全 文 ：微体古生物学报　 2000年 6月 ; 17( 2): 228- 233
Acta M icr opala eonto log ica Sinica　 17( 2): 228- 233; June, 2000
* 国家教委优秀年轻教师基金资助项目
收稿日期　 2000-01-13
毛冠菊属的花粉形态和结构及其系统学意义*
张小平
(安徽师范大学生物学系 ,芜湖 241000)
提　　要
　　对国产菊科特有属毛冠菊属的花粉形态和结构进行了光学显微镜、扫描电镜和透射电镜的
观察和研究。该属的花粉可以分为圆球形、扁球形和长球形三种类型。种之间的花粉差异主要表
现在大小、形状和外壁纹饰的细微变化等方面。与紫菀族、千里光族和旋覆花族三个族的代表种
的花粉特征进行对比分析 ,并结合其它生物演化证据 ,可得到以下结论: ( 1)支持 Hand-Mazzetti
的分类意见 ,毛冠菊属应归于紫菀族中 ; ( 2)毛冠菊属的花粉可划分为圆球形、扁球形和长球形
三种类型 ,其中圆球形占多数 ,并且是较原始的类型 ,由此演变成其它两种形状的花粉 ; ( 3)根据
花粉由近球形到长球形或扁球形的演化趋势 ,赞成传统形态分类将 Nannoglottis gynura归于较
原始的类群 -狭苞组的分类处理。
关键词　　菊科　紫菀族　毛冠菊属　花粉形态和结构　系统位置
POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF THE
GENUSNANNOGLOTTIS AND ITS SYSTEMATIC IMPLICATIONS
Zhang Xiao-ping
( Departmen t of Biology , Anhui Normal Universi ty , Wuhu 241000 )
Abstract
　　 This study presents a detailed surv ey and discussion on the pollen mo rpholo-
gy and st ructure of Nannoglottis with light microscopy ( LM ) , SEM, and TEM.
Measurements o f the polar axis and equa to rial diameter, exine thickness, spine
leng th, and spine number at each lobe are made. Dif ferences w ere observed bo th
betw een representativ es of dif ferent t ribes and between species of the same
genus. The principal dif ferences in pollen among the va rious species w ere in size,
shape and subt le va ria tions of the surface ornamentation. Three kinds of po llen
ty pes can be recognized in the w hole genus, namely, spheroidal-type,
prola te -type and obla te-type. Combined wi th bo th palynolo gical and biosystem-
atic evidences, the data suppo rt Hand-Mazzetti s taxonomic t reatment that put
Nannoglottis in the tribe Astereae, ra ther than under the t ribes Senecioneae or
Inuleae.
　　 Key words　 compositae, astereae, Nannoglottis , po llen mo rpholog y, systematic
posi tio n
1. Introduction
　　 The genus Nannoglottis Max im , which is endemic to the no rthw est of China, includes
only 8 species, all of which are perennial herbs and are mainly found under high mountain
fo rrest at about 3 000 meters above the sea lev el. The genus is t raditional ly diagnosed by
i ts heterogamous capitula wi th one series of peripheric fertile ligula te pisti llate flow ers,
and many sterile hermaph rodite f lo rets, in combina tion wi th 3 - 4 series of invo lucral
bracts and the rounded anther bases. N annoglot tis has a very iso lated posi tio n in Composi-
tae due to i t s rest ricted geog raphic dist ribution and speciali zed g rass mo rpholog y described
above. The question to which t ribe Nannoglot tis should be assigned has remained unre-
solv ed so fa r. Three kinds of view points can be summarized for the tax onomic t rea tments
o f Nannoglottis . 1. M aximov icz ( 1881) considered N annoglot tis as an abno rmal g enus o f
the tribe Inuleae on the basis o f it s g eneral appearance somewhat simi lar to that of some
genus in Inuleae, although the anther base is wi thout tail; 2. Hof fman ( 1894) , how ever,
did no t ag ree wi th Maximovicz opinion, and paid much at tention to the signi ficance of the
cha racteristics of the untai led anther base and the steri le hermaphrodi te f low ers, t ransfer-
ring N annoglott is into subtrib. Senecioninae o f the t ribe Senecioneae f rom the t rib. In-
uleae. This sy stematic t reatment w as accepted and adopted by Ling Rong and Chen Yi-ling
( 1965) , who noted that one o f the major generic fea tures, viz. , the presence of inner
tubulate pisti llate f low ers in Nannoglottis , w as neglected by all early bo tanists. 3. Han-
del-Mazzet ti ( 1937) indicated that the flowers o f N annoglot tis are t rimorphous, and com-
pared this character wi th that o f Conyza L. And Erigeron L. of the tribe Astereae, thus,
he inclined to place Nannoglottis in the tribe Astereae but wi thout knowing its clo se rela-
tiv es in that tribe. In thei r cladistic analy sis of Astereae Zhang Xiaoping and Ka re Bremer
( 1993) follow ed Hand-Mazzet ti , reco gnizing N annoglot tis as an isola ted genus in Aster-
eae.
