全 文 ：广 西 植 物 Guihaia 29(3)：285— 295 2009年 5月
天仙子族 (茄科)的花粉形态补充
研究及其系统学意义
张志耘1，杨冬之 ，Joongku Lee3 ，李 良千1*
(1．中国科学院 植物研究所 系统与进化植物学国家重点实验室，北京 100093；2．郑州大学 生物： 翌系，
郑州 450001；3．Korea Research Inslitute of Bioscience and Biotechnology，Daejeon 305—806，Korea)
摘 要：在扫描电镜下研究了茄科天仙子族 7属及颠茄属、茄参属和枸杞属 3属共 1O属 23种植物的花粉形
态学特征。结果表明：天仙子族的花粉粒形状为球形、近球形、扁球形和长球形 ；萌发孔类型有无萌发孔、不规
则的拟孔、3沟、4沟、3～4沟、散沟和 3孔沟．夕 壁纹饰为小刺状、瘤状、各式条纹状 、条纹一穴状、条纹一网状、
皱波状、细网状和网状。沟膜近光滑、具小颗粒、具小刺状突起或具瘤状突起。各属植物的花粉形态在萌发孔
的有无、萌发孔的类型和外壁纹饰等方面有较大的差异 ，可以作为探讨属问分类和系统关系的重要依据。天
仙子族植物花粉萌发孑L的演化趋势为：无萌发孔一3、4沟一3孔沟。无萌发孔的山莨菪属是原始类群，三孔沟
的马尿泡和天仙子等属是进化类群。还讨论了与前人研究结果不同之处和可能的原因。
关键词：花粉形态；天仙子族 ；茄科；系统学意义
中图分类号：Q94l 文献标识码：A 文章编号：1000—3142(2009)03 0285—11
Supplemental study on the pollen morphology of
the tribe Hyoscyameae(Solanaceae)and
its systematic significance ● ● - ●一●
ZHANG Zhi～Yun ，YANG Dong-Zhie，JOONGKU Lee3 ，LI Liang-Qian
(1．Herbarium (PE)＆ State Key Laboratory of Systematic and Ez~dutionary Botany，Institute 0，Bolany，the Chinese
Academy 0厂Sciences，Beijing 100093．China；2．Department 0r Bioengineer ing，Zhengzhou Uni ， sit3,，Zhengzhou
t50001，China；3．Korea Research Institute of Bioscience and Biote~hnolog3t，Daejeon 305 806，Korea)
Abstract：Polen grains of 23 species in seven genera of the tribe Hyoscyameae(Sohmaceae)and three related genera
(Alropa，Mandragora，Lycium)were im，estigated under SEM．The polen grains are globose，subglobose，oblate or
prolate in shape．Germination apertures were absent or present and are irregularly poroid，3-colpate，4-colpate，3-or 4一
colpate，pantocolpate or 3-colporate．The exine ornamentation was spinulose，tuberculate，variously striate，striate-fo—
veolate，striate-retieulate，rugose-sinuate，minutely reticulate or reticulate．The colpal membrane was nearly smooth
but with granular，spinulose Or tuberculate protuberances．The pollen grains show great、tariation between genera in
the Hyoscyameae in the presence and type of a germination aperture and exine ornamentation．These may be impor—
rant characters in rex，ealing phylogenetic relationship in the tribe．The evolutionary trend in germination apertures
ranges from absent or presence to 3-or,i-colpate then to 3-colporate．Anisodus，which lacks germination apertures，is
considered to be primiti~，e，while genera with 3-colporate polen grains，such as Przezc,alskia and H3，osc3’aIlUS are be—
lieved to be advanced．
Key words：polen morphology；tribe Hyosc5rameae；Solanaceae；systematic significance
收稿日期 ：2008—10—30 修回日期：2009—02—18
基金项目：lhe Plant 1)iversity Researcb Center ot the 21ST Frontier Research Program Funded by the Ministry of Education and．~ience and Technology
of Korea(PF 061O&O0)；因家 自然科学基金(30270094，30530050)
作者简介：张志耘(1950一)，女，广东广卅I市人，博士，研究员，主要研究方向为系统与进化植物学，(E-maiD zhangzy@ibcas．ac．cn。
。通讯作者(Author for corres1．,ondence，E mail：Iqli@ibcas．ac_cn)； ‘并列通讯作者(Amhor for c(卜corresf M】ndel1cc，E-mail：joongku@kribb．re．kr)
286 广 西 植 物 29卷
1 Introduction
Pollen morphology can be useful for the analyzing
taxonomic relationships in a wide variety of plant fami—
lies．Recent studies in the Solanaceae have shown pol—
len morphology to be of considerable systematic signif—
icanee，both at the generic and specific levels．
The tribe Hyoscyameae(Hunziker，2001)comprises
approximately 40 species in 7 genera~Anisodus Link(4
