Table Of Content植物研究雑誌
J. Jpn. Bot.
66: 199-204 (1991)
Cytogenetic Studies on Wild Chrysanthemum sensu lato in China1). 1.
Karyotype of Dendranthema vestitum
恥1asashiNAKATAa，D e-yuan HONOb，J in-zhuan QIUb，H iroshi UCHIYAMAa，
a b
Ryuso TANAKAand Sing-chi CHEN
aBotanical Institute，Fa culty of Science，H iroshima University，H iroshima，73 0 JAPAN;
bLaboratory of Systematic and Evolutionary Botany，a nd Herbarium，In stitute of Botany，
Academia Sinica，X iangshan，Be ijing 100093，T he People's Republic of CHINA
中国産野生キク属(広義)の細胞遺伝学的研究 1.
1)
Dendrαnthemαvestitum (ウラゲノギク)の核型
中田政司a，洪徳元b，郎均専\内山寛a，田中隆荘a，陳心啓b
a広島大学理学部植物学教室 730広島市中区東千田町 1-1-89
b中国科学院植物研究所系統及進化植物学開放研究実験室
(Received on March 21，1 991)
The chromosome number of 2n = 54 and the meiotic chromosome configuration of 2711 at diakinesis
and metaphase 1in Dendranthema vestitum (Hemsl.) Ling，an endemic species of China，ar e reported
here for the first time. This species is similar to D. occidentali-japonense (Nakai) Kitam.
(= C hrysanthemum japonense Nakai) of Japan with r田pectto chromosome number (2n =5 4 =6 x) and
morphology of flowerhead，but the karyotypes of the two species are different from each other. On
回
the other hand，D. vestitum is relatively similar to the diploid (2n =18 ) D. japonicum (Makino) Kitam.
(=C . makinoi Matsum. et Nakai) of Japan with the common characters of the white-colored ligule
of the flower-head，th e simple-shaped leaf，mo rphology of the sat-chromosome，an d presence of sub-
terminal centromeric chromosomes. Since the chromosome complement of the hexaploid D. vestitum
is almost equivalent to three times as that of the diploid D. japonicum，t hese two species might be
closely related to each other.
Dendranthemαvestitum (Hemsl.) Ling (Fig. 1) in having white ligules and simple-shaped leaves.
is restricted to Henan，Hu bei and Anhui Provinαs， It seems valuable to compare karyotypes between
China (Shih and Fu 1983). This species is paid these two species when we detect and speculate
much attention to be studied in cytology because evolutionary history of Dendranthema in East
it has some resemblance to a Japanese diploid Asia.
(2n= 18) species，D . japonicum (Makino) Kitam. Wer eveal and report here the chromosome
(= C hrysanthemum mαkinoi Matsum. et Nakai)， number of 2n= 5 4 (hexaploid) and the karyotype
-199ー
200 植物研究雑誌第66巻第4号 平成3年8月
Fig. 1. A chromosome voucher (2n =5 4，N akata 9302) of Dendranthema vestitum collected
in Huanghuachang，n ear Yichang City，H ubei Province，C hina.
of D. vestitum in comparisons with those of Dendranthema vestitum in two populations near
Ja panese Dendranthema species. Yichang City，Hu bei Province; 18 plants in Jiang-
jiawan，l Okm northeast of Yichang，2 30 m in
Materials and Methods altitude，an d 12 plants in Huanghuachang，20 km
During the second field survey担China泊 1989 northeast of Yichang，1 90 m in altitude. The
for the 恥1onbusho International Research former population was found on ar oadside-slope
Program-J oint Research， Nakata， Qiu and along with an open pine forest associated with
Uchiyama collected 30 living individual-plants of Quercus，V ite.芯 Artemisia，Senecio，X anthium，
August 1991 Journal of Japanese Botany Vol. 66 No. 4 201
Kalimeris，P atrinia and so on. The latter popula- Measurements of chromosome lengths were
tion was seen on another roadside-slope with sne made on photographs magnified x3 ，000 by a
国
but more sunny environment than the former one， digitizer-personal computer system (Uc hiyama et
with association of Senecio，B idens，M iscanthus， al. 1988). Nomenclature for centromeric positions
Eupatorium and so on. Thirteen individual-plants ofthe chromosomes followed Levan et al. (1964).
of D. vestitum were introduced to Japan for the The voucher specimens were deposited in the
first time and cultivated in the experimental herbarium，In stitute of Botany，A cademia Sinica
greenhouse of Hiroshima University for cytological (PE).
studies.
