全 文 ：植　　　物　　　研　　　究
BULLETIN OF BOTANICAL RESEARCH
第 19 卷　第 1 期 1999 年　1 月
Vol.19　No.1 Jan., 　1999
东北蒿属植物比较形态学研究
Ⅰ.种皮结构
张大维
(哈尔滨师范大学生物系 , 哈尔滨　150080)
摘　要　本文通过光学显微镜和扫描电子显微镜 ,对东北蒿属植物莳萝组(Sect.
Absiuthium DC.),艾蒿组(Sect.Abrotanum Bess.),艾组(Sect.Artem isia),龙蒿
组(Sect.Draunculus Bess),牡蒿组(Sect.Artm isia)35种代表植物种皮的形态特
征进行了观察。其中瘦果形状多为倒卵形 、长圆形及其它变形 ,但立体构形多有变
化。瘦果表皮细胞均有次生加厚:之一是表皮细胞垂周壁和平周壁极度加厚 ,凸起
相互联接构成网状结构;之二是果皮细胞壁次生加厚也比较强烈 ,在果实的表面形
成密集的纵棱轻微的纵向隆起 ,领细胞的层数和直径也有一些变化。这些微形态学
特征在组间存在着某些差异 ,具有一定的分类学和生态学意义。
关键词　蒿属;种皮形态;种皮微形态;扫描电镜
COMPARATIVE STUDYON MORPHOLOGYOF ARTEMISIA
PLANTS FROM NORTHEASTERN CHINA
I.MORPHOLOGY OF SEED COAT
Zhang Da-wei
(Department of Bio logy , Harbin Normal University , Harbin 150080 P.R.China)
Abstract　The morphology of seed coat f rom 35 representative species in g roups of
Sect.Absiuthium DC., Sect.Abrotanum Bess., Sect.Artem isia , Sect.Dracun-
culus Bess and Sect.Iati lobus Y.R.Ling from Artemisia were observed using both
optical microscope and scanning electron microscope.The shape of most achenes w as
inversely-ovate , lengthened-round or in other variable forms.However , the stereo-
哈尔滨师范大学校科研基金资助项目。
　蔡火石生物科学发展基金资助项目。
　收稿日期:1997-6-12
scopic configuration varied to certain ex tent.All the epidermal cells of achenes had sec-
ondary thickening.First of all , the vertical and the horizontal perines of epidermal cells
thickened ext remely and they pro truded so as to interconnect as netted structure.Sec-
ondly , the relatively intensive secondary thickening took place in the cell w all of the
seed coat so that the concentrated longitudinal ridges fo rmed on the surface of seed.
Certain variations oceurred interms of the diameter and the layer number of the collar
cells.The difference in these microst ructure characters among groups may be of impo r-
tance in plant taxonomy and ecology .
Key words　Artem isia;Morphology and microst ructure of seed coat , Scanning elec-
t ron microscope(SEM)
Artemisia , a genus in subtrib chrysanthem inae , Trib Arthemideae , composatae , covers
approximate 380 species in the wo rld.Almost 200 species of this genus exist in P.R.China in
w hich most of these species grow in the north and some areas of the southwest of the country.
About 40 species distribute in the northeast of China〔5〕.With the applicat ion of scanning elec-
tron microscope , more at tention is paid by plant tax onomists to the seeds of plants for their
complexici ty and large diversity of morphological and micromorphological characters 〔1-3〕.I t has
been shown that the achenes of Artemisia bear the tissue in the epidermis w hich w as retained
by the ripe plants of last generation , manifesting certain t raits of the ripe plants.The morpho-
logical study of achenes from Artemisia reflects the aff inity relationship of macrobiocoenosia
w hich formed during the sy stemat ic development of the genus.The present study w hich mainly
employed sacnning elect ron microscope , besides sibesides optical microscope , focused on sys-
tematically the morphological characters of epidemis of achenes in Artem isai plants located in
northeastern China and characterized the seed coat s decorat ive specif icity of this genus and the
discussion of the taxonomic importance of the specifici ty as w ell.