species)in China，Bhutan，India and Nepal；Archihyo—
scyam“s八 M Lu(1 species)in Western Asia(Kurdis—
tan)：Atropanthe Pascher(1 species)endemic to China；
tTIyoscyamus L (abo ut 23 species)widely distributed in
Northern Africa．Asia and Europe；Physochlaina G．Don
(about 11 species)in Asia：Przewalskia Maxim．(1 spe-
cies)endemic to China at high elevation on the Qinghai-
Tibetan Plateau：and Scopolia Jacq．(2 species)in Asia
and Europe． The center of diversity of the tribe is in
eastern Asia．with many species na tive to China．The
Hyoscyameae forms the only tribe in the family centered
in the northern hemi sphere． The diversity has been
linked to the Hima layan uplift(Lu& Chang。1986)．
The delimitation of the tribe Hyoscyameae has been
debated widely(Bentham，1876；Tetenyi，1987；D’Arcy,
1991：Hoare＆ Knapp，1997；Otmstead et a1．，1999；
Hunziker，2001)．The focus of this argument has been
on the affinity of Atropa to the traditionaly recognized
Hyoscyameae．Traditional classifications(Kuang& Lu,
1978：Tetenyi，1987；D’Arcy，1991)treated Atropa as
separate from the Hyoscyameae on the basis of the
tieshy，indehiscent fruits in Atropa and the unusual cir—
cumscissile capsules in the other genera．
The pollen grains of only a few taxa of Hyoscya—
rrleae have been examined in pre~ous studies(Basak，
1967：Punt&Clarke，1980；Sandina et a1．，1982；Diez&
Ferguson，1984；Chang& Lu，1984；Lu 8L Chang，1986；
Ying “Z．，1993；Hoare＆ Knapp，1997)and it has not
been evaluated for its taxonomi c utility wi thin a sub—
tribe, the tribe or within the Solanaceae as a whole
The present paper reports on the pollen grains of
10 genera，including 23 species of Hyoscyameae and
three related genera of Solanaceae(Atropa，Mandrag—
ora，Lycium)to assess its utility in providing a better
understanding of the systematic relationships of the
species within the Hyoscyameae and with other genera
in the family．
2 Materials and Methods
Pollen grains were removed from collections depo s—
ited in the herbarium of the Institute of Botany，the Chi—
nese Academy of Sciences(PE)，Nijmegen Botanical Gar—
den，Nimegen University(NL)，and the Natural History
Museum(P)，Paris，France(Appendix)．For scanning e—
lectron microscope(SEM)examination，the polen grains
were collected from drY specimens，mounted on stubs
with double-sided adhesive tape，then coated with gold
for 3—4 min．Observations were ma de with a Hitachi S-
800(SEM )． The pollen terminology follows Erdtman
(1952)．
3 Results
Characteristics of the
bers of the Hyoscyameae
pollen grains studied of mere—
and related taxa are舀ven in
Table 1．
The pollen grains are spheroidal，subspheroidal，ob—
late or prolate in shape．Germination apertures are ab—
sent or present and irregularly poroid，3一colpate，4-col—
pate，3一or 4-colpate，pantocolpate or 3-colporate．The ex-
ine sculpture is spinulose，tuberculate，variously striate，
striate-foveolate，striate-reticulate，rugose-sinuate，min—
utelv reticulate or reticulate．The colpal membrane is
nearly smooth but with granular，spinulose or tubercu—
late Drotuberances．Features of the po llen grains for the
various genera are as follows．
1$copolia(Plate I：1-3)
Polen grains spheroidal，3一colpate or rarely pantocol—
pate，exine sculpture densely rugose．
2 Anisodus(Plate II：13—16)
Polen grains subspheroidal or spheroidal，nonapertu—