Freshly growing root tips (ca. 1c m long) were Results and Discussion
obtained from the cultivated plants and pretreated Dendranthema vestitum showed morphological
in 8m M8 ・hydroxyquinolineat 200C for 3h r. variation: the individual plants of the Jiangjiawan
They were fixed in Farmer's fixative (3: 1m ixture population displayed that flower heads varied from
of ethanol and glacial acetic acid) at 50Cf or 20 hr 27 to 47 mmi n diameter. They also exhibited
before they were macerated in 2: 1m ixture of 450/0 variations of lengths (7.5-17.5 mm)，w idths
acetic acid and 1N HCl at 600C for 10 sec and (3.5-5.5 mm) and number (13-21) of ligulate
stained with 1% aceto-orcein for 10 min. Then， flowers，a nd of shapes (elliptic，o blong-ovate，
the meristematic tissu (ca.1m ml ong) of the root ovate or rhombate) and bases (acute，o btuse，
邸
tips were squashed with 1% aceto-orcein for attenuate or truncate) of leaves.
chromosome observation. At mitotic metaphase，2 n =5 4 chromosomes
For karyotype study，th e enzymatic maceration- (Fig. 2A) were observed in the 18 individuals of
flame drying method was also applied. Fixed root D. vestitum collected in the two populations
tips were transferred to deionized water through (Accession numbers: Nakata 9261，9 265，9 268，
decreasing ethanol series (70，5 0，3 0，a nd 15%). 9269，9270，9271，9272，9273，9278，9279，9280and
Then，th eir meristematic tissues (ca. 1m ml ong) 9281 from the Jiangjiawan population; and 9291，
were cut and placed onto slides and added one drop 9297，9 298，9 299，9 300 and 9302 from the
of enz戸nemixture (4% cellulase“ONOZUKA" Huanghuachang population). At diakinesis (Fig.
RS and 2%p ectolyase Y-23，p H4 .8). The slides 2B) and meiotic metaphase 1，2 n = 2711 chromo-
were incubated at 370Cf or 40 min in am oistened somes were also observed. Thus，th e above is the
petri dish. The root tips were rinsed with deionized first report of the chromosome number of D.
water and re-fixed with Farmer's fixative before vestitum. Since the basic chromosome number of
flame-drying. The chromosomes were stained with Chrysanthemum sensu lato is x= 9 ( Shimotomai
1% aceto-orcein for 5m in，ri nsed with tap water， 1933)，D . vestitum is ah exaploid (2n =5 4 =6 x)
and then air-dried. specIes.
Yo ung flower heads were fixed in modified Figure 2C shows as et of the individual chromo-
Carnoy' s fixative (2: 1: 1 mixture of ethanol， somes of D. vestitum arranged in decreasing order
chloroform and glacial acetic acid) and their of the length. Thus，t he 2n =5 4 chromosomes
anthers were squashed in 1% aceto-orcein.恥1eiotic varies gradually in length from 6.9 to 4.3μm. Six
chromosomes were observed at diakinesis and medium-to small-sized subterminal centromeric
metaphase 1i n pollen mother cells (PMC's). (st) chromosomes (Fig. 2C) were easily identified
202 植物研究雑誌第66巻第4号 平成3年8月
())cll'
c:Jlllil
f~'(lrJ:Þ，抑制{削減
Fig. 2. Chromosomes of Dendranthema vestitum. A. Mitotic metaphase chromosomes in
root tip cells of ap lant from the Jiangjiawan population (Nakata 9278). B. Meiotic
chromosomes at diakinesis (2n = 27II = 54) in PMC's of ap lant (Nakata 9265) from the
Jiangjiawan population. C. Individual chromosomes of ap lant (Nakata 9265) prepared by
enzymatic maceration-flame drying method. Asterisks show subterminal chromosomes. Stars
show submedian chromosomes with am inute satellite. All bars indicate 5μm.
from the others. Satellites often occurred on the (1981) described that the karyotype of D.