1.Materials and Methods
35 species in Artem isia were selected in this wo rk.The ripe achenes of these species were
all collected from the w ild field.The healthy , unbroken and naturally dry achenes w ere t reated
w ith 70%ethanol before microscopic observ ation.With optical microscope , the size(leng th by
w idth)of the achenes w as measured 30 seeds per species using eyepiece micromeasure.The av-
erage value w as taken from reading s of 30 samples.Weight of 100 seeds was measured with
balance.For preparation of the materials used in elect ron microscopic observation , the dried
samples were g lued with double-side glue tape to the sample holder of the scanning elect ron
microscope.The gilded samples w ere observed using Hitachi S-520.
2.Results
2.1　Observation wi th optical microscope
Achenes of Artemisia were mostly inversely -ovate , ellipt ic , leng thened-round or in
112 植　　物　　研　　究　　　　　　　　　　　　　　　19 卷
o ther varied shapes.Most of these achenes were red brow n , brown yellow or brown in colour.
The size of them did not differ significant ly.Some of the achenes w ere slightly f lat w ith elliptic
cross-section.Those that bear the above characters include A.adamsil Bess.A.annua
Linn.Some of other achenes w ere not flat but with closely round cross section.The species
that have such features include A .subulata Nakai , A .rubripes Natkai , A .argyi Leavl , et.
van , A.integri folia Linn.Some achenes formed several ridges in longitudinal direction with
i rregular polygonal cross section.The typical species in such catego ry is A .f rigida willd.
2.2　Observation wi th scanning elect ron microscope
In the present study , the terms from W.Barthlo tt were employed.That is the type of
cell s arrangements and the shape of the cells , includng the outline of cell w all , the st ructure of
the vertical perines , the curvature of the outw ard perines , etc.was called the primary decora-
tion.The microrelief sculpture , including the decorat ion of the cuticular membrane and the sec-
ondary thickening of the cell w all , was named as the secondary decorat ion.The observation of
different achenes w ith scanning elect ron microscope show ed that the decorations of achenes
from Artemisia plants g rowing in the no rtheast of China can mainly be classified as three cate-
go ries , i.e.netted type , longi tudinal-rib type and smooth type.
a.Net ted type
The achenes of this category were shaped in net ted form , the vertical and the horizontal
perines of the cell thickened ex tremely , prot ruded so as to form an irregular netted conf igura-
tion w ith other structures existing on the surface of the achene.The species w ith such features
include Artem isia sieversiana Ehrhart ex w illd , A .anethoides Mat t f , A .zhaodongensis G.
Y.Chang et M .Y.Lion , A .anethi folia web.ex.stechm and northeastern A adamsil
Bess.
b.Longitudianl-rib type
The decoration of achenes in this category w as characterized w ith intensive secondary
thickening of the cell w all , the prot ruding of the vertical perines and the forming of many co r-
rugation ridges in the horizontal perines.The achenes of this g roup can be further devided into
tw o subtypes:
transversely-lined and indist inct ly t ransversely -lined achenes.
(Ⅰ)Transversely-lined
Intensive secondary thickening of the cell s horizontal Perines took place in the achenes of
this subtypes.Many transverse lines can be seen on the surface of the achenes.The species in
this subtype include A .mongolica (Fisch.ex Bess).Nakov i , A .sellengensis Turcz .ex
Bess , A .igiaria Maxim , A .lavand ulaefolia DC , A.lancea van and A .annua Linn.
(Ⅱ)Indistinctly transversely-lined
The secondary thickening of the cell s ho rizontal perines was less intensive than that of the
achenes in the first subtype.In addition , the t ransverse lines were not so clear or observable as
those in the firs subtype comparably .The species of this subtype cover A.subulata Nakai ,
A .integrifolia Linn.A .sylvatica Maxim , A .argyi Leavl , et.van , A .keiskeana Miq
1131 期　　　　　　　　　　　　张大维:东北蒿属植物比较形态学研究
　　Table 1 　Morphological and micromorphological features of seed wat in Artemisia
plants from northeastern China
Name of Plant
Morphological features
colour shape size(mm) shape of cross
section
S tct.
Absinthium
DC
Ar temisia siever-
siana Ehrhartex.
w illd.
b row n - red
w ith black spots
inversely-ovate 1.583×0.764 app.round
A.anethi folia web.
ex.Stechm. g rey-yellow ellipt ic or lengthend 1.492×0.789
inregular plygou
or app.round
A. zhaodongensis
G.Y.Chang et.M.
