全 文 ：中国横断山区狭义百合科四属部分植物核型研究?
高云东 , 周颂东 , 何兴金??
(四川大学生命科学学院 , 四川 成都 610065)
摘要 : 用细胞压片法对分布于中国横断山区的百合科 4 属部分植物进行了核型研究 , 其中百合属 ( Lilium)
中除卷丹 ( L . tigrinum) 为三倍体 2 n= 36 外 , 大部分都是 2 倍体 , 2 n = 24 ; 贝母属 ( Fritillaria) 全为 2 倍
体 , 2 n = 24 , 其次缢痕较多 , 且比较明显 ; 假百合属假百合 ( Notholirion bulbuliferum) 同时存在 2 倍体和 3
倍体 , 2 n= 24 以及 2 n= 36; 洼瓣花属 ( Lloydia) 只研究了一种 , 即西藏洼瓣花 ( Lloydia tibetica) , 发现 2 n
= 23。4 个属的核型区别明显 , 为确定属间亲缘关系提供了一定的参考。其中 , 马塘百合 ( L . matangense)
和西藏洼瓣花的核型是首次报道。
关键词 : 染色体 ; 细胞分类学 ; 百合科
中图分类号 : Q 942 文献标识码 : A 文章编号 : 0253 - 2700 (2009) 05 - 399 - 07
Karyotypes of Four Genera in Liliaceae ( s. str .) from
Hengduan Mountains of Southwestern China
GAO Yun-Dong, ZHOU Song-Dong, HE Xing-Jin
* *
( Collegeof LifeScience, Sichuan University, Chengdu 610064 , China)
Abstract : Chromosome numbers andmorphologyof 4 generaof Liliaceae ( s. str .) fromHengduanMountainsof southwest-
ern China were studied . Almost all populations in Liliumwere diploid, with 2 n = 24 , except that populations of L . tig-
rinumwere found triploid, with 2 n= 36 . All taxa in Fritillaria investigated were diploid, with 2 n= 24 and the secondary
constrictionswereobvious and abundant . Diploidandtriploid exist in different populations of Notholirionbulbuliferum, with
2 n= 24 and 2 n= 36 , and there was one population contained both ploidy types . For Lloydia, onlyonetaxon Lloydia ti-
betica was studied, with2 n= 23 . All taxa studiedwerecollected in Hengduan Mountains, andthe differences amongthese
genera were obvious, which would contribute to thefuture taxonomic study . The karyotypes of L . matangenseand Lloydia
tibetica were documented for the first time .
Key words: Chromosomes; Cytotaxonomy; Liliaceae
The family Liliaceae ( s. str .) consists of 9 gen-
era ( Gegea, Lloydia, Tulipa, Erythronium, Fritil-
laria, Notholirion, Cardiocrinum, Liliumand Nomo-
charis) , and approximately 550 species distribute pri-
marily in the north temperature zone (Liang, 1995 ) .
Although plants of four genera ( Lloydia, Fritillaria,
Notholirion, and Lilium) dealt in this research were
studied before, karyotypic analyses are still limited .
For example, investigations in Lilium were conducted
in only some limited regions of China, andmostly con-
cernedwith a few populations of some species ( Li and
Gao, 1991; Tolgor and Liu, 1996; Wang et al. ,
云 南 植 物 研 究 2009 , 31 (5) : 399～405
Acta Botanica Yunnanica DOI : 10 .3724?SP. J . 1143 .2009.08179
?
?? ?Author for correspondence; E-mail : xjhe@ scu. edu. cn
Received date: 2008 - 09 - 12 , Accepted date: 2009 - 02 - 22
作者简介 : 高云东 (1985 - ) 男 , 在读研究生 , 主要从事染色体进化研究。 ?