rate，exine irregularly tuberculate or poroid．
3 Atropanthe(Plate II：17—18)
Polen grains spheroidal，3-colpate，colpi long，exine
curved striate．
3期 张志耘等：天仙子族(茄科)的花粉形态补充研究及其系统学意义 287
A}lis0d z rid s Subspheroidal Nonaperturate or occasionaly poroid
A．tanguticus
Atropanthe sinensis
Przetmlskia tangutica
Physochlaina praealta
P．physaloides
P．infundibularis
P．capitata
H．desertorum
H．pusillus
H．bipinnatisectus
H．muticus
H．seTwcionis
H．algFdlgS
H．albus
H．nige~‘
H．reticulates
Archihyoscyamus
calyx
Atropa belladonna
Spheroidal
Spheroidal
Spheroidal
Subspheroidal to
prolate
Spheroidal
Spheroidal
Subsphemidal to
prolate
Prolate
Prolate
Subspheroidal
Spheroidal
Spheroidal
Prolate
Prolate
Prolate
Prolate
lepto— Prolate
A~ndragora caulescens
A chinghaiensis
Lycium barbarum
Nonaperturate or po roid
3-colpate
3-colporate
3-colporate
4-colpate
3-colpate and 4-colpate
Indistinctly 3-colpate
3-colporate
3-colporate
3-colporate
3-colporate
polar area
3-colporate
polar area
3-colporate
3-colporate
3-colpo rate，
polar area
3-colpo rate，
polar area．
3-colporate
Subspheroidal or 3-eolporate
oblate
Spheroidal
Spheroidal
Prolate
Tuberculate，tubercles unequal in size，densely gran
ulate on surface
Tuberculate，tubercles unequal in size
Curveel striate
Reticulate
Shortly striate，occasionaly punctate，indistinctly tu—
berculate on striae
Rather densely spinose and irregularly foveolate or
punctate together
Densely spinose and sparsely foveolate together
Rugose，irregularly spinulate and punctate together
Striate and
Striate and
Striate and
colpi long，nearly reaching Striate and
sparsely punctate together
sparsely punctate together
sparsely punctate together
indistinctly foveolate together
colpi long nearly reaching Striate and sparsely punclate together
colpi long，nearly reaching
colpi long，nearly reaching
Nonaperturate
Nonaperturate
3-colporate
Striate-foveolate，striae low
Indistinctly striate to rugose，and sparsely punctate
together
Finely reticulate，muri wide and sparsely verucate
together，luminae smal，unequal in size
Finely reticulate，muf wide and sparsely verrucate
together，luminae sma ll，unequal in size
Striate，irregularly punctate among striae，sparsely
granular on surface of striae
Striate-reticulate，nearly smooth on striate surface
Spinulate and tuberculate
Spinulate and tuberculate
Striate，irregularly foveolate among striae，irregular
ly granular at puncta bottom
4 Przewalskia(Plate I：4-6)
Pollen grains spheroidal，3-colporate，exine retic—
ulate．
5 Physochlaina(Plate III：19—30)
Pollen grains spheroidal，subspheroidal to prolate，3一
colpate，3-or 4-colpate，exine spinose or rugose．
6 Hyoscyamus(Plate IV：31—36；Plate V：37—45；Plate
VI：46～5l；Plate VII：55—60)
Pollen grains spheroidal to prolate，3一colporate，exine
striate or rugose．
7 Archihyoscyamus(Plate VI：52—54)
Pollen grains prolate，3一colporate，exine striate．
8 Mandragora(Plate II：10—12)
Pollen grains spheroidal， nonaperturate， exine
sparsely spinulate and tuberculate．
9 Atropa(Plate I：7-9)
Pollen grains subspheroidal or oblate，3-colporate，
exine densely tuberculate．
10 Lycium (Plate VII：61-63)
Pollen grains prolate，3一colporate，exine striate．
Based on aperture，the pollen grains of Hyoscya—
meae and related genera may be placed in four groups．
