全 文 ：广西植物Guihaia28(2)：143—147 2008年3月
DistylyinForsythiasuspensa(Oleaceae)
RUANCheng—Jiang
’
(KeyLaboratoryofBiotechnology＆Bio-ResourcesUtilization．S缸据EthnicAffairsCommission
andMinistryofEducation，DalianNation litiesUniv rsity，Dalian116600，China)
Abst瞰t：DistylyprovidesprecisepollentransferwithouttheCOSTSthatareassociated、Il，ithsexualinterferenceand
self-pollination．InthirtypopulationsofdistyiousF r ythiasuspensa，style-morphratiosvariedfromisoplethy(1．00
long-morph：1．00short-morph，n=17opulations)toL-biased(1．14long-morph：1．00short-morph，n=13opula—
tions)．Underopenpollinationcondition，therewasnosignificantdifferenceinfruitsetbetweenlo g-(9．11％士
0．01％)andshort-(8．93％士O．06％)styledmorpha(F(x．39)=2．47，P=0．25)．Thelowerfertilizationsofco -
trolledintramorphpollinations(2．71％土O．02％inlong-longand2．58％土O．01％inshort-shortcombinations)in—
dicatedhatF．suspen$aisintramorph-incompatible．Therew snosignificantdiff renceinfruitsetofcross-pollina—
tionbetweenlo g-short-(36．410A士o 07％)andshort-long-(36．27％士O．03％)styledmorphscombinations
(Fo，39)=1．38，P----0．14)．Theresultsofartifidalouterossingin如tedtIlatpollinatorlimitationmayccurinthe
artificiaIpopulationsofF．suspezly硷innortheastofChina，thismaybebecsusethathelowtemperaturendmuch
windyinthearlyspringfluencepollinators’kindsandactivities．
Keywords：distyly；Forsythiasuspcrl$al floraItr it
(L(：Number：Q944．58DocumentCode：AArticleID：1000—3142(2008)02_0143—05
Heterostylyisusuallycomposedofa syn—
dromeoftraitsincludingfferencesinstyle ngth
andantherheight，asuiteofancillarypollena d
stigmapolymorphisms(Barretteta1．，2000)．This
sexualpolymorphismhasevolvedin ependentlyin
atleast28animal—pollinatedangiospermfamilies
(Webb&Lloyd，1986；Guitidnetal．，1998；Han＆
Kim，1999；Barretteta1．，2000；Arroyo&Barrett，
2000；P4rezetal．，2004；Massingaeta1．，2005)．In
heterostylousspecies，populationsarecomposedof
two(distyly)orthree(tristyly)floralmorphs(Bar—
retteta1．，2002)．MOstheterostylousspecieshave
strictintramorph-incompatibilitysystem(Massin-
gaeta1．，2005)，orsomehavestrorigbutnotstrict
intramorph—incompatibilitysys em(Jacquemynet
口Z．，2002)．Xueeta1．(2000)studiedthatthetime
andspacer lationso theglycoproteinforming
fromself-incompatibilitystyleofShatinyu(Citrus
grandisvar．shatinyu)．Distylousfrowersdiffer
reciprocallyinstigmaandantherposition，thelong—
andhort—styledmorphs(hereafterL—andS-mor—
phs)aresimilarinmorphologytotheapproachand
reverseherkogamousconditions(Barretteta1．，
2000)．Distylyprovidesprecisepollentransfer
withoutthecostshatareassociatedwithsexual
i terferenceandself_Dollination(CharlesworthD
&CharlesworthB，1979；Kohn＆Barrett，1992；
Paillereta1．，2002；Barretteta1．，2002)．Theper—
sistenceofdistylydependsontightlinkageofthe
genesaffectingantherheightandstigmaheight，
andasinglediallelicMendelianocuswithdomi—
nanceo trolsdistyly(Ganders，1979)．
Distylyhasbeenreportedinsomespeciesof
Oleaceae，suchasForsythia(Darwin，1877；Samp—
Receiveddate：2007—01—20Accepteddate：2007·08—09
Fotmdationit咖：SupportedbythChina-BelgianCooperationProject(200441505)
Biography：RUANCheng-Jiallg(1972’)，PKD，Professor，mainlyengagedinplantevolutionaryecology，molecular，systematic酬opment
andevolvement，andgeneticandbreeding，(E-mail)ruan@}dlnu．edu．cIL
万方数据
144 广西植物 28卷
son，1971；Han&Kim，1999)，Abeliophyllumdis— inceofChinain2005，and20populations(over500in-
tichum(Kangeta1．，2000)，Jasminum(Verdoorn，dividualsperp pulation)inXJ ya gCityofHenan
1963；0rnduff，1974)，andSchrebera(Verdoorn，
1963)．Darwin(1877)reportedthatth um(short-
styledmorph)anthersweresmallerthanpin(10ng-
styledmorph)anthersinForsythiasuspensa．The
alleled terminingthrumformis—dominantinFor-
sythia(Sampson，1971)．Flowercolorofshort—
styledflowerwasmoreyellowthanlong—styled
flowerinForsythiakoreanaNakai，andtheflower-
ingofshort--styledflowerwasearlierthanlong‘。
styledflower(Han&Kim，1999)．However，there
