全 文 ：广 西 植 物 Guihaia 24(5)：422—425 2004年 9月
丰都车前的细胞学研究，兼论它的多倍体起源
王恒昌，孟爱平，李建强*，王 勇，
(中国科学院武汉植物园，湖北武汉 430074)
陶 勇
摘 要：对长江三峡库区特有濒危植物丰都车前(Plantago erosa var．fengdouensis)进行了细胞学研究，包括
有丝分裂和减数分裂两方面。丰都车前可能为一异源四倍体。其核型公式为 2n⋯4x 24 12m(2sat)+
12sm(2sat)，属于 2A型。极端的环境压力尤其是季节性的水淹可能是此四倍体物种形成的主要原因。对同
属内的另一物种车前(P．asiatica)进行了比较观察，确证车前为六倍体。该实验为丰都车前的保育遗传学研
究提供了细胞学证据。
关键词：丰都车前；三峡库区；多倍体起源
中图分类号：Q943 文献标识码：A 文章编号：1000—3142(2004)05—0422—04
Cytological studies of Plantago erosa var．
，· - ● ●J一 ● l ^en 2a0Uensls，WltJ Specia re／erence
t0 its polyploid origin
WANG Heng—chang，MENG Ai—ping，LI Jian—qiang ，
WANG Yong，TAO Yong
(Wuhan Botanical Garden，Chinese Academy of Sciences，Wuhan 430074．China)
Abstract：Cytological studies including mitosis and meiosis of Plantago erosa var．fengdouensis，an endangered
plant native to the Three-Gorge Reservoir Area of China，have been investigated．It may be an allotetraploid．
The karyotype can be formulated as 2n=4x=24= 12m(2sat)+ 12sm(2sat)，belonging to 2A type．Extreme
environmental impetus especially the seasonal flooding may play a principal role in the polyploid formation of
it．Chromosome number of P．asiatica is reconfirmed，and P．asiatica is a hexaploid．This study will supply a
cytological evidence for the further genetics conservation research of Plantago erosa var．fengdouensis．
Key words：Plantago erosa var．fengdouensis；Three-Gorge Reservoir Area；polyploid origin
1 Introduction
Plantago L．(Plantaginaceae)is a cosmopolite
genus．There are more than 1 9O species in total ge—
nus，and 2O occur in China(HU et a1．，2002)．Ac—
cording to the classification standard of Zheng et
a1．(1992)，P．erosa Wal1．ex Roxb．var．feng—
douensis Z．E．Zhao et Y．Wang was described．The
new variety is characterized by lanceolate，subin—
cised and trinerved leaves，flowers wit hout pedicel，
the styles of flowers being 4—～7 mm long and gla—
Received date：2003·‘09·‘24 Accepted date：2003·-12-24
Foundation item：Wuhan Botanical Garden director fund(OlO35lO8，010151lO)；Chinese Academy of Sciences(KSCX2一Sw一104)．
First auther：Wang Heng-chang(1967一)，Male，born in Xiantao city，Hubei Province，Doctoral student，Associate professor，
working in plant systematics and taxonomy．*Author for correspondence E-mail：jianqiangl@hotmail．com
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5期 · 王恒昌等：丰都车前的细胞学研究，兼论它的多倍体起源 423
brous，and flooding tolerant(Zhao et a1．，2002)．
Only two populations of the variety were discov—
ered in two small islands of two counties along the
Yangtze River． One is located in Fengdou of
Chongqing city，another in Zhongxian． There are
over 100 km between the two localities，which are
located in a submergible plain of the Three—Gorge
Reservoir Area of China．In December，2002，be—
fore the famous Dam began to retain water，one of
authors of this paper transplanted most of wild in—
dividuals of the variety into W uhan Botanical Gar—
den，Chinese Academy of Sciences(CAS)，and culti—
vated them under a semi—natural condition
． In
Plantago，the basic chromosome number is x=6 for
most species and polyploids are common from 2n=
4x一24 to 2n= 16x一 96(Fedorov，1974)．P．erosa
is a tetraploid with 2n=4 x=24(Hong and Zhang，
1990)．This study aims at through a cytological re—
search on the variety，(1)providing additional evi—
dence for the further research on genetic conserva—
tion and taxonomical re-evaluation of this endemic，
rare and endangered taxon of the Three-Gorge Res—
ervoir Area；(2)presenting an example for probing
into the origin and evolution of tetraploid group
that confined to some constricted ecological habi—
tat．For the later purpose，another species P．asi—
atica L．is also investigated．