(1)Nonaperturate：Mandragora；(2)Nonaperturate or
occasionally irregularly poroid：Anisodus；(3)3一colpate，
4-colpate or pantocolpate：Scopolia，Atropanthe and
Physochlaina (except P．praealta)；(4)3-colporate：
Przewalskia，Hyoscyamus，Arc^ ^ osc “， “ ，Phys—
ochlaina praealta，Atropa and Lycium．
288 广 西 植 物 29卷
Plate I 卜3．Stopuli“l7【{ nic“；4-6．Pr： “如kiatangutic“；7-9．At㈨pa bell“d Ⅲ“；1，4：bar一1 7·7 mFi1
2：bar=2O mm；5，7，8：bar= 16 film；3，6，9：bar=20 nlm．
Based on exine sculpture，the pollen grains of Hy—
oscvameae and related genera can be divided into four
types as folows：(1)Rugose or tuberculate：Scopolia，
A sodus；(2)Striate，curved striate，striate-foveolate，
str’1at reticu1ate，striate~rugose：Atropanthe，Physoch—
ZⅡ 扎口praenh“Hyoscyamus(except H．niger and H．
r ￡ f“Z“t“ )，Archihyoscyamus，Atropa and Lycium；
(3)Spinose：Physochlaina(except P．praealta)，Man—
dragora；(4)Finely retictdal e，reticulate：PrzewaIskia，
Hyoscyamus niger and H．reticulatus
4 Discussion and Conclusions
Polen grains of Hyoscyameae vary greatly among
the genera in the presence or absence and type of ger—
mination aperture and exine sculpture．These features
can therefore be important in considering the phyloge—
netic relationships of the tribe．
3期 张志耘等：天仙子族(茄科)的花粉形态补充研究及其系统学意义 289
PlateⅡ 10 11．^ ， c，， ⋯ ““ 【 ，J ：12．^五 ，2(1n r1 (h， ^ir， i』；13一】4
． ／l，? ￡j ]ur ，“ ：15一】6
．
／l j “ ，“， ，“￡
17—18 it rfJ mthe t nsis． 10：bar一 1 7
． 7 nlIl1；1I，12，14，16，18：bar=20 nlIl1；13，17：b∈lr=t 6 nlnl；15：bar_2O Hln1
．
The exine sculpture of Anisodus and Sf(JP0li(』is
tuberculate，but the pollen grains of Anisodus are nonap—
erturate or occasionaly irregularly poroid(Hoare ＆．
Knapp，1997)，while in Scopolicl，the pollen grains are 3一
colpate or pantocolpate．Morphologicaly，the flowers of
both genera are radial symmetrical，their stamens are e—
qual in length and inserted at the base of the∞rolla
tube，indicating that tile tWO genera hax，e a close re1a
fionship．Altisodus，howex-er，is illore primitive tt】a【l Sf(J—
polia． The nonaperturaIe anti t uberculate exine polen
grains of Scopolia(’arnioli~oides(Chang ．I u，1984)are
identical with the pollen grains of Anisodus，but different
from other species of Scopolia
． Data from polens there—
fore support the incorporation of S
． carniolicuide in
Anisodus(D’Arey Zhang，1992)．
The 3一colpate apertures of Atropanthe are similar
tO tlOSe in SOI-le species of Ph、 (J(^laina
． But in Atr(J—
panthe，the colpi are long and COIl＼，erge in the polar area
290 广 西 植 物 29卷
PlateⅢ 19 21．Ph3，sochlaina praealta；22—24．Physochlaina physaloid ；25—27．Physochl“ nⅡ 舶 “缸
19：bar=24 mrlrl；20，22，23。25，26：bar=17．7 mm；21，24，27：bar=20 mn1．
and the exine sculpture is curved striate，which iS dif—
ferent from that of Anisodus and Scopolia。The flow—
ers of Atropanthe，Anisodus and Scopolia are solitary，
but in Atropanthe the corolla is distinctly zygomorphic
and the stamens are unequal in length．Combined with
characteristics of the polen morphology，we considered
the genus Atropanthe to be more advanced than Aniso—
dus and Scopolia．
Przevaalskia is quite different from other genera of
Hyoscyameae in morphological and palynological fea—
tures．Przewalskia is endemic to Qinghai，Gansu，Si—
chuan and Xizang(Tibet)provinces，China，growing in
cold，arid environments at 3 2OO一5 000 m The axillary