short arms of the “st" chromosomes and/or small- occ identafi.てjaponense(as C. japonense) had eight
sized submedian centromeric chromosomes (Fig. or ten subterminal centromeric chromosomes and
2C). four median centromeric chromosomes with
With respect to flower head with white-colored minute satellites.
ligules and the chromosome number of 2n =5 4，D. Dendranthema vestitum is morphologically
vestitum of China and D. occidentaliてjaponense similar to D. japonicum，a d iploid (2n =18 ) species
(Nakai) Kitam. (=C hrysanthemum japonense of Ja pan，bu t th位 chromosomenumbers are quite
Nakai) of Japan are closely related to each other. different from each other. According to Tanaka
However，th eir leaf shapes are quite different from (1959) and Nakata (unpublished data) ，D .
each other; the leaf of the former species is not japonicum has two pairs of diagnostic chromo-
lobed，w hile that of the latter species is cleft or somes often exhibiting satellites; a pair of
parted. Moreover，t he karyotypes of the two subterminal centromeric chromosom回 (mediumto
species are different from each other; Watanabe small in size) and ap air of submedian to sub-
August 1991 Journal of Japanese Botany Vol. 66 No. 4 203
terminal centromeric chromosomes (smal1 in size). (1978， 1981) for the Japanese genera Dend-
Thus，D . vestitum and D. japonicum have the αrnthema， Tanacetum， Leucanthemella and
common feature in karyotype. Tripling of the Nipponanthemum，al l segregated from Chrysan-
diploid chromosome complement of D. japonicum themum sensu lato for convenience. The referable
might simulate the hexaploid karyotype of D. names under Chrysαnthemum sensu lato used in
vestitum. Dendranthema vestitum seems to be our previous series (cf. Nakata et al. 1987) are also
karyomorphologically a close species to D. given in parentheses here.
japonicum concerning ap olyploid relation. This study was financial1ys upported by Grant-
Dendranthema japonicum is distributed in the in-Aid for the Monbusho 1nternational Research
southwestem part of Japan except for Kyushu and Program-Joint Research， No. 01044099 for
the westernmost-part of Honshu (Nakata and 1989-1991 (the representative; Ryuso Tanaka) of
Tanaka 1989). 1n contrast，D . vestitum grows on the Ministry of Education，Sc ience and Culture of
foothills or low mountains (340-1，500m in Japan.
altitude) in Henan，H ubei and Anhui Provinces in
China (Shih and Fu 1983). The distribution of D. References
vestitum seems to correspond to the transitional Hou H.Y. 1983. Vegetation of China with re-
subtropics from the w m-temperateregions (Hou ference to its geographical distribution. Ann.
紅
1983)，wh ile that of D. japonicum correspond to Missouri Bot. Gard. 70: 509-548.
the climatic regions of warm temperate to sub- Kitamura S. 1978. Dendranthema et人中'J)onanthe-
tropics in Jap an. The similarity of these distribu- mum. Acta Phytotax. Geobot. 29: 165-170.
tions may also be correlated with th位 phylogenetic 一一一1981.Compositae. In: Satake Y. et al.
relationship. (edよWildFlowers of Japan III. Heibonsha，
Tokyo (in Japanese).
Wew ish to express our gratitude to Mr. Yang Levan A.，F redga K. and Sandberg A. A. 1964.
Wang，M r. G凶-fuWang and Mr. Yan Chen， No menclature for centromeric position on
1nstitute of Botany，Ac ademia Sinica，Be ijing，fo r chromosomes. Hereditas 52: 201-220.
their help in the field work and cultivating Nakata M.， Tanaka R.， Taniguchi K. and
materials. We are indebted to Prof. Katsuhiko Shimotomai N. 1987. Species of wild Chrysan-
Kondo，H iroshima University，fo r critical reading themums in Japan: Cytological and cyto
国
of the manuscript. genetical view on its entity. Acta Phytotax.
Geobot. 38: 241-259 (in Japanese).
