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Leaf epidermal features of 21 medicinal plants of Euonymus in China

国产21种卫矛属药用植物叶表皮形态特征研究



全 文 :Leafepidermalfeaturesof21medicinalplantsofEuonymusinChina
ZHANGYifeng,HEPing,ZHANGChunping,HEJunxing
(KeyLaboratoryofEcoenvironmentsofThreeGorgesReservoirRegionMinistryofEducation,Schoolof
LifeScience,SouthwestUniversity,Chongqing400715,China)
[Abstract] Objective:ToaccumulatetaxonomicdataoftheleafepidermalfeaturesofthemedicinalspeciesofEuonymus.
Method:Twentninematerialsof21taxa(including16species,4varietiesand1form)representing5sectionsofEuonymusareex
aminedbyusingbothoflightmicroscopyandscanningelectronmicroscopy.Result:TheformofepidermalcelsinEuonymusisusual
lypolygonaloriregular.ThestomatawereanomocyticinalthespeciesexaminedexceptE.maackiandE.bungeanusvar.semiper
sistens.Stomataltypesofalspeciesstudiedmaybeanomocytic,anisocytic,cycolocyticorthetransitionaltypesamongthem.Conclu
sion:Theresultsshowthatsomecharacteristics(includingcuticularmembrane,shapeofguardcels,innermarginofouterstomatal
rim,outerstomatalrimandstomatatype)oftheleafepidermiscanprovidesomeanatomicalevidencefortheclassification.Thechar
acteristicsofleafepidermissupportfolowingtreatments:E.acanthocarpusvar.longipes,E.acanthocarpusvar.scandensandE.ac
anthocarpusvar.sutchuanensisshouldbemergedintoE.acanthocarpus;E.bungeanusvar.semipersistensshouldbemergedintoE.
maacki;E.hamiltonianusf.lanceifoliusshouldbemergedintoE.hamiltonianus.
[Keywords] Euonymus;medicinalplants;leafepidermis;stomatalapparatus;identification
[Accepteddate] 20080901
[Authorforcorrespondence]  HePing,Prof.Email:He
ping196373@126.com.
[Author] ZhangYifeng,Ph.D.,interestedintheresearchofconser
vationofmedicinalplants.Email:zhangyifeng_007@163.com.
  EuonymusL.isagenusofdeciduousorever
green,shrubsorshrublikeplants(ascendingorclimb
ingstemwithadventitiousrootsonothertreesorrocks
whenavailable)consistingof129speciesdistributedin
alovertheworld,primarilyinAsia[1],morethan100
speciesarefoundinChinaMedicaluses(especialy
intraditionalChinesemedicine)ofthespeciesfrom
EuonymushavealonghistoryThespeciesfromthis
genuscontainabundanttypesofmedicinalingredients,
suchassesquiterpenes[23],flavonoids[4],triterpenes
[5],steroids[6],cardiacglycoside[7],andsoon.
Pharmacologicalactionsofthoseconstituentsfrom
plantsofEuonymusincludingefectsofantitumor,an
tithrombotic,hypoglycemic,reducingbloodlipid,an
tiinflammatory,analgesia,antiAIDS,immunosup
pressionandinsecticidalactivity[8].
Scientificand systematicclassification isthe
premiseconditionofutilizationofwildmedicineplant
resources.Sometimes,itisverydificulttorecognize
speciesfromitsrelativespeciesaccuratelybasedon
grossmorphology.Misusedofmaterialsintraditional
Chinesemedicinemayleadtoseriousconsequences.It
isverynecessarytoaccumulatemicroscopictaxonomic
dataofmedicinalplantsfromthisgenus.Ithasbeen
repeatedlyconfirmedthatleafepidermalfeaturescan
helptoelucidatetaxonomicrelationshipsatdiferent
levels[912].However,theleafepidermisofEuonymus
hasbeenlitleexamined.Thepurposeofthisstudyis
tofindsignificanttaxonomiccharactersoftheleafepi
dermisforabeterunderstandingofthegenusandto
providesomereferencesforidentificationofmedicinal
plantsfromEuonymus.