flowers are solitary or 2 or 3 together and pedunculate or
not．The short stamens are adnated to the base of the
corolla tube，not attached to the corola throat．The ap—
erture is 3-colporate，which is the same as in Hyoscya一
US．The exine sculpture，however，iS reticulate and dif—
3期 查耘等：天仙子族(茄科)的花粉形态补充研究及其系统学意义 291
PlaleⅣ 川 ’7 yso ina“ pit ￡；3】
一 33． Ⅲ ；34 36
． Ⅲ
． 28 1
29，34：b 一l 7
- 7 n1n1；31，32，3S bar=】6 n1n1 30
．33，36：baf：2。n1n1．
ierent}rO1TI that of Hyoscyamus，which is st rjate
． We
considered Pr；re~rdskia to be a distinct，ad、，anced tax0rL
In Hyoscyamus the pollen grains are identica11v 3一
colP。rate and with a variously striate，or occasiona11v
t “。lY neulate exine sculpture
． The differences in de～
an be used for delineating species
． Morphologica卜
ly ll0wets are distinctly secund，sessile or subses—
slle,and lhe lower ones axillary and solitary with th
e
uPper ones lorming a dense racemose Or subscorDi0id
。ym。·The stamens are inserted near the middle。f the
coro1a tube． Hyoscy“ “ can be considered rather ad—
ranted in these features
．
1he polen grains of chi}lyOsc e 3-col∞ 一
ral e‘the same as in Hyoscyamus
，but the exine scu1Dture
strlate,ir egularly punctale among the striae and
parsely granular on surface of the striations
． T s obvi—
OUS dlt±ererlc~from Hyosc3 ㈣ “ supports tk：e re∞gni—
tlOn of Archihyoscyamus(Lu
，1997)．
292 广 西 植 物 29卷
Plate V 37—39．H3-OS(j “7n“ desertorum；40—42．Hyosc3一“，n“j reticulatus；43—45．H)一u“3一[1 7nus aurgu3．37，44：bar=24 FflFn
40．41，43；bar=20 mm；38：bar=17．7 mil；39，45；bar=20 1Tim；42：bar=15．8 toni．
The apertures of Physochlaina are usually 3一col—
pate or 4-colpate． The exine sculpture is spinose or
spinulate，and obviously different from other genera in
the Hyoscyameae．In P．praeaha，however，the aper—
tures are 3一colporate and the exine sculpture is shortly
striate，which is similar to that of Hyoscyamus and
may indicate that P．praealta is transitional between
Physochlaina and Hyoscyamus．
The pollen grains of Atropa are similar to that of
Hyoscyamus and supports treating it aS a member of the
Hyoscyarneae．In Mandragora，the nonaperturate pollen is
identical with Anisod“s，but the exine sculpture is spinulate
and tuberculate，which is different from Amisodus．In Ly—
cium，the exine sculpture is striate，irregularly foveolate a—
mong the striae and irregularly granular at the bottom of
the punetae．These features differ from members of the
Hyoscyameae．Lycium therefore has a distant relationship
wi th Hyoscyameae．
期 张志耘等：天仙子族(茄科)的花粉形态补充研究 墨竺兰 293
PlateⅥ 46-48．Hyoscyamlg6~l№ 49—51．协+州 -I， Ⅲ ；52 54． Ⅲn ·46，49，50Iba 2。
47：bar一 16 n1m；52：bar：1 3．6 mn1；53：bar=12 mrfl；48：bar一10 mm；51，54：b 2O ·
Chang I u(1 984)examined the pollen g m
of 12 species in 6 genera in the subtribe Hyoscyameae·
In the largest genus，Hyoscyamus，they only st dl d
tWO sDecies，making it difficult to deternfine the full ex—
tent of diversity and 、，ariation within the tribe．The
Dresent research found differences in the exlne “ P
ture from the previous report by Chang& 1 u(1984)，
details of which are gix ell in table 2．
1n the present study，the exine sculpture of Atro—
panthP sinensi is curved striate，while not shortly cla—
vate． In Physochlaina praeaZta，the exme sculpture s
