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Study on Floral Dynamic and Sexual Reproduction of Swertia mileensis, a Rare and Endemic Medicinal Plant

青叶胆开花动态及有性生殖特征的解剖学研究



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,2016,36(6):1146-1154
犃犮狋犪犅狅狋.犅狅狉犲犪犾.犗犮犮犻犱犲狀狋.犛犻狀.
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犛狋狌犱狔狅狀犉犾狅狉犪犾犇狔狀犪犿犻犮犪狀犱犛犲狓狌犪犾犚犲狆狉狅犱狌犮狋犻狅狀狅犳犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊,
犪犚犪狉犲犪狀犱犈狀犱犲犿犻犮犕犲犱犻犮犻狀犪犾犘犾犪狀狋
LILi1,LONGHua1,ZHANGAili2,HUANGHengyu2
(1KeyLaboratoryofPlantResourcesConservationandUtilizationofColegeofHunanProvince,JishouUniversity,Jishou,Hu
nan416000,China;2EngineeringResearchCenterforReproducingFineVarietiesofChineseMedicinalPlants,YunnanUniversi
tyofChineseTraditionalMedicine,Kunming650500,China)
犃犫狊狋狉犪犮狋:Thebloomingofthesingleflower,thedevelopmentofmicrosporangium,malegametophyte,me
gasporogenesis,femalegametophyteandembryogenyof犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊,aspecialherbendemictoYun
nan,werestudiedinthisarticle.(1)Thereproductivegrowthof犛.犿犻犾犲犲狀狊犻狊beginsatlateAugustorear
lySeptemberandthebudstageislastforapproximately35days,floweringtimeislastforonly2-3days
whilefruitstageisthelongest,approximately40-45days.(2)Thereareaseriesofmechanismtoguar
anteethecrosspolinationin犛.犿犻犾犲犲狀狊犻狊,suchastheversatileanther;thedichogamyinwhichthematu
rityofstamenwasabout23hoursearlierthanthematurityofpistil,thatmeanstheoverlappingtimeof
thematurestamenandpistilisonly1-2hours.Furthermoreanewmechanismwasobservedinwhichthe
stamenandpistilwouldchangetheirpositioninspacetoavoidselfpolination.Atfirstantherclingsto
stamenandthenturnaroundonthebackofthepistilaftera180°reversion,whilefilamentdeviateleadto
theantherbentdownwith210°.Sectionshows:anthersaretetrasporangiate.Thedevelopmentofanther
walsconformstothebasictypeandcomprisesofepidermis,endothecium,twomiddlelayersandtapetum
atthematurestage.Thetapetumcelshavedualoriginandbelongtotheglandulartype.Theantherloc
uleshavedegeneratingtapetumnucleiinthemiddlewhicharefromtheearlydividingandintrudingofthe
tapetum.Twomiddlelayersareephemeral;endotheciumreducesandepidermispersistsanddevelopsto
becomefibrousthickening.Thecytokinesisofthemicrosporemothercelinmeiosisisofthesimultaneous
type.Mostofthemicrosporetetradsaretetrahedralandpolengrainsare2celedand3celedwhenshed.
Theovaryisbicarpelate,unilocularwithsomeseriesofovuleslocatedattheparietalplacentation.Theo
vuleisunitegmic,enuinucelarandanatropous.Theonechalazalmegasporeinlineartetradbecomesthe
functionalmegasporeandthedevelopmentoffemalegametophyteisthepolygonumtype.Threeantipodal
celshavemultiseriatenucleianddistincthaustorium,andpersistlikealayerof“outerendosperm”justout
oftheendospermnucleuslayer.Fertilizationisporogamousandofpremitoticsyngamytype.Thedevelop
mentofendospermconformstonucleartypeandtheembryogenycorrespondstothesolanumtype.The
embryoisatthelateearlyheartshapedstagewhenseedsreleasedfromthecapsule.Thedevelopedantipo
dalcelsandsuspensorstructurehasmagnificentsignificanceforthereproductiveadaptationandevolution
oftheseedafterripeningof犛.犿犻犾犲犲狀狊犻狊.