　　 Obviously, in o rder to understand clea rly the phylog eny of N annoglot tis , the o ther
data besides the gro ss morpho logical evidences have to be pursued. The aim o f our present
investigation w as at examination o f the po llen wa ll morpho log y and ult rastructure o f 8
species o f N annoglot tis , to see if they have the regularly perfo ra ted inner layer of the
229　 2期 张小平: 毛冠菊属的花粉形态和结构及其系统学意义
Astereae proper, the character which is the key to distinction betw een Astereae, Inuleae
and Senecioneae. It is our belief tha t this study will furnish the relevant informa tion to the
understanding of i t s sy stematic po sition in the Compositae, and to the elucidating the
species relationships wi thin the genus.
2. Materials andMethods
　　 All o f the pollen grains examined w ere taken from the anthers of specimens mounted
on herba rium sheets. Pollen w as prepared for LM , SEM , TEM observ ations and measure-
ments. Fo r LM , the standard aceto lysis method dev ised by Erdtman( 1969) was used. The
pollen samples w ere acetoly zed and mounted in glycerol jelly fo r observ ations, some LM
photog raphs of the microcha racters w ere taken. Fo r SEM , nonacetoly zed pol len g rains
w ere placed on double stick tape and then sent into a vacuum chamber and coated wi th
gold-palladium , observa tions and pho tog raphs w ere taken w ith a JSM -6300 scanning elec-
tron micro scope. For T EM , mature anthers w ere placed in 1% OSO4 buffered wi th sodium
cacodylate, pH6. 8, at ro om tempera ture fo r 18 hours. Following dehydration in an ace-
tone series, the anthers w ere embedded in Spur r medium, sectioned wi th g lass kniv es,
stained wi th urany l aceta te in combination w ith lead cit rate, and observ ations and trans-
mission electron micro graphs w ere made wi th a HITACHI H-600 elect ron microscope.
Table 1　 List of pollen materials for this study
Taxon Locali t y No. of specim en Sou rce
N . ca rpesioides Yuzh ong County, Qingh ai 3325 PE
N . dela vayi Northw est Yunnan 12626 PE
N . gynura Long Chang val ley, Sichuan 9626 PE
N . h ieraciphylla Northw est Yunnan Unknow n PE
N . lati squama Huang Jia Ping , Yunnan 24749 PE
N . macrocarpa Eas tern Xizhang 5539 PE
N . sou liei Wes t Sichuan 73064 PE
N . yuannanensis Northw est Yunnan 1875 PE
3. Results
　　 Table 2 presents the quanti tativ e measurements and quali ta tiv e description for po llen
fea tures of all the species studied. With LM and SEM th ree di fferent shapes of the po llen
occur in Nannoglott is , they a re spheroidal ( see PlateⅠ , fig s. 6- 9; PlateⅡ , fig s. 1-
10; PlateⅢ , fig s. 6- 10; Pla teⅣ , figs. 1- 5) , pro la te( Pla teⅠ , fig s. 1- 5; PlateⅢ ,
figs. 1- 5) , and oblate( PlateⅣ , fig s. 6- 10) , av eraging 29. 15× 28. 22μm in size. The
pollen o f all species is trico lpo rate wi th mo re or less lalongate o re. The surface is unifo rm-
ly spinulose, w ith 22 spines in equa to rial cross-section view and 5 spines a t each lobe in
polar view , and the spines are g radual ly pointed, lanceo late, and some of them possess
basal perfo rations ( po res) ; wi th TEM , the fine st ructure of Nannoglottis pollen w all i s
230 微　体　古　生　物　学　报 17卷　
consistent fo r all species studied( see Pla teⅤ , fig s. 1- 6) . Exine caveate( Pla teⅤ , figs.