sh0rtly striate，not reticulate．In P．physaloides，the ex—
ine sculpture is densdy spinose，not striate-reticulat ·In
Hy(J cv“l?TltS pusillus，t1e exine sculpture 1S strlate’not
reticulate( bk 2)． lhe differences rrmy be due to the
re1atively few species studied by Chang& Lu(1 984)．
The evolutionary trend in the germnatlon aperture
of the polen grains in the Hyoscyameae may be in the
294 广 西 植 物 29卷
Plate 55—57．F{ 。“3w，月“ bipinnafise
．
c
：
t
：
us；58-60．H
：
yoscyamus m“
5
ti
7
tu
，6
s
。
；
，
6
6
1
3
-
：
6
b
3
a
．
r
L
：
y
2
c
。
iu
m
m
m
ba
．
r6“r“?H-55，56：b。 17·7“
58 59 61 20 wtnl；62 bar=24 mm； ， ， ： ，’oU ’nJ u 【一 ⋯
directi0n from absence of germination aperture to pres—
ence 0f 3一or 4一colpate，to 3 colporate germma 0n aper
ture．A dus，which has nonaperturate Poll n gr m ’
should be pri tive，while genera vdth 3-colporate po儿en
grai嘶 such as PrzewaIskia and Hyoscyamus’shouId be
advanced．This evolutionary trend is basically consistent
with the interpretation from morphology-
Appendix：List of taxa examined
1．Scopolia加ponica
Japan：H。nshu，T．Yahata “f．，5936，1980—04—26(PE)
2．A isodus luridus
China：Cult．In Northwest-Plateau Inst．Bot-CAS，L Anming
01，1974—07—28(PE)
3．A．tanguticus
TheNetherl础 ：CultinBot．Card．NijmegenUBiv,13o．96／124(
4 AtYo n ￡^e sinensis
The N therIands：cult in B0t．Gard．N megen UniV·no·8g1。。
(NL)
5 Przez~xdskia tangutica
China：sang xiong，Xizang，Fu Guoxun 184，1960—06—6(PE)
3期 张志耘等：天仙子族(茄科)的花粉形态补充研究及其系统学意义 295
6．P．praealta
China：Malashan，Xizang，Wu Zhengyi et a1．，75-456，197,5 07—0(PE)
Table 2 Diferences in exine sculpture between
this study and previous report by Chang 8．
Lu(1 984)in the tribe Hyoscyameae
7．P．physaloides
Mongolia：colecting place unknown，八 八 Hatob 17533，19,14(PE)
8．Physo~hlaina ifundibularis
China：Huashan，Shaanxi．Yao Fuhan et( f．，sine no．(PE)
9．P．f“pitata
China：Gongliu．~njiang，Zhou Tmyan et a1．，650279，1965一O5_1O(PE)
1O．H．dese~ torut?t
Colecting place and date unknown．H．Gombavh 4983(P)
l1．Hyoscyanlus pusilus
China：Xinyuan，Xinjiang．colector unknown 69112-009，l969_O 27(PD
l2．H．b pinnaticase
Iran：South part．Th．Kotschy sine no．1842-06—10(P)
13．H．mutic“
Iran：South part．R．F．Hohenacker l8_I5，l842—03—13(P)
l4．H．seneciosis
Iran：North part，J．Bornmuler 7648，1902～06—08(P)
1 5．H3roscyamus aureus
Cyprus：collector an(I no．unknown，1880．date unknown(P)
16．H．albus
Syna：F F Maristesd’AI en．colector unknovcn 1947．dale u．nle~ n(P)
17．H．niger
China：Fukang．Xinjiang．colector unknown 4256，1959-09-19(PE)
l8．H．retieMates
Iran：East part，I ．Sintenis．sine no．1 89 i．date unkno~r(P)
1 9．Archihyoscyamus leptocalyx
Kurdistan：P．Sintenis 1000．1909一O5—16(P)
2O．Atropa belladomza
China：Cult．In Nanjing Bot．Garden．Yue Junsan 0358．1954—06一
O5(PE)
21．Mandrogora cau[estells
China：Zhongdian，Yunnan．T．T．Yu 11327．1937—05 17(PE)
22．M chinghaiensi~
China：Seda．Sichuan，colector unknown 06584(PE)
23．Lyeium barba； Ull
China：Cult．In BO1．Gard．Inst．BO t．CAS．Zhang Zhiyun＆．Wen
Jie 001，2002—06 05(PE)
*NL：Botanical Garden，Nijmegen University，Nijmegen，Neth—
erlands．
Acknowledgments
We greal y thank Dr．D．E．Bo ufor(1 of Harvard University
for valuable improving English language and Mr Xiao Yin Hou，
Mr Yang XueJian，Miss Yang Wen—Jing of Institute of Botany，
the Chinese Academy of Sc[ences for their kind help．
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