犓犲狔狑狅狉犱狊:犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊;flowering;embryology;犛狑犲狉狋犻犪;ecologicaladaptation
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犘犾犪狋犲Ⅰ Thedurationofthesingleflower
A.Tetramerousflower;B.Inflorescenceemergence;CF.Thebudtime,whichlastsforabout20days;G.Thepositionofsta
mensinunopenedflower;H.Theflowerbudopens;I.Theanthersturnoutwards;J.Theantherpositionisverticaltopistiland
higherthanstigmafor60minutes;K.After30minutes,theantherkeeponturningoutwards90°tofaceagainstthepistil,thepolen
beginfalingwhilethestigmaisstilunreceptive;L.Thesheddingofpolenpeaksat23hoursafteritstartswhilethestigmaisstil
unreceptive;M.4hourslatertheanthersareinthelatestageofpolensheddingwhilethestigmaisreceptive;N.Thestigmaisopti
malforpolination2hourslater;O.Ovaryshapeafterpolination
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犘犾犪狋犲Ⅱ Microsporegenesisanddevelopmentofthemalegametophyte(ScaleBar:A =100μm;BE,J=20
μm;FI,KO=10μm)
A.Thetransectionofanther,showingfouranthers;B.Primaryparietalcels(PP)andprimarysporogenouscels(PS);C.An
therwalinthetimeofsecondarysporoginouscel:epidermis(EP),outlayer(OL)andinnerlayer(IL)withitsdivisionphase;D.
Antherwalinthetimeofmicrosporemothercels:epidermis(EP),endothelium(EN),middlelayer(M),tapetum(T);E.“Pla
centoid(PL)”fromthetapetum;F.MeiosismetaphaseⅠ;G.MeiosisanaphaseⅠ;H.MeiosismetaphaseⅡ;I.Meiosistelophase
Ⅱ;J.“Placentoid(PL)”fromthetapetuminthetimeoftetrads;K.Microsporehadjustformedfrommicrosporetetrads;L.Some
smalvacuolesappearinthemicrospore;M.Vacuolateperiodofuninucleatemicrospore;N.Early2celedpolens,showingobvious
celwal;O.Mature3celedpolens
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[1] c+%ec#$RSTUV!.c#$R(62»)[M].
"W

+%†X
,1988:392.
[2] YZ[,\Çc.wxy#$c¾^]-{ê^Gwxyê
_¾ŒH
[J].b/#$HS,1984,6(3):325328.
NIERL,HERY.TheErythrocentaurinandSwermirinfrom
犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊[J].犃犮狋犪犅狅狋犪狀犻犮犪犢狌狀狀犪狀犻犮犪,1984,6(3):
325328.
[3] `ab.ûy+[M]//c1d.ÔÞ\e? (g2f)
[M].»f:»f+%gh†X,1991:387402.
[4] mij,§k/.wxyT¤{|«tl~120z&m[J].
cfg
,1990,13(8):3738.
CHENGSM,ZHOUGN.犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊capsuleinthe
treatmentofacuteenteritis(areportof120cases)[J].犑狅狌狉
狀犪犾狅犳犆犺犻狀犲狊犲犕犲犱犻犮犻狀犪犾犕犪狋犲狉犻犪犾狊,1990,13(8):3738.
[5] no^,npq,Õ 3,d.wxy€‚ƒ[\DaHS
[J].cyºdfrR,2003,61:107109.
YANGYH,YANGLF,FANJ,犲狋犪犾.Strategyonpersis
tentutilizationof犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊[J].犆犺犻狀犲狊犲犑狅狌狉狀犪犾狅犳
犈狋犺狀狅犿犲犱犻犮犻狀犲犪狀犱犈狋犺狀狅狆犺犪狉犿犪犮狔,2003,61:107109.
[6] stp,7X.wxy{|«t€¬t}~422z|­Š‹
[J].b/cdrR,1981,3:3537.
DUGM,LIGY.犛狑犲狉狋犻犪犿犻犾犲犲狀狊犻狊inthetreatmentofacute
virushepatitis(ananalyticreportof422cases)[J].犢狌狀狀犪狀
35116š             7 8,d:wxy|{ËC“¦tBu1¾ÛÜ%HS
犑狅狌狉狀犪犾狅犳犜狉犪犱犻狋犻狅狀犪犾犆犺犻狀犲狊犲犕犲犱犻犮犻狀犲犪狀犱犕犪狋犲狉犻犪犕犲犱犻
犮犪,1981,3:3537.