Foun ?dation items: National Natural Science Foundation of China (grant no . 30670146 ) , National Infrastructure of Natural Resources for Science
and Technology ( grant no . 2005DKA21403)
1993; Yang et al. , 1997; Yu et al. , 1996a) . All the
previous researches confirmed that the number of the
chromosomes was 2 n = 24 , except Lilium tigrinum
with 2 n= 36 . Thegenus Notholirion contains fivespe-
cies (Liang and Minoru, 2000) . Threeof themaredis-
tributed in China ( N. bulbuliferum, N. campanulatum
and N. macrophyllum) , and they all are found in
Hengduan Mountains . Research on this genus was
scant, and the published data showed that Notholirion
bulbuliferumwas diploid (Wang et al. , 1993; Xu et
al. , 1986; Yu et al. , 1996b) . Triploids were detect-
ed in N. campanulatum ( Yu et al. , 1996b) , and
nearly no records were available for N. macrophyllum
yet . Fritillaria is confronted with thesame situation as
Lilium does, there was some but not sufficient re-
searches on it (Duan and Zheng, 1987; Wang, 1992;
Wang et al. , 2006; Zhang et al. , 1992 ) . Lloydia
was much less investigated due to the difficulty of ob-
taining samples .
Buxbaum (1936) studied the phylogenic relation-
ships among thesegenera . Hesuggested that Gegeawas
the most primitive genus and Lloydia originated from
Gegea and probablywas the ancestor of the rest genera
in the group . Liang (1995 ) supposed that there were
very close relationships among Lilium, Fritillaria,
Notholirion, Cardiocrinum and Nomocharis . In this
study, karyotypes of plants of Lloydia, Fritillaria,
Notholirion, and Lilium collected from the Hengduan
Mountains, a region recognized as a‘biodiversity hots-
pot’in the world (Myers et al. , 2000 ) , were studied
to find out the differences among genera as well as to
accumulate karyotypic data .
1 Material and methods
All the samplesstudied were collected in thefield (Table 1)
andcultivated in pots for cytological study . For each population,
roots were obtained from at least two individuals . Fresh root tips
were collected between8: 00 and10: 00 am, and pretreated in a
1∶1 mixtureof 0.1% colchicine and p-dichrlorobenzene at ambi-
ent temperature (10 - 20℃) for 8 - 10 hours, thenfixed in Carnoy
I ( glacial acetic acid: 100% ethanol = 1∶3 ) at 4℃ for 2 - 24
hours . They werethenmacerated in1 mol L - 1 HCl at 60℃ for 6
minutes, and stained and squashed in Carbolic acid Fuchsin .
Table 1 Origins, habitats and vouchers of species and populations of Liliaceae ( s. str .) examined
Species Habitat and Altitude ( m) Location in Sichuan Province Collector & voucher ( SZ)
Lilium
L ?. matangense Thicket slope, 3188 4Matang, Maerkang Y . D . Gao G07009
L ?. duchartrei Forest margin, 2790
Moist place in forest, 3100 ?Daofu, Ganzi Y . D . Gao G07039
L ?. regale Rocky slope, 1250 ?Wenchuan Q . Wang G07026
L ?. tigrinum Hillside, 1260 Beichuan Q . Wang G07029
Grass slope, 2850 ?Songpan H .Y . Liu G07040
L ?. lophophorum Alpine slope, 3650 ?Daocheng Y . D . Gao G07042
Fritillaria
F ?. cirrhosa Alpine grassland, 3560 Tangfang, Wolong Y . D . Gao G07035
Alpine thicket, 4230 OQuershan, Dege Y ?. D . Gao G07037
F ?. unibracteata Alpine thicket, 3660 OMengbishan, Maerkang Y . D . Gao G07019
Forest, 3800 UMucheng, Maerkang Y ?. D . Gao G07018
Alpine meadow, 4120 oZhegushan, Maerkang Y ?. D . Gao G07015
Notholirion
N ?. bulbuliferum Forest margin, 3600
Thicket, 3600 ?Huangtuliang, Pingwu Q . Wang G07003
Lloydia tibetica Alpine grassland, 3550 ?Tangfang, Wolong Y . Yu G07028
004 云 南 植 物 研 究 31 卷
Karyotype formulae were based on measurements of fine
metaphase chromosomes taken from photographs . The chromo-
somes of at least 30 cells were counted and the measurements of
at least 3 cellsweremade . Nomenclaturefor thecentromeric po-
sitions of chromosome introduced by Levan et al. (1964) were
followed: m= median centromeric chromosome with arm ratio of
1 .0 - 1 .7 ; sm= submedian centromeric chromosomewitharmra-
tio of 1 .7 - 3 .0 ; st = subterminal cetromeric chromosome with
arm ratio of 3 .0 - 7.0 ; t= terminal centromeric chromosome with
arm ratio of morethan7 . The karyotype classification of Stebbins
(1971) and the indexof asymmetry (AsK% ) defined by Arano
(1963) were adopted .
