species have similar karyotype,es pecially altitude comparing with the other species and its in meta-or submetacentric chromosomes. In radicalleaves persist partially at the flowering time. morphological features,r osettes consisting of The third subgroup,w hich includes C. Table 2. Karyotypes and morphological characteristics of Japanese Carpesium. Species m sm st m-sm No. of Karyo- Morphological largest types characteristics chromosomes C. abrotanoides 20 8 2 10 2(m)コ一一-一ー A Sessile heads 1 C. rosulatum 14 22 4 • Rosette radical ぬm)_1- C. glossophyllum 16 16 8 • 2(Sm) A leaves 向 2コ C. macrocephalum 30 8 2 • Winged petioles & half-globose - B1 C. cernuum 28 8 4 • heads C. triste 22 16 2 • 4(sm)一一一 B Semirosette 2 radical leaves C. divαricatum 26 10 4 • 44〈{ssmm))-1ト-B3 Wingless petioles C. divaricatum 24 12 4 • & ovate or ovate 同 var. matsuei oblong low leaves C. koidzumii 26 10 4 • 4(sm) 34 植物研究雑誌第66巻第1号 平成3年2月 divaricatum,C. divaricatum var. matsuei and C. References koidzumii,is characterized by four large submeta- Arano H. 1962. Cytotaxonomic studies in subfam. centric chromosomes (nos. 1-4) and four small Carduoideae of Japanese Compositae,I V. subtelocentric chromosomes in the karyotypes. Karyotype analysis and its karyotaxonomic These three species have commonly ovate or considerations in genus Carpesium. La oblong-ovate lower leaves which are abruptly Kromosomo 53-53: 1762-1777 (in Japanese). narrowed on wingless petioles. Arano H. 1965. The karyotypes and the speciation Karyotypes and morphological characteristics in subfamily Carduoideae (Compositae) of of Japanese Carpesium are summarized in Table Japan. Jap. Journ. Bot. 19(3): 31-67. 2. Meiotic studies and more detail karyotype in- Kitamura S.,M urata G. and Hori M. 1981. vestigations will help to demonstrate the relation- Colored illustrations herbaceous plant of Japan ship among these species. 1. Revised ed. 71-72. Hoikusha Publishing Co.,L td. Osaka (in Japanese). 1a m grateful to Prof. M. Ono of Tokyo Shimakawa S. 1952. Cytotaxonomic studies of Metropolitan University,fo r his guidance and en- genus Cαrpesium. J. Biol. Hiroshima Univ. couragement during the course of this work and 4(1): 29-31 (in Japanese). also for his critical reading of the manuscript. My 要 旨 thanks are due to Assoc. Prof. M. Wakabayashi for his valuable comments and suggestions on this 日本産ヤブタバコ属Carpesium植物8種 1変 study. 1e xpress mys incere appreciation to all the 種について核型分析の再検討を試みた.従来,未 members of the Makino Herbarium,T okyo 角卒+斤であったホソバガンクビソウC.koidzumii Metropolitan University, for their generous において, 2n=40の染色体数が確認された. ま support,c ollections of plant materials and en- たヤブタバコc.αbrotanoidesは, 従来2n=36 couragements. Thanks are also due to Prof. W. と報告されてきたが,観察したすべての個体が2 Kim of Kyungpook National University in Korea, n=40で, 2n=36のものは確認できなかった. for his advice during this study. 核型分析より, この属の諸種は染色体組を構成す る特に長い染色体の数によって,大きく 2つの群 に分けることができた. 1つは2本の長い染色体 Endnote をもっ群で, ヤブタバコ ヒメガンクビソウc. 1) Present address: Department of Biology, rosulatum及びサジガンクビソウC.glossoρhyl- College of Natural Sciences,K yungpook Na- bumである.もう 1つは4本の長い染色体をもっ tional University,B uk-ku,T aegu,7 02ー701, 群で, オオガンクビソウC.macrocephalum, コ Korea.韓園大郎市慶北大肇校 自然科皐大皐 ヤブタノイコC.cernuumガンクビソウC.divαri- 生物撃科. αctum,ノッポロガンクビソウC.divαricatum var. matsuei及びホソパガンクビソウが属する. この 2つの群は長い染色体の形態によって, それぞれ 2つ及び、3つの亜群に分けることができる.

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