Y.Liou
g rey-w hite ellipt icall inversely-ovate 1.377×0.707 app.elliptic
A.anethoides Mat-
tf. yellow-w hi te ellipicall inversly-ovate 0.860×0.437 ellpit ic
A.f rigida willd. b row n-red ellipicall inversly-ovate 1.526×0.774
inreyular poly-
gou or app.
round
Sect.
Abrotanum
Bess
A.adamsii Bess. silvery-grey ellipt ically inversely - o-
vate
1.530×0.605 ellipt ic or app.
round
A. tanacet i folia
Linn.
silversy brow n
-red
ellipt ically inversely - o-
vate
1.537×0.967 ellipt ic or cres-
cent
A.la ti fol ia Ledeb. yellow ellipicall inversly-ovate 1.828×1.015 app.elliptic
A.annua Linn.
light brow n -
yellow w rith
shine
ellipicall inversly-ovate 0.917×0.424 elli tic
A. sacrormun
Ledeb. g rey-brown inversly ovate 0.806×0.421 ellipt ic
A. sacrormun
Lebeb.var.messer-
schmidt iana (Bess)
Y.R.Ling
g rey-brw on lengthened inversely -o-
vate
0.715×0.423 ellipt ic or app.
round
A.palustri s Linn. b rw on inversely ovate 0.912×0.534 ellipt ic or app.
round
A.aurata Komar. black-brow n inversely ovate 0.821×0.492 ellipt ic or app.
round
Sect.
Artmisia
A. li ttoricola Ki-
tam. b row n ellipt ically inversely ovate 1.123×0.5935
ellipt ic or app.
round
A. lavandulaefolia
DC. b rw on-yellow
lengtheened ovate or in-
versdy-ovate 1.282×0.520 app.round
A.la ncea van. b row n-yellow lengthened elliptic or lev-
gthened round
1.065×0.324 app.round
A.rubr ipes Nakai. b row n-red legthened elliptic or shu t-
t le-like 1.404×0.420 round
114 植　　物　　研　　究　　　　　　　　　　　　　　　19 卷
Micromorphological features
rat io of
length to
w idth
w eight of a
hundred
seeds
conjugat ion of
pericarp and seed
coat
surface decorat ion
type
number of layers
of callar cell
diameter of col-
lar cell(um) plate No
2.07 24.08 relaxed netted type 3 120.4 1
1.87 29.98 relaxed netted type 3 84.1 2
1.94 23.48 relaxed netted type 3 102 3
1.97 8.62 easily separated netted type 3 66 4
1.97 28.36 relaxed I rregular bulge sub-
t ype
4 60
2.53 16.50 relaxed netted type 4 72 6
1.89 40.50 relaxed Bulge - present sub
type
4 120.4 7
1.80 48.73 relaxed Bulge - present sub
type
4 129 8
2.16 5.12 relaxed t ransversely Lined 3 48 9
1.91 8.20 relaxed netted type 7 81 10
1.69 19.25 relaxed netted type 5 72 11
1.70 21.32 relaxed indistinctly tram s
versely-Lined 3 96 12
1.66 10.96 relaxed indistinctly trans-
versely-Lined 4 72 13
1.87 18.65 relaxed netted type 3 78 14
2.47 16.70 relaxed t ransversely-Lined 4 126 15
3.29 8.11 relaxed t ransversely-Lined 5 102 16
3.35 14.77 relat ively tigh t indistinctly trans-
versely-Lined 7 100 17
1151 期　　　　　　　　　　　　张大维:东北蒿属植物比较形态学研究
Name of Plant
Morphological features
colour shape size(mm) shape of cross
section
Sect.