[7] NOP,N u,vwx,d.wxyäÕ|€*+FŒ#2äB8$¶›HS
[J].c\f,2016,
47(3):480487.
HUANGHY,HUANGJ,WANGMR,犲狋犪犾.Comparison
onprogenyregenerationcapacityamongdifferentmatingpat
ternsandoptimizationofcultivationconditionin犛狑犲狉狋犻犪
犿犻犾犲犲狀狊犻狊[J].犆犺犻狀犲狊犲犜狉犪犱犻狋犻狅狀犪犾犪狀犱犎犲狉犫犪犾犇狉狌犵狊,2016,
47(3):480487.
[8] \}´,~3à.#$lR§÷HS¾¤Ã‡ãD€[J].
#$BC%&
,2003,27(2):151163.
HEYP,LIUJQ.Areviewonrecentadvancesinthestudies
ofplantbreedingsystem[J].犃犮狋犪犘犺狔狋狅犲犮狅犾狅犵犻犮犪犛犻狀犻犮犪,
2003,27(2):151163.
[9] ]b,‚ƒP,7„….†+、ه†+#$lR§÷DÜ
¯B$%¾HS‡ã
[J].#$Eë%&,2005,43(6):
574585.
GAOJY,RENPY,LIQJ.Advancesinthestudyofbreed
ingsystemandpolinationbiologyofgingers(Zingiberaceae
andCostaceae)[J].犃犮狋犪犘犺狔狋狅狋犪狓狅狀狅犿犻犮犪犛犻狀犻犮犪,2005,43
(6):574585.
[10] THORNERF.Proposednewrealignmentsintheangio
sperms[J].犖狅狉犱犻犮犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,1983,3:85117.
[11] THORNERF.Anupdatedphylogeneticclassificationof
flowingplants[J].犃犾犻狊狅,1992,13(2):365389.
[12] XUECY,HOTN,LIDZ.Embryologyof犛狑犲狉犻犪(Genti
anaceae)relativetotaxonomy[J].犅狅狋犪狀犻犮犪犾犑狅狌狉狀犪犾狅犳狋犺犲
犔犻狀狀犲犪狀犛狅犮犻犲狋狔,2007,155:383400.
[13] SHAMROVII.Theovuleof犛狑犲狉狋犻犪犻犫犲狉犻犮犪 (Gentianace
ae):structuralandfunctionalaspects[J].犘犺狔狋狅犿狅狉狆犺狅犾狅犵狔
,1991,41(3&4):213229.
[14] SHAMROVII.Ovuledevelopmentandsignificanceofits
featuresforGentianaceaesystematics[J].犗狆犲狉犪犅狅狋犪狀犻犮犪
犅犲犾犵犻犮犪,1996,7:113118.
[15] ~3à,ˆb‰,\Š‹.ŒfŽXYZ¾„…%HS[J].
!"ÕV%%&

)*+%†
),1988,34(4):5966.
LIUJQ,XUEC Y,HE T N.Embryologyof犛狑犲狉狋犻犪
犳狉犪狀犮犺犻狋犻犪狀犪,afamousTibetanmedicine[J].犑狅狌狉狀犪犾狅犳
犖狅狉狋犺狑犲狊狋犖狅狉犿犪犾犝狀犻狏犲狉狊犻狋狔 (NaturalScience),1988,34
(4):5966.
[16] \Š‹,ˆb‰,~3à.^3XYZ¾„…%[J].!"#
$%&
,1999,19(1):7680.
HET N,XUEC Y,LIUJQ.Embryologyof犛狑犲狉狋犻犪
犈狉狔狋犺狉狅狊狋犻犮狋犪 Maxim[J].犃犮狋犪犅狅狋犪狀犻犮犪犅狅狉犲犪犾犻犗犮犮犻犱犲狀狋犪
犾犻犪犛犻狀犻犮犪,1999,19(1):7680.
[17] ˆb‰,\Š‹,~3à.(XYZ¾„…%“­§÷%
VW
[J].#$Eë%&,1999,37(3):259263.