The voucher specimens were deposited in the Herbarium of
Sichuan University (SZ) , andthe voucher numberswere listed in
Table 1 .
2 Results and Discussion
The chromosomes counts, ploidy levels and karyotype
formulaeof all populations examined are listed in Table 2 .
The chromosomesof all species areillustratedinFigs.1 - 3,
which reveal their morphological traits and size .
2 . 1 Lilium matangense J . M . Xu
The karyotype of this species described by Xu
(1985) is reported for the first time . The 3A type and
relatively low asymmetry index ( 79 .3% ) showed dif-
ferences from other species in Lilium . It indicates that
L . matangensewas primitiveaswell as L . lophophorum
(Table 2 ) in Lilium .
2 . 2 Chromosome number and size
It is clear that these four genera inour study held
asame base number with n = 12 . All studies showed
that Lilium, except L . tigrinum with 2 n = 36 , were
diploids with 2 n = 24 . The diploid L . tigrinum was
only distributed in Korea Peninsula and Japan (Noda,
1986; Kim et al. , 2006) . All the populations of this
species fromChina were triploids (Li and Gao, 1991;
Tolgor and Liu, 1996; Yang et al. , 1997 ) . Based on
the number of chromosomes and its distribution pat-
tern, we concluded that L . tigrinumwas originated in
Korea Peninsulaor Japan, and dispersed into mainland
of China . Noda (1986 ) had studied the origination of
L . tigrinum and believed that this species came from
the hybridization between diploid L. tigrinumand L. max-
Table 2 Karyotype structureof populations and species of Liliaceae ( s. str .)
Species Karyotype Type As K% ( % ) Voucher ( SZ)
Lilium
L ?. matangense 2 ?n= 2 x= 4m+ 14st ( 2sat ) + 6t 3 lA 79 .2 G07009 Q
L ?. duchartrei 2 ?n= 2 x= 2m+ 2sm ( 2sat ) + 6st + 14t (2sat) 3 nB 84 .0 G07010 Q
2 ?n= 2 x= 2m+ 2sm ( 2sat ) + 8st + 12t (2sat) 3 nB 83 .0 G07013 Q
2 ?n= 2 x= 2m+ 2sm ( 2sat ) + 6st + 14t (2sat) 3 nB 83 .6 G07022 Q
2 ?n= 2 x= 2m+ 2sm ( 2sat ) + 8st + 12t (2sat) 3 nB 82 .6 G07024 Q
L ?. davidii 2 ?n= 2 x= 4m+ 8st (2sat ) + 12t 3 nB 81 .3 G07021 Q
2 ?n= 2 x= 2m ( 2sat ) + 2sm+ 6st (4sat ) + 14t ( 2sat) 3 nB 83 .0 G07032 Q
2 ?n= 2 x= 2m ( 2sat ) + 2sm+ 8st (4sat ) + 12t ( 4sat) 3 nB 82 .5 G07039 Q
L ?. regale 2 ?n= 2 x= 2m ( 2sat ) + 2sm ( 2sat ) + 6st ( 2sat ) + 14t (2sat) 3 nB 83 .5 G07026 Q
L ?. tigrinum 2 ?n= 3 x= 6m ( 5sat ) + 12st + 18t 3 nB 82 .3 G07029 Q
2 ?n= 3 x= 3m ( 3sat ) + 3sm ( 3sat ) + 12st (3sat ) + 18t (3sat) 3 nB 83 .1 G07040 Q
L ?. lophophorum 2 ?n= 2 x= 2m ( 2sat ) + 2sm+ 14st (4sat ) + 6t 3 lA 79 .0 G07042 Q
Fritillaria
F ?. cirrhosa 2 n= 2 x= 2m+ 4sm ( 2sat ) + 4st (2sat ) + 14t ( 2sat) 3 nB 80 .4 G07035 Q
2 ?n= 2 x= 2m+ 2sm ( 1sat ) + 10st (6sat ) + 10t ( 2sat) 3 lA 80 .6 G07037 Q
F ?. unibracteata 2 n= 2 x= 2m+ 2sm ( 1sm ) + 6st (2sat ) + 14t ( 2sat) 3 nB 81 .6 G07019 Q
2 ?n= 2 x= 4m+ 6st (2sat ) + 14t (2sat) 3 nB 81 .5 G07018 Q
2 ?n= 2 x= 2m+ 2sm ( 1sm ) + 6st (2sat ) + 14t ( 2sat) 3 nB 81 .5 G07015 Q