2 Material s and Methods
The plants investigated were cultivated in W u—
han Botanical Garden，CAS．Young growing inflo—
rescences were obtained for cytological observation
and the voucher specimens were deposited in HIB．
For somatic chromosome observation，the ma—
terials were pretreated in a saturated solution of P—
Dichlorobenzene for 2．5 h before fixed in Carnoy
I(glacial acetic acid：absolute ethanol一1：3)at
20℃ for 1 h．For meiotic process investigation．
the young anthers were fixed in Carnoy I for 0． 5
h．Then we used wall degradation and hypotonic
treatment to prepare chromosome samples． More
than 10 metaphase cells were detected to determine
the chro
．
mosome number．The symbols classifica—
tion standard used to describe the karyotypes fol—
lowed Stebbins(1971)and Levan et a1．(1964)，the
karyotype asymmetry index followed Arano
(1963)．
3 Resuhs
3．1 Mitosis
The chromosome number of P．erosa var．
fengdouensis investigated in the metaphase was
counted to be 2n=4x=24(Plate I：3)．It would be
a tetraploid．The whole 24 chromosomes could be
roughly arranged into 6 groups of 4 homologues
but there existed obvious heteromorphy between
the two pairs of homologues in the third and sixth
groups(Plate I：10)．According Tanaka(1971，
1977)，the resting nuclei and prophase chromo—
somes of P．erosa var．fengdouensis were catego—
rized to be complex chromocenter type and intersti—
tial type respectively(Plate I：1，2)，the karyotype
can be formulated as 2n一 4x= 24— 12m(2sat)+
1 2 sm (2 sat)． The chromosome morphology and
karyotype are shown(PlateⅡ：10)．The karyo—
grams are listed in Table 1．The ratio of the lon—
gest chromosome to the shortest(L／S)is 1．44，and
the percentage of Arm ratio over 2：1 is 0． 375，so
the karyotype belongs to 2A type．The index of
karyotype asymmetry is 63．4 1 ．
3．2 Meiosis
The chromosome number of P．erosa var．
1engdouensis investigated at metaphases is consist—
ently counted to be 2n=4x=24(Plate I：5，6)．At
the prophase of meiosis，the chromosomes are dif—
fuse with chromatin slender and winded each other
(Plate I：4)．At metaphase I。the configuration of
P．erosa var．fengdouensis is almost uniform(Plate
I：5，6)．No tetravalent but bivalents occur，and
the bivalents are sticklike．It seems that P
． PrDsc￡
var．fengdouensis is more an alotetraploid than an
autoploid or a segmental allotetraploid．We also in—
vestigate the stages of anaphase I and anaphaseⅡ
of P．erosa var．fengdouensis。the division proces—
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424 广 西 植 物 24卷
ses are found to be normal(Plate I：7，9)，while
very seldom lagging chromosomes are observed
(Plate I：8)．Meanwhile，we make fl statistic of the
pollen grain fertility and find they are almost nor—
mfll also．
Table l The karyomorphological parameters
of P．erosa var．fengdouensis
The value is based on the means of measurements of the best five
cels． chromosome with satelite
Additionally，we observe the meiosis of P．ero—
sa var．fengdouensis and confirm the bivalent
counts of it are 18．The species would be fl hexa—
ploid with 2n=6x一36(Plate II：1l，12，13，14)．
4 Discussion and Conclusion
Confined to fl labile biota，P．erosa var．feng—
douensis is not an aggressive taxon，and may at
most be fl kind of moderate success of tetraploid．
The two seasonal islands(app．1 500× 150 m。)
where the variety is located remain to be land most
time of fl year while submerge when flooding oc—
curs from June to late August．The aerial portions
of the plants of the variety die back and disarticu—
late from the rootstocks at autumn with seedlings
that keep evergreen from winter to spring
． The
growth habits of P．erosa var．fengdouensis hence