Artemisia
A.igiaria Maxim. dark b row n -
yellow
ellipt ic or lengthened 1.437×0.517 app.round
A.argyi Levl.et
van. b row n-yellow
lengthened elliptic or
langthened round
1.191×0.452 round
A. mongolica(Fisch. ex Bess.)
Nakai.
g rey brow n
lengthened in versely -o-
vate
1.395×0.620 round
A. integr i folia
Linn.
g rey brown -
yellow
inversely - ovate or
langthened round
1.628×0.570 round
A.subulata Nakai. b row n-yellow lengthened elliptic or
langthened round
1.440×0.507 app.round
A.selengensis Tur-
cz.ex.Bess. light-yellow
ellipt ically or langthened
round
1.838×0.522 round
A.keiskeana Miq.
silvery yellow -
b row n or silvery
red-b row n
inversely-ovate elliptic 1.623×0.891 app.elliptic
A. sylva tica Max-
im. b row n-yellow
ellipt ically or langthened
round
1.316×0.468 round
A. vir idi ssim a
(Komar.)Pamp. g rey-brown
ellipt ically inversely - o-
vate
1.215×0.4056 round
Sect.
Dracunculus
A. dracumculus
Linn. red-b row n
narrow ly inversely -ovate
or narrow ly ellipt ic
1.238×0.486 app.elliptic
A. ha lodend ron
Turcz.ex Bess. yellow-brow n inversely-ovate 1.998×0.685 app.elliptic
A. oxycephala
Kitag. g rey - yellow n-b row n lengthened inversely -round 1.594×0.657 app.elliptic
A. pubescens
Ledeb. b row n-yellow inversely-ovate 1.350×0.602 app.elliptic
A.scopar ia Waldst
et.Kit.
light.yellow -
b row n
inversely-ovate 0.742×0.338 app.elliptic
A. deser torum
Spreng.
dark b row n -
red
inversely - ovate or
lengthened in-versely -
ovate
1.245×0.689 app.elliptic
Sect.
lat ilobus
Y R.Ling
A.japonica Thunb. b row n-red inversely-ovate or ellipti-
caly in-versely-ovate 1.001×0.556 app.elliptic
A. manshurica
(Komar.)Komar.
dark b row n -
red
inversely-ovate or ellipti-
cally in-versely-ovate 1.195×0.631 app.ellipt ic
A. l it tor icola Ko-
mar.
yellow - brow n
or brow n-red inversely-ovate 1.191×0.648 app.round
116 植　　物　　研　　究　　　　　　　　　　　　　　　19 卷
Micromorphological features
rat io of
length to
w idth
w eight of a
hundred
seeds
conjugat ion of
pericarp and seed
coat
surface decorat ion
type
number of layers
of callar cell
diameter of col-
lar cell(um) plate No
2.78 17.05 relaxed indistinctly trans -
versely-Lined 4 102 18
2.84 10.30 relat ively tigh t indistinctly trans -
versely-Lined 4 90 19
3.02 12.39 relat ively tigh t t ransversely-Lined 4 127.5 20
2.85 23.69 relat ively tigh t indistinctly trans-
versely-Lined 4 129 21
2.84 17.53 relat ively tigh t indistinctly trans-
versely-Lined 4 145.2 22
3.52 14.71 relat ively tigh t t ransversely-Lined 7 906 23
1.82 41.07 relaxed indistinctly trans-
versely-Lined 4 206 24
2.81 11.35 relat ively tigh t indistinetly trans-
versely-Lined 4 90 25
2.99 12.65 relat ively tigh t indistinctly trans-
versely-Lined 8 172 26
2.55 10.13 tight Bulge - in - g roove
subtype
6 84 27
2.92 35.50 tight Bulge - in - g roove
subtype
3 120.4 28
2.43 19.90 relaxed Bulge - in - g roove
subtype
3 126 29
2.24 20.70 tight I rregular bulge sub-
t ype
4 120 30
2.19 2.28 tight I rregular bulge sub-
t ype
3 60 31
1.87 24.37 tight Bulge - in - g roove
subtype
6 84 32
1.80 14.00 tight Bulge - in - g roove
subtype
4 102 33
1.89 22.24 tight Bulge - in - g roove
subtype
3 84 34
1.84 24.37 tight bulge - in - g roove
subtype
5 90 35
1171 期　　　　　　　　　　　　张大维:东北蒿属植物比较形态学研究
and A.rubripes Nakai.
c.Smooth type
The secondary thickening of the cell w all in this type of achenes w as less intensive than
that of the netted type or longitudinal type.A few or no long itudinal ridges could be observed
on the ho rizontal perines.Certain vert ical perines may pro trude slightly.Clear and regular
transverse-lines in the form of “ ladder” fo rmed as a result of secondary thickening of the hori-
zontal perines.Based on the characters described above , achenes of this type can be further de-
vided into three subtypes , i.e.bulge-present subtype , bulge-ing roove subtype and irregular
bulge subtype.