1 MaterialandMethods
Matureleavesof21plantsfromChina,represen
tingalfivesectionswithinthegenus,areinvestigated
underlightmicroscope(LM)andscanningelectron
microscope(SEM).Alofthesamplesareobtained
fromthesouthwestuniversityofChina(SWU).Al
voucherspecimensaredepositedinSWU.Species
identifiers:ProfessorHePingandDengHongping
(southwestuniversityofChina).
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ThematerialsforLMstudyareboiledinwaterbe
forebeingmaceratedin20% NaClOsolution.Pieces
ofleafepidermisarestainedinasolutionof1% safra
nin(in50% alcohol)beforebeingmountedonmicro
scopeslides.Tochecktheconstancyofepidermal
structure,atleastfiveslidesaremadefromdiferent
partsofasingleleaforfromdiferentleavesofthe
samespecies.Thestomatalindexiscalculatedusing
formulaS/(E+S)100%.ThematerialsforSEM
oberservationarerinsedin75% alcoholsolutionbefore
beingsputercoatedwithgoldpaladiumusingaHita
chiE1010.ObservationsaremadeusingaHitachiS
30000Nscanningelectronmicroscopeat18kV.The
sectionalsystemofclassificationofCheng[13]isfol
lowed.TheterminologyfolowedthatofWilkinson[14]
andofDilcher[15].
2 Results
Thestomataexistedonabaxialepidermisinthe
speciesexaminedhere.Leafepidermalcelswereusu
alypolygonalinshapeofalspeciesexceptinE.ver
rucosoides.Thepaternsofanticlinalwalswereslightly
straight,repandorsinuate.Stomataltypesmaybe
anomocytic,anisocytic,cycolocyticorthetransitional
typesamongthem.Theoutlinesofthepairofguard
celsareusualynearlyroundedtowidelyelipticas
seeninsurfaceview,withlength/width(L/W)ratio
102156.Cuticularmembraneandouterstomatal
ledgearenealysmoothorerose.Outerstomatalrimare
raisedornot.Waxornamentation,ifpresent,isflake
likeorgranular.Thedetailcharacteristicsofleafepi
dermisinEuonymusunderLMandSEMaresumma
rizedinFig.1,Table1andTable2.
Table1 Species,localitiesandvouchersusedfortheleafepidermalstudiesinEuonymus
Taxa Locality Voucher
Sect.Ilicifolia Euonymustengyuehensis Lijiang,Yunnan W.C.Chen55360
  E.tengyuehensis Ganzhi,Sichuan F.W.Shi457825
  E.japonicus Beibei,Chongqing G.Q.He80001
  E.japonicus Mt.Emei,Sichuan P.Heetal.81456
  E.hederaceus Baoxing,Sichuan S.X.Guoetal.4372
Sect.Echinococcus E.angustatus Mt.Emei,Sichuan W.P.Fang7356
  E.angustatus Yuanjiang,Yunnan Q.RZhang07568
  E.wilsoni Wuxi,Chongqing S.X.Guoetal.6342
  E.chui Withoutpreciselocality S.X.Guoetal.00895
  E.acanthocarpus Mt.Emei,Sichuan S.X.Guoetal.570452
  E.acanthocarpusvar.longipes Nanchuang,Chongqing K.J.Shuetal.0660264
  E.acanthocarpusvar.scandens Lijiang,Yunnan P.Heetal.81659
  E.acanthocarpusvar.sutchuenensis Withoutpreciselocality W.P.Fang57896
Sect.Euonymus E.yunnanensis Withoutpreciselocality S.X.Guoetal.0026
  E.yunnanensis Lijiang,Yunnan W.P.Fang680047
  E.myrianthus Mt.Baita,Sichuan S.X.Guoetal.14563
  E.grandiflorus Mt.Emei,Sichuan S.X.Guoetal.570093
  E.maacki Yunnan S.X.Guoetal.84064
  E.bungeanusvar.semipersistens Lijiang,Yunnan W.P.Fang680126
  E.hamiltonianus Mt.Jinyun,Chongqing W.P.Fang0003
  E.hamiltonianusf.lanceifolius Qingyinge,Sichuan S.X.Guoetal.570064
Sect.Melanocarya E.verucosoides Mt.Emei,Sichuan S.X.Guoetal.570072
  E.verucosoides Guilin,Guangxi S.W.Wangetal.0125
  E.alatus Qingyinge,Sichuan S.X.Guoetal.570079
  E.alatus Yunnan Q.RZhang07524
Sect.Uniloculares E.schensianus Withoutpreciselocality Y.F.Zhang0543
  E.schensianus Daocheng,Sichuan S.X.Guoetal.2277
  E.giraldi Wenshan,Yunnan W.Wang426933
  E.giraldi Mt.Emei,Sichuan S.X.Guoetal.570236
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1.Euonymustengyuehensis.2.E.tengyuehensis.3.E.japonicus.4.E.japonicus.5.E.hederaceus.6.E.angustatus.7.E.angustatus.8.E.