1, 3) ; endexine tw o-layered, upper lamella te, low er homogeneous but disrupted near
pores; foo t lay er thin, uneven, about one-third thickness of endexine; bases of co llumel lae
spreading or div ided, joined to fo rm a thick basal layer that is occasionally discontinuous;
w ith sub-columellae o ften apparent beneath spines; internal fo ramina most ly elonga ted
common throughout co llumellae and tectum( Pla teⅤ , figs. 1- 6) .
Table 2　 The features of pollen morphology of Nannoglott is species
Taxa
Mean
polar axis
(μm )
Mean
equatorial
axi s (μm)
P/E
Pollen
shape
Am b Aperture
Spin e
leng th
(μm)
Exine
thickness
(μm)
Spine
number
at each
lob e
N . capesioides 31. 9 27. 0 1. 18 Prolate 3-lobed ci rcu lar 3-colporate 3. 6 2. 3 5
N . delavayi 28. 5 26. 3 1. 09 Sph eroidal 3-lobed ci rcu lar 3-colporate 3. 6 3. 1 5
N . gynura 25. 7 26. 4 0. 98 Sph eroidal 3-lobed ci rcu lar 3-colporate 2. 9 2. 1 5
N . hieraciph yl la 31. 9 29. 5 1. 08 Sph eroidal 3-lobed ci rcu lar 3-colporate 2. 3 2. 2 5
N . lat isquama 28. 5 24. 0 1. 19 Prolate 3-lobed ci rcu lar 3-colporate 3. 8 2. 0 5
N . macrocarpa 28. 1 27. 4 1. 03 Sph eroidal 3-lobed ci rcu lar 3-colporate 3. 3 2. 6 5
N . soul iei 31. 5 34. 2 0. 92 Sph eroidal 3-lobed ci rcu lar 3-colporate 2. 7 2. 5 5
N . yuannanensis 27. 0 31. 1 0. 87 Oblate 3-lobed ci rcu lar 3-colporate 2. 7 1. 9 5
4. Discussion and Conclusion
　　 For th rowing some ligh t on questions of the posi tio n o f Nannoglott is w ithin the Com-
posi tae and of i ts phylo geny wi thin the genus, the comparison of pollen morpho log y of the
genus wi th tho se of sev eral rela ted tribes w as conducted ( see Table 2 and Table 3) . As
show n in the table, the po llen mo rpholo gy of N annoglott is is much closer to that of the
t ribe Astereae than to tha t of the t ribe Inuleae and Senecioneae in the pollen shape, size,
and spine number at each lobe, especially completely identical to Astereae rather than to
Inuleae and Senecioneae with reg ard to the fine structures of the pollen wal l which a re em-
ployed by Skvarla et al. ( 1977) as the tw o key cha racters in classi fica tion of po llen pa t-
terns in Compositae. The fi rst one is internal foramina by which Helianthoid pa ttern is dis-
tinguished from the Senecioid pat tern that is cha racterized by the absence of internal
fo ramina. According to Skvarla et al. , the t ribe Astereae belongs to Helianthoid pat tern
w hi le the t ribe Inuleae and Senecioneae to Senecioid pat tern. From our observa tion,
Nannoglottis perfo rates wi th conspicuous elongated internal foramina th roughout columel-
lae and tectum. This na ture o f the pollen w all indica ted tha t Nannoglottis has very close
af fini ty wi th the t ribe Astereae, both of them are assigned to Helianthoid pat tern. The
second one is the spine shape. The surface of the po llen in Nannoglottis is echina te wi th
thin, g radually sharp-pointed f rom the ba se to the tip, lanceola te spines which is homolo-
gous wi th that o f Astereae, but not wi th that o f the t ribe Inuleae and Senecioneae which
have stout, ba se-turgid, conical spines.