(i)Bulge-present subtype
The neigboring cells bound closely so that some borders between cells could not be clearly
discriminated.On certain horizontal perines bulged the longitudinal ribs.The species of this
subtype contain A .lati fol ia Ledeb.and A.tanacet i folia Linn.
(ii)Bulge-in-groove subtype
The border between neigboring cells w as in the form of g roove in w hich there w as the pro-
truding longitudinal-rib structure and the arrangements of thread-like cutin.The shape of
epidermal cells w as clearly observed.The species wi th such features include A.japonica
Thunb.A.desertorum spreng , A .l ittorcola Kitam , A .manshurica (komar.)Komar , A .
dracunculus Linn , A .halodondron Turcz.ex Bess , A.oxycephala Kitag and A.f rigidaw
w illd.
(Ⅲ)Irreqular bulge subtype
The ho rizontal perines of the cells were concavely or convexly uneven or prot ruded irregu-
larly in the form of longi tudinal rib.The vertical perines may protrude as longi tudinal rib.The
species with this characters include A .pubescens ledeb and A .scoparia waldst.et.Kit.
The morphological and the micromorphological structures of achen s seed coat were listed
in Table 1.
3.Discussion
The experimental materials in this study w ere mainly obtained from the northeast of Chi-
na.Even though they accounted for only a small portion of this genus so that the representativ-
eness of the materials for the genus w as still limited to certain ex tent and such that the complete
deco ration of the seed coat of the achenes w as partially studied f rom the above point of view .
Nevertheless , it can still be show n that dif ferences in the achene s seed coat in Artemisia
plants from northeastern China exist between different groups of the genus based on the results
obtained here.The following comes from the comprehensive analysis of the results mostly f rom
an evolutionary view point.
3.1　The majori ty of the biocoenosia which w ere closely related in af finity relationship by con-
vent ional classification beared no ticeable similarity in terms of seed coat s decoration〔4〕.One
example of this is that in g roup of Sect.Absinthium DC.except A .f rigida willd , the rest of
118 植　　物　　研　　究　　　　　　　　　　　　　　　19 卷
4 species showed relaxed conjugation betw een the pericarp and the seed coat , and the ext reme
thickening of the pericarp , indicating the close genetic relationship of these species and their
relatively primitive types.However A.f rigida Willd w as much dif ferent f rom the other 4
species in the g roup in that a close conjugation betw een the seed coat and the pericarp , and less
intensive secondary thickening of the pericarp were observed.In addition , surface deco ration of
smooth type w as observed w ith SEM , show ing the mo re evolutionary properties.Moreover ,
the ratioes of leng th to w idth in g roup of Sect.Absinthium DC.were very close , varying f rom
1.90 to 1.99 , show ing the ed evolutionary feature.The 8 species from g roup of Sect.
Abrotanum Bess gave a nature of easy separation betw een the seed coat and the pericarp.This
g roup w ere primitive from revolutionary point of view , and large differences in microstructure
of seed coat existed among species of this group.For instance , A.Adamisii Bess had the net-
ted type of surface decoration , while A .Annua Jinn gave the longitudinal-rib type.Both A .
Lati fol ia Ledeb and A .Tanaceti folia Linn belonged to smooth type of the decoration.Also ,
the ratio of length to w idth varied to large deg ree betw een 1.60 to 2.50 , indicating the occur-
rance of ex tensive dif ferentiation in this group.
3.2　The analysis of the achene s mo rphological characters supported the point of view pro-
posed by Lin Youran that Sect.Artemisia , Sect.Dracunculus , Sect.lat ilobus Y.R.Ling
etc.were evoluted f rom Sect.Abrotanum Bess.Thirteen species f rom Sect.Artemisia stud-
ied here beared the longitudinal-rib type of the microst ructure on the surface of the f ruit.This
g roup , on the one hand , retained the primitive properties , i.e.the cross section of the f ruit
w as mostly round except A.Keiskeana Miq.On the o ther hand , they took the evolutionary
feature , i.e.the ratioes of the f ruit s length to w idth were all relatively large(more than 2.80
in most cases)and most species had tight conjugat ion between the pericarp and the seed coat
except A.keiskeana M iq.