wilsoni.9.E.chui.10.E.acanthocarpus.11.E.acanthocarpusvar.longipes.12.E.acanthocarpusvar.scandens.13.E.acanthocarpusvar.
sutchuanensis.14.E.yunnanensis.15.E.yunnanensis.16.E.myrianthus.17.E.grandiflorus.18.E.maacki.19.E.bungeanusvar.semi
persistens.20.E.hamiltonianus.21.E.hamiltonianusf.lanceifolius.22.E.verucosoides.23.E.verucosoides.24.E.alatus.25.E.alatus.
26.E.schensianus.27.E.schensianus.28.E.giraldi.29.E.giraldi.Scalebar=50μmunderLM,Scalebar=10μmunderSEM.
Fig.1 Adaxialepidermisof29speciesunderboththelightmicroscope(A)andscanningelectronmicroscope(B)
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Table2 CharactersofleafepidermisofEuonymusunderLMandSEM
Taxa
Cuticular
membrane
Shape
of
guard
cels
Inner
margin
ofouter
stomatal
rim
Outer
stomatal
rim
Wax
ornamentation
Stomata
type
Sizeof
stomata/μm
Stomatal
index
Sect.Ilicifolia Euonymustengyuehensis smooth nrerose raised flakeslike ano 2633×2409 176
  E.tengyuehensis smooth nrerose raised invisible ano 2568×2435 197
  E.japonicus smooth nrerose raised Invisible cyc 3815×3709 183
  E.japonicus smooth nrerose raised invisible cyc 3755×3652 167
  E.hederaceus smooth nrerose raised invisible ano 3128×2553 213
Sect.EchinococcusE.angustatus smooth nrerose notraised granular ano 2602×2493 218
  E.angustatus smooth nrerose notraised granular ano 2534×2476 158
  E.wilsoni striate nrerose notraised invisible ani 2336×2129 123
  E.chui smooth nrerose notraised granular ani 2723×2329 202
  E.acanthocarpus smooth nrerose notraised granular ani 2553×2319 145
  E.acanthocarpusvar.longipes smooth nrerose notraised granular ani 2633×2357 128
  E.acanthocarpusvar.scandens smooth nrerose notraised granular ani 2598×2467 147
  E.acanthocarpusvar.sutchuanensissmooth nrerose notraised granular ani 2497×2285 206
Sect.Euonymus E.yunnanensis smooth weerose notraised flakeslike ano 2838×2128 138
  E.yunnanensis smooth weerose notraised invisible ano 2925×2237 127
  E.myrianthus smooth nrerose notraised flakeslike ano 2954×2768 158
  E.grandiflorus smooth weerose notraised granular ano 3297×2234 150
  E.maacki striate wenearlysmoothraised invisible cyc 3023×2328 213
  E.bungeanusvar.semipersistens striate wenearlysmoothraised invisible cyc 2835×2267 189
  E.hamiltonianus smooth nrerose raised invisible cyc 3037×2897 186
  E.hamiltonianusf.lanceifolius smooth nrerose raised invisible cyc 2979×2788 178
Sect.MelanocaryaE.verucosoides smooth weerose raised flakeslike ani 3439×2537 108
  E.verucosoides smooth weerose raised invisible ani 3227×2248 158
  E.alatus smooth nrerose raised granular ano 2892×2649 129
  E.alatus smooth nrerose raised flakeslike ano 2787×2665 117
Sect.UnilocularesE.schensianus striate wenearlysmoothraised invisible ani 3342×2235 178
  E.schensianus Striate wenearlysmoothraised invisible ani 3955×2526 132
  E.giraldi striate weerose notraised invisible ani 4048×2789 152
  E.giraldi striate weerose notraised invisible ani 4364×2813 139
  Notice:wewidelyeliptic;nrnearlyrounded;anoanomocytic;anianisocytic;cyccycolocytic.