XUECY,HETN,LIUJQ.Embryologyof犛狑犲狉狋犻犪狋犲狋
狉犪狆狋犲狉犪 Maxim.(Gentianaceae)anditssystematicimplica
tion[J].犃犮狋犪犘犺狔狋狅狋犪狓狅狀狅犿犻犮犪犛犻狀犻犮犪,1999,37(3):
259263.
[18] ˆb‰,\Š‹,7‘.!/XYZ¾„…%“­§÷%
VW
[J].b/#$HS,2002,24(1):7581.
XUECY,HETN,LIDZ.Embryologyof犛狑犲狉狋犻犪犮犻狀犮狋犪
(Gentianaceae)anditssystematicvalue[J].犃犮狋犪犅狅狋犪狀犻犮犪
犢狌狀狀犪狀犻犮犪,2002,24(1):7581.
[19] 7 8,NOP.üýXYZ#$FQBG€F‚ƒ
[J].#$HS,2006,26(4):452460.
LIL,HUANGHY.Thegenesisofmicrosporeandthefor
mationofmalegametophytein犛狑犲狉狋犻犪犱犪狏犻犱犻犻Franch[J].
犅狌犾犾犲狋犻狀狅犳犅狅狋犪狀犻犮犪犾犚犲狊犲犪狉犮犺,2006,26(4):452460.
[20] NOP,û ”,1’’,d.XYZ¾„…QB[J].#$
HS
,2009,29(6):665673.
HUANGHY,LONGH,YITT,犲狋犪犾.Embryogenesisof
犛狑犲狉狋犻犪犫犻犿犪犮狌犾犪狋犪[J].犅狌犾犾犲狋犻狀狅犳犅狅狋犪狀犻犮犪犾犚犲狊犲犪狉犮犺,
2009,29(6):665673.
[21] NOP,û ”,1’’,d.XYZ#$FQBG€F
‚ƒ
[J].§!#$,2010,30(5):584593.
HUANGHY,LONGH,YITT,犲狋犪犾.Microsporogenesis
andthedevelopmentofmalegametophytein犛狑犲狉狋犻犪犫犻犿犪犮
狌犾犪狋犪[J].犌狌犻犺犪犻犪,2010,30(5):584593.
[22] DAVISGL.SystematicEmbryologyoftheAngiosperm [M].
NewYork:JohnWileyandSonsIIncorporated,1966:7071.
[23] SAMKARARK,NAGARAJM.StudiesinGentianaceae:
embryologyof犛狑犲狉狋犻犪犿犻狀狅狉(Gentianaceae)[J].犆犪狀犪犱犻犪狀
犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,1982,60:141151.
[24] 7 8,NOP.üýXYZ(ûy+)¾„…%HS[J].§
!#$
,2015,35(5):631640.
LIL,HUANGHY.Embryologyof犛狑犲狉狋犻犪犱犪狏犻犱犻犻(Gen
tianaceae)[J].犌狌犻犺犪犻犪,2015,35(5):631640.
[25] CAMERONBG,PTAKSH N.Occurrenceofgiantantipo
dalsinthefemaleGametophyteofAustralianBossiaceae,In
digofereaeand Mitbelieae(Leguminosae)[J].犃狌狊狋狉犪犾犻犪狀
犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,1990,38(4):395401.
[26] WILLEMSE M T,WENTJL.TheFemaleGametophyte
[M]//JohriB N.Embryologyofangiosperms.Berlin,
Heidelberg,NewYork:Springer,1984:8891.
[27] AKHALKATSM,WAGNERJ.Comparativeembryologyof
threeGentianaceaespeciesfromtheCaucasusandtheEuro
peanAlps[J].犘犾犪狀狋犛狔狊狋犲犿犪狋犻犮狊犪狀犱犈狏狅犾狌狋犻狅狀.1997,
204:2948.
[28] SCHWARTZBW,YEUNGEC,MEINKED W.Disrup
tionofmorphogenesisandtransformationofthesuspensorin
abnormalsuspensormutantsof犃狉犪犫犻犱狅狆狊犻狊[J].犇犲狏犲犾狅狆
犿犲狀狋,1994,120:32353245.
[29] SONGGQ,WANGXG,LIAQ,犲狋犪犾.Researchprogress
insuspensorofangiosperms[J].犆犺犻狀犲狊犲犅狌犾犾犲狋犻狀狅犳犅狅狋犪
狀狔,2012,47:188195.

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