Endnote 一一一-and Tanaka R. 1989. Species of Chrysan-
1) A new series attached to the previous series themum in Japan in the study of chromosomes.
entitled “Cytogenetic studies on wild Chrysanthe- In Hong D.Y. (edよ PlantChromosome
mum from China (1-V)" (Tanaka and Taniguchi Research 1987. p. 17-22. Organizing Com-
1987，Ta niguchi and Tanaka 1987，T anaka et al. mittee of the SinoJapaneseSymposium on
・
1987a，1 987b，19 90)，si nce we follow here Shih and Plant Chromosomes，Be ijing and Hiroshima.
Fu's taxonomic treatment (1983) for the Chinese Shih C. and Fu G. X. 1983. Compositae (3)，
genera Dendranthema，A jania，T anacetum and Anthemideae. Flora Reipublicae Popularis
Leucanthemella and Kitamura's new treatment Sinicae 76(1). Science Press， Beijing (in
204 植物研究雑誌第66巻第4号 平成3年8月
Chinese). 要 旨
Shimotomai N. 1933. Zur Karyogenetik der Dendrαnthemα uestitum (Hemsl.) Ling
Gattung Chrysanthemum. J. Sci. Hiroshima (ウラゲノギク)は白色舌状花をもっ中国固有の
Univ. Ser. B，D iv. 2，2 : 1-100. キク属植物である.湖北省宜昌市近くの2集団で
Tanaka R. 1959. Ont he speciation and karyotypes
30個体のD.uestitumを採集し，観察した18個体
in diploid and tetraploid species of Chrysanthe-
の全てにおいて体細胞染色体数2n=54を算定し
mum II. Karyotypes in Chrysanthemum
た.また，花粉母細胞の減数分裂移動期，中期I
Makinoi (2n= 18). J. Sci. Hiroshima Univ. Ser.
において， 27個の二価染色体を観察した.キク属
B，Di v. 2，9: 17-31.
(広義)の染色体基本数は x=9であるので，D.
一一一-and Taniguchi K. 1987. Cytogenetic studies
estitumtま6倍体である.D. uestitumの染色体
on wild Chrysanthemum from China 1.
組には，他の染色体と容易に区別できる中型ない
Karyotype of Ch. lavαndulaefolium
し小型の次端部動原体型染色体が6個あり，付随
sianense. CIS 42: 10-12.
一一一一，一一一-andAoyama M. 1987a. ditto. 111. 体はこの次端部動原体型染色体と小型の次中部動
Karyotype of Ch. arisanense. CIS 43: 20-21. 原体型染色体にしばしば観察された.D. uestit-
一一一一， 一一一一， Nakata M. and Aoyama M. umtま白色舌状花を持ち， 6倍体である点で日本
1987b. ditto. IV. Karyotype of Ch. morii. CIS 産ノジギクD.occidentali-jαponense (N a kai )
43: 21-22. Kitam. (=C hrysαnthemum jα:ponense Nakai)
一一一一一， Kawasaki S.，Y onezawa Y.，T aniguchi と似ているが，葉の形態と核型は異なっていた.
K. and Ikeda H. 1990. ditto. V. Fchybrids of
一方，白色舌状花を持ち，葉の形が単純な点で，
Chrysanthemum lavandulaefolium var.
D. uestitumは日本産リュウノウギク D.jαponi-
siαnense x Ch. boreale. Cytologia 54: 365
cum (Makino) Kitam. (=C. mαkinoi Matsum.
372.
et Nakai)にも似ている. リュウノウギクは2倍
Taniguchi K. and Tanaka R. 1987. ditto. II.
体 (2n=18)であるが，次端部動原体型染色体
Karyotype of Ch. lavandulaefolium. CIS 43:
が2個観察され，付随体染色体の形態も共通して
18-19.
おり (Tanaka1959)， リュウノウギクの染色体
Uchiyama H.，N akata M.，M orihiro S. and
Tanaka R. 1988. Use of personal computer and 組を3倍すると， 6倍体の D.uestitumに似た
digitizer for karyotype analysis. La Kromosomo 染色体組になる.核型的に D. uestitumはリュ
1150:1652-1658 (in Japanese). ウノウギクと倍数性の関係を持つ近縁種で、あるこ
・
Watanabe K. 1981. Studies on the control of とが示唆される.
diploid-like meiosis in polyploid taxa of
Chrysanthemum 1. Hexaploid Ch. japonense
Nakai. Cytologia 46: 459-498.