3 Discussion
Theexistenceortypesofwaxornamentationin
samesepeciesfromdiferentplacecouldbediferent
(Table2,suchasE.tengyuehensis,E.yunnanensis,
E.alatus).Thesizeofstomataandstomatalindex
couldchangebetweenthesamesepeciesfromdiferent
place(Table2,suchasE.alatus,E.yunnanensis,
E.giraldi).Waxornamentation,sizeofstomataand
stomatalindexarenotconstantwithinspecies,and
theymayberelatedtohabitat,butfurtherinvestigation
isrequired.Someotherepidermalfeatures(Including
cuticularmembrane,shapeofguardcels,innermar
ginofouterstomatalrim,outerstomatalrimandsto
matatype)inEuonymusappeartobeconstantwithin
species,andthuscanbeusedfordistinguishingsome
species.Theresultsofourstudyshowthatthecharac
teristicsoftheleafepidermiscanprovidesomeanatom
icalevidencefortheclassificationofEuonymus.There
areafewsubtleappearancediferencesbetweenE.
tengyuehensisandE.japonicus,soitisdificultto
identifytheformerfromthelaterbasedongrossmor
phology.Presentstudyrevealsthatsizeofstomatain
E.tengyuehensis(2633μm×2409μm)ismuch
smalerthanE.japonicus(3815μm×3709μm);
flakeslikewaxarefoundinE.tengyuehensisbutin
visibleinE.japonicus;anomocyticstomataisthe
mainlytypeofE.tengyuehensis,butcycolocyticstoma
tadominantinE.japonicus.Thus,itisveryeasyto
distinguishthemfromeachotherbasedontheleafepi
dermalfeatures.E.acanthocarpusvar.longipes,E.
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acanthocarpusvar.scandensandE.acanthocarpus
var.sutchuanensiswerereducedassynonymsofE.ac
anthocarpusbasedongrossmorphologyanalysis[1].
Presentstudyrevealsthatthecharacteristicsofleafepi
dermissupportthistreatment:cuticularmembrane,
shapeofguardcels,innermarginofouterstomatalrim
andouterstomatalrimofthosespeciesareverysimi
lar.Characteristicsofleafepidermisprovidedleafana
tomicalevidenceforthemergeofE.bungeanusvar.
semipersistensandE.maacki;ofE.hamiltonianus
andE.hamiltonianusf.lanceifolius.
[Acknowledgements] ThecuratoroftheherbariumofSouth
westUniversityforprovidingspecimens.Prof.Luolingfei,Mrs.
Hufei,Mrs.Majun,Mrs.Huangsizhou,Mrs.Lichun,Miss
YuanliliandMissPujiwenforthehelpduringtheprocessof
developingfilms.
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国产21种卫矛属药用植物叶表皮形态特征研究
张益锋,何平,张春平,何俊星
(1.西南大学 生命科学院 三峡库区生态环境教育部重点实验室,重庆 400715)
[摘要] 目的:为了积累卫矛属EuonymusL.药用植物叶表皮形态特征方面的分类学资料,以探讨和解决分类学中的一
些问题。方法:29份材料包括卫矛属5个组的21种植物(16种,4变种,1变型)叶表皮进行了光学显微镜(LM)和扫描电子显
微镜(SEM)下的比较研究。结果:表皮细胞一般呈不规则皱波状或者多裂深波状。气孔只存在于下表皮细胞(白杜和半常绿
白杜除外)。气孔类型有无规则型、不等型、环列型及其过度类型。结论:研究表明,一些叶表皮形态特征(角质膜、保卫细胞
形状、气孔外拱盖内缘、气孔外拱盖表面、气孔类型)包括能为卫矛属植物鉴定提供一些解剖学证据,支持如下分类处理:长梗
刺果卫矛、刺果藤杜仲、疏花刺果卫矛应并入刺果卫矛;半常绿白杜并入白杜;披针叶卫矛应并入西南卫矛。
[关键词] 卫矛属;药用植物;叶表皮;气孔;鉴定
[责任编辑 吕冬梅]
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