3.3　Sect.Dravnclus Bess and Sect.Lat ilobus Ling lay at the advanced stage of evolution.
Both g roups gave the relatively small achenes wi th large ratio of length to w idth.In addition ,
the secondary thickening of the cell wall w as less intensive than that of the Sect.Abrotanum
Bess and Sect.Artemisia.The cell s surface w as smooth and only in the horizontal perines
formed the lateral lines and slight longitudianl-bulges.From the relativity of the characters ,
those w ith less intensive secondary thickening of the pericarp s cell w all had close conjugation
between the pericarp and the seed coat , and they had the high dif ferentiation of the cells , Based
on the characters described above , there w as no sifnificant difference betw een the tw o groups so
that it may be more proper fo r these tw o groups to be considered as one biocoenosium from the
viewpoint of the f rui t s structure.
Reference
1.Lin Lin , Lin S houquan.A preliminary study on the microstructural features of seed surfaces of Chinese Glycyrrhiza.Wuhan
Plant Research.1992 , 10(3):295～ 296
2.Yang Ji , Li Yiuhong , Wang Jinw u.A p rimary study on the microstructureal features of achene surfaces and i ts taxonomic
1191 期　　　　　　　　　　　　张大维:东北蒿属植物比较形态学研究
improtance to Polygonum , Acta Botanica Boreali-occideutalia S inica.1991 , 11(1):1～ 7
3.Huang Yuuping , Yin Zutang.The priliminary study on the micromorphlogical features of achene surfaces of Saussurea , Bul-
letin of Botanical Res , 1994 , 14(4):395～ 400
4.Ling Yeou-ruenn.On the system of the genus Artemisia Linn.and the relationship with it s allies.Bulletin of Botanical
Res.1982 , 2(2):1～ 60
5.Flora.RELPUBLICAE POPULARIS SINICAE.Delectis Florae Reipublicae Popularis S inicae Agende Academiae Sinicae
Edit.Tomus 76 Science Press 1991
Explanation of Plates
Fig.1～ 35.S canning elect ron microg raph sof seed coat svent ral 1.Ar temisie sieversiana Ehrhart ex W illd.2.A.anethi folia
Web.ex Stechm.3.A.zhaodongensis G.Y.Chang et.M.Y.Liou 4.A.anethoides Mat t f.5.A.f r igida Willd.6.A.
adamsii Bess.7.A.tanacet i fol ia Linn.8.A.la ti fol ia Ledeb.9.A.annua Linn.10.A.sacrorm um Ledeb.11.A.
sacrorm un Ledeb.var.messerschmid tiana (Bess)Y.R.Ling 12.A.palustri s Linn 13.A.aurata Komar.14.A.li t-
toricola Kitam .15.A.lavandu laefol ia DC.16.A.lancea Van 17.A.rubripes Nakai.18.A.igiaria Maxim.19.A.
argyi levl.et van.20.A.mongol ica(Fisch.ex Bess.)Nakai.21.A.integri fol ia Linn.22.A.subula ta Nakai.23.A.
selengensis Turcz.ex Bess.24.A.keiskeana Miq.25.A.sylvatica Maxim .26.A.viridisima(Komar.)Pamp.27.A.
d racuncu lus Linn.28.A.halodendron Turcz.ex Bess.29.A.oxycephala Kitag.30.A.pubescens Ledeb.31.A.sco-
paria Waldst.et Kit.32.A.deser torum Spreng.33.A.japonica Thunb.34.A.manshu rica(Komar.)Komar.35.A.
lit toricola Kitam .
120 植　　物　　研　　究　　　　　　　　　　　　　　　19 卷
张大维等:东北蒿属植物比较形态学研究Ⅰ .种皮结构 图版Ⅰ
Zhang Da-wei et al.:Comparative Study on Morphology of Artemisia Plants from Northeastern China Ⅰ .
Morphology of Seed Coat PlateⅠ
See explanation at the end of text
张大维等:图版Ⅱ Zhang Da-w ei et al.:PlateⅡ
See explanation at the end of text
张大维等:图版Ⅲ Zhang Da-w ei et al.:PlateⅢ
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