Notholirion
N ?. bulbuliferum 2 n= 3 x= 3m+ 3sm+ 12st + 18t 3 nB 83 .6 G07001 Q
2 ?n= 2 x= 2m+ 8st + 14t 3 nB 84 .3 G07004 Q
2 ?n= 3 x= 3m+ 3sm+ 9st + 21t 3 nB 84 .0 G07004 Q
2 ?n= 3 x= 3m+ 3sm+ 12st + 18t 3 nB 84 .6 G07005 Q
2 ?n= 2 x= 2m+ 2sm+ 6st + 14t 3 nB 84 .2 G07006 Q
2 ?n= 3 x= 3m+ 3sm+ 9st + 21t 3 nB 84 .1 G07007 Q
2 ?n= 2 x= 2m+ 8st + 14t 3 nB 84 .5 G07003 Q
Lloydia tibetica 2 ?n= 2 x= 1m+ 8sm+ 8st (2sat ) + 2t 3 nC 70 .9 G07028 Q
1045 期 GAO Yun-Dong et al. : Karyotypes of Four Genera in Liliaceae ( s. str .) fromHengduan Mountains . . .
Fig . 1 Karyotypeof Lloydia tibetica ( G07028 , 2 n= 23 ) .
Scale bar = 10μm
imowiczii , which had a close phylogenic relationship
with diploid L . tigrinum . But therewas controversy on
this conclusion because there was no evidence showing
that these two species shared habitats ( Kim et al. ,
2006) . Our research results support Noda′s opinion .
Wefound both two populations held the same trait that
the second pair ( Fig. 2 : 3 , Table 2 ) included two
chromosomes out threewith a secondary constrictionon
their short arms, while the rest one had none . It indi-
cates that the two had secondary constrictions which
probably came from the diploid L . tigrinum while the
other one was not . Others′research ( Li and Gao,
1991) also showed this phenomenon . Thus, we infer
L . tigrinum is an allopolyploid .
Six populations of Notholirion bulbuliferum in this
study contained both diploid and triploid samples . The
differences among diploids and triploids were few
(Fig.2 : 5 , 6 , Table 2) .Thekaryotypes weresimilar to
thoseof Lilium which confirmed the close relationship
between the two genera . Thedifferences among popula-
tions were not obvious . There was a stable trait on the
second pair, which included the same two sm chromo-
somes and a smaller one . This could be the result of
chromosome deletion, but also indicates the possibility
of hybridization in this complex . However, theZhuokeji
(G07005) population′s second pair combined three dif-
ferent sm chromosomes . This might be the result of
translocation . This pattern could be formed by three
equal chromosomes, inwhich a fragment of one chromo-
some is transferred to a different chromosome . These
findings imply that N. bulbuliferumis an autoploid .
In this study Lloydia tibetica was a dysploid, with
2 n= 23 . There was a metacentric chromosome, and this
longest one couldn′t be paired (Fig.1) .This chromosome
could be formed by centric fusion of two subterminal or
terminal chromosomes . Theoriginal number should be 2 n
= 24 . It′s typed 3C, which mean it′s more asymmetry
than othergenerastudied by us . However, the As K% =
70.9% , which was obviously fewer than the others . The
asymmetry indexwas well inaccordancewith the relation-
ship of four genera studied, which showed that Lloydia
was themost ancestral one (Liang, 1995) .