231　 2期 张小平: 毛冠菊属的花粉形态和结构及其系统学意义
Table 3　 Comparison of the pollen characters of Nannoglottis with Astereae, Inuleae and Senecioneae
Taxa Pollen shape
Mean
diameter
(μm)
Spine
leng th
(μm)
Spine
n umber at
each lobe
Internal
foramina
Spine sh ape Cav us
N annog lott is Prolate, oblate, sph roidal 28 3. 0 5— 6 perfo rate un tu rgid at
spin e bas e
caveate
Astereae Prolate, oblate, sph roidal 27 3. 0 5— 6 perfo rate un tu rgid at
spin e bas e
caveate
Inuleae Prolate, sph roid al 26 3. 5 4— 5 nonperforate tu rgid at
spin e bas e
Rarely
caveate
Senecioneae Prolate, s phroidal 30 3. 1 3— 4 nonperforate tu rgid at
spin e bas e
caveate w ith
short bacula
　　 Thus, it can be, f rom a po llen point o f v iew , concluded that Nannoglot tis should be
placed correctly in the tribe Astereae, the posi tion of Nannoglottis in the t ribe Inuleae or
Senecioneae did not obtain the suppo rt f rom the evidences o f the pollen study.
　　 As fo r the relationship betw een species w ithin the genus, three types o f the po llen
shapes can be found: 1. prolate, P /E> 1. 14, the colpus membrane unseen, including N .
carpesioides , N . latisquama; 2. Oblate, P /E < 0. 88, the colpus membrane immersed in,
g ranular. Only N . yuannanensis included; 3. Spheroidal, P /E= 0. 88- 1. 44, the colpus
membrane immersed in, ei ther smoo th or granula r, most of the species covered, they a re
N . delavayi , N . gynura, N . macrocarpa, N . hieraciphyl la , N . soul iei . According to the
evo lutionary t rends o f pollen type of composi tae f rom spheroidal th rough obla te to pro la te,
the result of the present study supports the t rea tment o f N . gynura as the only member o f
Sect. Stenolepis Ling and Y. L. Chen, the rath er primi tive tax on in the genus.
References
Erdtman, G. , 1960. The acetolysis method-Svensk Botanisk Tidsk ri ft , 54: 561- 564
Handel-Mazzet ti , H. , 1937. Vierhapperia . Not izb l. Bot . Gart. Berlin. 13: 627
Hoffmann, O. , 1894. Composi tae in Engler, A. And K. Prant l, Pf lanzenfamilien IV , 5: 269- 394
Ling Yo ng & Chen Yi-ling, 1965. Genera nova vel minus cog ni ta familiae Composi tarum , II. Acta Phytotaxonomica Sini-
ca, 5( 1): 91- 104
Maximovicz. , C. J. , 1881. Diagnoses plantarum novarum Asiaticurum LV. Bull. Acad. Imp. Petersb. , 27: 480
Skvarla, J. J. , Tu rner, B. L. , Patel, V. C.& Tomb, A. S. , 1977. Pollen mo rphology in the Composi tae and in mo rpho-
logically related families. In: The Biology and Chemistry of the Composi tae 1( ed. V. H. Heywood, J. B. Harborne&
B. L. Turner ) . pp. 141- 248, Academic Press, London
Zhang Xiaoping & Kare Bremer, 1993. A cladistic analysis of the tribe Astereae ( Asteraceae) wi th notes on their evolu-
tion and subtribal classif ication. Pl. Sys t. Evol. , 184: 259- 283
图 版 说 明
PlateⅠ
1- 5. Nannoglot tis carpesioides
　　 1. Ploar view , SEM× 2700. 2. Equato ril view , SEM
× 2700. 3. Detail of s pines showing the spine shape
and indist inct perforations on the base of spine, SEM
232 微　体　古　生　物　学　报 17卷　
× 6000. 4. Polar cross s ection view , LM× 400.
5. Equatorial cross s ection view , LM× 400.
6- 10. Nannoglot tis delavayi
　　 6. Ploar view , Dis tinct perfo rations on th e bas e of
spine and tectum , SEM× 3000. 7. Equato ri l view ,
SEM× 3000. 8. Equatori l view at low focus , LM×
400. 9. Equato ri l view at high focus , LM× 400. 10.
Polar view at low focus , LM× 400.