present an adaptive mechanism of counter-seasonal
growing． Transplanted in W uhan Botanical Gfir—
den，it doesn’t spread beyond the place it is grown．
The average of seed germination is about 20 and
greatly lower than that of P．asiatica(app．95 )．
Except P．asiatica，fl vigorous and fertile weed eve—
rywhere，no other Plantago species were found
companying with the varieties all over the wild re—
gion．The two Plantago groups have fl vicariance
distribution．It is interesting whether the two taxa
have relation to each other in evolutionary origin．
The chromosome numbers of P．asiatica reported
by various authors seemed to be ambiguous(Chen，
1988，1989，2n=2x=12；Matsuo& Noguchi，1989；
Nishikawa，1990，2n一4x一 24；Probatova，1996，2n
一 6 x=3 6)．Thereby fl re—examination of its ploidy
is needed．
Since P．erosa var．fengdouensis is proposed to
be an allotetraploid，it may not derive from the dip—
loid simple doubling of P．asiatica even though the
later may possess diverse ploidy populations inclu—
ding diploid as Chen(1988，1989)reported．Obvi—
ously different in morphological characteristic and
ecological habit，genetic introgression in tetraploid
level between the two taxa seems to be impossible
．
Moreover，we examine P．asiatica is fl hexaploid，so
this assumed possibility can basically be excluded
．
Briefly，the ancestral diploid progenitors of P
． erosa
var．fengdouensis would have been extinct．Inter—
estingly，there are two P．erosa var．fengdouensis
populations distributed in two relatively isolated
islands along the Yangtze River，where the ecologi—
cal factors are found to be very similar
． This seems
to infer that environmental impetus especially the
extreme flooding has played an important role in
the origin of the tetraploid P．erosa var
． feng—
douensis．
The taxonomical position of P．erosa var
．
fengdouensis should be recognized，though wheth—
at it is fl variety or fl subspecies needs fl further
critical circumscription． From the cytogenetic
point of view analysed above，the taxon possesses
some relatively independent cytological，as well as
distinct morphological and ecological features．
Nonetheless，as Stebbins(1 979)pointed out，these
kinds of populations need to be studied more care—
fully，particularly with respect to differences in
genes coding for isozymes(or plus molecular mark—
er)and ranges of ecological tolerance．
Acknowledgements We gratefully thank Prof．Z．
E．Zhao for providing detailed field information．
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5期 王恒昌等：丰都车前的细胞学研究，兼论它的多倍体起源 425
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王恒昌，等：丰都车前的细胞学研究．兼论它的多倍体起源
WANG Heng-chang，et 8 ．：Cytological studies of Plamago ，_0sⅡvar fengdouensis
，with special reference to its polyploid origin
图版 I
Plate I
I一
- l|
·。 ● ’
。 r
一 一 ■—--L l 一 _
’—l r一 —I 1
l 嘲 ’，
，C、
恼 ＼ ▲ 广1I
-■■-J， ■，_r
-■I- 一{
_ _ — — ． J7 -◆1 - 一 IF ．▲ ’ lIr
一
◆
6 I
· 4 ．．◆ 5
◆ 够 毋
，
．
● ‘
’
’ ， 8 9
1～ 3．Mitosis of somatic chromosomesia P var
． fengdouensis-1．Resting nueteus；2．Prophase
3 Metaphase(Arrows show chrDIlr]O$oin~s with sateltite)，2n一4x一24 4～ 9 Meiosis stages
4．Prophase；5．Lateral view of Lietaphase l(Showlrtg 6 pairs of bivalents)}6．Apical view；
7·8．Anaphase I(Arrow showing lagging chromosome)：g．Anaphase I．
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王恒昌，等：
WANG Heng—chang．et 4
图版 Ⅱ
PlateⅡ
I ‘I 一：lI；jI
m sm m sm
I：Il：Il Ic i t,1]Il¨ jIf
m sm m sm
10
广 ≮
咚 §
4
10．Karyogram ofP．e,~o5~2 r． ngdouensi~．¨ ～14．bleiosis chromosome of P． osiatica．11，13．metaphasel
l2—14．Anaphase I．(Photos magnification：for Figs 1～10．×2 000·for Figs．11～14．×1 810)．
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