2 . 3 Secondary constrictions
Inour research, therewerenosecondary constric-
tions in Notholirion . Though colchicines treatment
might not be very suitable for showing secondary con-
strictions (Aarestrup et al. , 2008 ) , we believe there
are no secondary constrictions . With the same treat-
ment, Lilium and Fritillaria showed more secondary
constrictions . The amount of secondary constrictions of
the species in Liliumstudied by us varies greatly . The
range of secondary constrictions is from 2 ( L . mat-
angense, Fig. 1) to 10 ( L . davidii , Fig. 3 : 2) . How-
ever, Fritillaria had more secondary constrictions and
mostly on the long arms . The only short-arm constric-
tionwas located on a chromosome of the second pair,
and this trait was stable in two species ( Fritillaria cir-
rhosa and F . unibracteata) in this genus ( Figs. 2 : 2 ,
4 , Table 2 ) . It was convincing that these two species
had a very close relationship . Wedisapproved that se-
condary constriction as a significant distinctive feature
among species, especially using traditional acetic orce-
in method . From the previous studies (Zhang et al. ,
1992; Wang et al. , 2006 ) and ours, it is clear that
the locations of secondary constrictions are unstable
among populations . However, there are some excep-
tions in somespeciesof Lilium . Thekaryotypes of four
populations of L. duchartrei were in accordance with
each other (Table 2) , yet another research on L . duc-
hartrei showed quite different karyotypes ( Huang and
Li , 1996) . Three populations of L . davidii seemed to
bedifferent because thelocation andnumber of second-
ary constriction ( Table 2 ) . Yu et al . ( 1996a) also
showed differences among populations .
204 云 南 植 物 研 究 31 卷
Based on the data available, it is clear that few
aneuploidy and polyploidy existed in Liliaceae, espe-
cially in genera Lilium and Fritillaria . Thus, aneu-
ploidy and polyploidy could not be the main driving
forceof the evolution in these two genera . The large
amount of secondary constrictions and their variable lo-
cations in Lilium and Fritillaria could be the evolu-
tionary mark, or, the result of differentiation . However,
Fig . 2 Chromosomes of Lilium, Fritillaria and Notholirion
1 . L . matangenes ( G07009) ; 2 . F . unibracteata ( G07015 ) ; 3 . L . duchartrei ( G07010 ) ; 4 . F . cirrhosa ( G07037 ) ; 5 . N. bulbuliferum
( G07004) , 2 n= 24; 6 . N. bulbuliferum ( G07004 ) , 2 n= 36 . Scale bars= 10μm . The arrows show the secondary constrictions
3045 期 GAO Yun-Dong et al. : Karyotypes of Four Genera in Liliaceae ( s. str .) fromHengduan Mountains . . .
Fig . 3 Chromosomes of Lilium
1 . L . davidii ( G07021) ; 2 . L . davidii (G07039) ; 3 . L . tigrinum ( G07040) , 2 n= 36; 4 . L . lophophorum ( G07042 ) ;
5 . L . regale ( G07026 ) . Scale bars = 10μm . The arrows show the secondary constrictions
there was little regularity which could help to tell the
differences between the two generamentioned above . It
might be caused by the defects of research methods .
As mentioned above, therewas no secondary con-
striction found in Notholirion . Yu et al . ( 1996b)
showed it did exist, although not obviously . In con-
trast, polyploidy was very common . Thus, polyploidy
might be amain force in differentiation of Notholirion .
404 云 南 植 物 研 究 31 卷
Although aneuploidy existed in Lloydia tibetica,
more investigations are needed to confirm it . Our data
suggested that Lilium, Fritillaria and Notholirion could
be parallel evolution after deriving from their common
ancestor, Lloydia, as suggested by Buxbaum (1936) .
2 . 4 Genera relationship
There was no doubt that Lilium, Fritillaria and
Notholirion were very similar to each other on karyo-
type, which indicates close relationships of these three
genera . Based on karyotypes, it suggested that Lilium
and Fritillaria were stable while Notholirion and
Lloydiawerestill under a differentiationprocess . How-
ever, the aneuploid Lloydia tibetica could be an indi-
vidual phenomenon, and more individuals are needed
to be investigated to find out if euploid exists . It was
clear that Lloydia tibetica was more symmetrical than
other genera, and, thus, could be more ancestral in
Liliaceae ( s. str .) according to Stebbins (1971) . The
main differences between Lloydia and the other three
were the asymmetry and the chromosome size as well as
the absence of two giant metacentric chromosomes,
which were present and stable in other three genera .
The only metacentric chromosome in Lloydia tibetica
could begenerated by centric fusion . So it waspossible
that the individual with 2 n = 24 existed in natural
world . If so, the karyotype of Lloydia tibetica would
formulate as 2 n= 2 x= 8sm+ 10st ( 2sat ) + 2t with-
out metacentric chromosomes . Further investigations
areneeded to confirmthis hypothesis .
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