PlateⅡ
1- 5. Nannoglot tis gynura
　　 1. Equatori l view , SEM× 3500. 2. Polar view , SEM
× 3300. 3. sho rt and s tout spines and wi th out perfo-
rat ion at the base of spines , and smooth tectum,
SEM× 9000. 4. Equatorial cross sect ion view , LM×
400. 5. Equatorial cros s s ection view at low focus,
LM× 400.
6- 10. N annog lotti s hieraciph yl la
　　 6. long and lanceoate spines and wi th out perforat ions
at th e bas e of spines, SEM× 9000. 7. Equatoril
view , SEM× 2700. 8. Equatoril cros s s ection view at
high focus, LM× 400. 9. Equato ril view at low fo-
cus , LM× 400. 10. Equatori l cross section view at
intermidium focus , LM× 400.
PlateⅢ
1- 5. Nannoglot tis lat isquama
　　 1. Equatori l view , SEM× 3000. 2. Polar view , SEM
× 2700. 3. Detail of s pines showing th e cu rve end of
th e spines and cours e tectum and indis tinct perfora-
tions on the base of spin e, SEM× 7500. 4. Equatori-
al view at high focu s, LM× 400. 5. Polar cros s sec-
tion view , LM× 400.
6- 10. N annog lotti s macrocarpa
　　 6. Show ing th e sharp end of th e spines and a few per-
fo rat ions on th e bas e of the spine, SEM× 9000. 7.
Equatori l view , LM× 400. 8. Polar cros s s ection
view , LM× 400. 9. Subequatori l view , SEM× 3000.
10. Polar view , SEM× 3000.
PlateⅣ
1- 5. Nannoglot tis soul iei
　　 1. Su bequatori l view , SEM× 3000. 2. Polar view ,
SEM× 3000. 3. Detail of s pines sh ow ing th e sharp
end of the spines and indistinct perforations on th e
bas e of spine, SEM× 9000. 4. Equatorial cros s sec-
tion view , LM× 400. 5. Polar cross s ection view ,
LM× 400.
6- 10. N annog lotti s yuannanensis
　　 6. Wi th out perforat ions on the bas e of spine and the
sm oo th tectum, SEM× 9000. 7. Polar view at low
focus , LM× 400. 8. Equato ril cross s ection view ,
LM× 400. 9. Sub equato ril view , SEM× 3000. 10.
Polar view , SEM× 2700.
PlateⅤ
1- 6. The f ine s t ructure of th e pollen wall under TEM.
　　 1. Nannoglot tis hieracip hylla, Sh owing sexine equal
to nexine and elongate internal foramina th roughou t
collumel lae and tectum. 2. N annoglott is gynura ,
Show ing sexine thicher than nexine and collumel lae
beneath spine. 3, 4. Nannoglott is yuannanensis, Indi-
cating cavus in area betw een apertures . 5.
Nannoglott is carpesioid es, Sh owing tectum and spine
base wi th indis tinct perforation s. 6. Nannoglott is
d elavayi , Sh ow ing spine base w ith dis tinct opens.
233　 2期 张小平: 毛冠菊属的花粉形态和结构及其系统学意义
张小平:毛冠菊属的花粉形态和结构及其系统学意义 图版 Ⅰ
Pollen M orpho logy and Detailed Structure o f the Genus N annoglot tis and i ts
Sy stematic Implications Pla teⅠ
张小平:毛冠菊属的花粉形态和结构及其系统学意义 图版 Ⅱ
Pollen M orpho logy and Detailed Structure o f the Genus N annoglot tis and i ts
Sy stematic Implications Pla teⅡ
张小平:毛冠菊属的花粉形态和结构及其系统学意义 图版 Ⅲ
Pollen M orpho logy and Detailed Structure o f the Genus N annoglot tis and i ts
Sy stematic Implications Pla teⅢ
张小平:毛冠菊属的花粉形态和结构及其系统学意义 图版 Ⅳ
Pollen M orpho logy and Detailed Structure o f the Genus N annoglot tis and i ts
Sy stematic Implications Pla teⅣ
张小平:毛冠菊属的花粉形态和结构及其系统学意义 图版 Ⅴ
Pollen M orpho logy and Detailed Structure o f the Genus N annoglot tis and i ts
Sy stematic Implications Pla teⅤ