isnoreporton hestyle-morph(L—andS-morphs)
ratiosintheF．suspensepopulations，floraltrait
differencebetweentwomorphs，andfertilizationof
intramorphandintermorphpollination．Thepur—
posesofthepresentcontributionaret supply
someofthemissingformationonstyle-morphra—
tiosandfloraltraitsofF．suspensai thepopula—
tionofF．suspensei NortheastChina，investiga—
ringfruitsetunderopenpollinationcondition，and
examiningtheconsequencesofintramorph-。pollina·-
tionbetweenintramorphandCROSS—-pollinationbe··
tweeni termorph．Thisdatamayprovidescientificbas-
icforfurtherunderstandingtheistylyofF．suspema．
1 MaterialsandMethods
Forsythiasuspensa，aperenniallydedduous
shrub，isnativetothenorthandcentralofChina．Itis
cultivatedbroadlyinC11inaas nornamental．Thecap—
suleistipandegg-shaped，thenumb rofventriclesis
two，with4——10ovulesineveryventricle．Forsythia
suspensaflowersduringspring(fromMa chtoMay)，
andeachflowerlasts5—15days．Twostamenswith
equalheight(occasionallyppearsthreestamens)atta—
chestothebaseoftheflowertubefusedbythebaseof
quatrefoilc rolla．Stylewiththebilobedstigmaisu—
rtited．
Weconductedanextensiveurveytomeasurethe
relativefr quenciesofthetwostylemorphsinthirty
populationsofF． uspensa(over400individualsper
population)growinginDalianC tyofLiaoningProv-
Pr vi ceofCIlinain2006．InthePaotaipopulation，
thefollowingtreatmentswerecarriedout：(1)50flow—
ersofeachofmorphsweres lectedfrom50different
individualstomeasuresevenfloraltraits，including
flowerlengthandwidth，petallobewidthandlength，
corollatubelength，stigmaandantherheights；(2)Fruit
setsofeachoftwostyledmorphswereinvestigated(n
----50individualspermorph，andi00flowersperindi—
vidualwereselected，taggedandharvestedafter60
days)；(3)Ar ificialpollinationtreatmentswerede—
signed：oneisintramorph-pollination，includingL．L
(pollenfromtheotherflowersofthesameindividual，n
=20individuals)andS-S(pollenfromtheotherflow—
ersofthesaln．eindividual，n=20individuals)combina-
tions；anotheriscross-pollination．includingL_S(pol—
lenfromtheS-morphflowers)(n一20individuals)and
SL(pollenfromtheL-morphflowers)(n=20individ—
uals)combinations．Emasculationwasdonebeforethe
flowersfullyopen；thenthetreatedflowerswere
baggedusingylonbags．Hand-pollinationwascon—
ducted6-7daysaftertheflowersfullyopen．Ineach
treatment，100flowersperindividualwereselected，
tagged，treated，andharvestedafter60days．
2 Results
TheflowersofF．suspensahavefouryellowpet—
als，eachntherhastwoanther-petals，andeachanther-
petalhastwoclinandriums．Dehiscentmodeofclina —
driumislongitudinaldehiscence．Dehiscentorientation
ofantheristowardtopetals，whichmayavoidpollen
dispersingitsownstigmatopreventself-pollination．
L-morphflowersa eprotogyny，andS-morphflowers
areprotandry．
AllfiftypopulationsofF．suspensasampledare
dimorphicforstyle ngth．InDalianCity，style-morph
ratiosvariedfromisoplethy(1．00L： ．00S)(n一17
populations)toL-biased(1．14L： 00S)(n=13pop—
ulations)；andinXinyangCity，style-morphratiowas
1．00L：1．00S．IntheL-morph(Fig．1：A)，stignm
heightwas6．12±0．05Ⅱlm(n=50)。withtheanther
万方数据
2期 阮成江：连翘的二型花柱 145
heightof2．354-0．04rnrn(n一50)(Fig．2)；whereas
antherheightin heS-morph(Fig．1：B)was6．02±
nifieantlyexceededtheL-morphflowerinsizeofmost
corolladimensions，includingflowerlen th(S／L=
0·06ITIITI(n250)，withthestigmaheightof2．23-I-1．34)andwi th(S／L=1．25)。petallobewid h(S／L
0·04ITlln(n250)(Fig．2)．TheS-morphflowersig一 =1．17)andle gth(S／L=1．11)，andCOrollatube
E
E
75
65
55
45
35
15
Fig．1DistylyinForsythiasuspensa
A，long-styledmorphB，short-styledmorphC。short-styledmorphwithinflatedanthers；D，Iong．styled
morphwithinflatedanthers}1，stigma，2。anther；3，inflatedanther8
如 ．morph ▲
L- s‘m叼h每雌雄 ^‘渊‰▲·，▲ -▲酱
≮‰ 厶址．鱼．量衄嚆笋裂嚣洋
Fig．2Therangeofvariationinstigma(△)andanther
(▲)heightsinthelong-andshort-styledmorphs
amongindividualsfromthePaotaipopulation
growinginthenortheastofChina
length(S／L=1．08)．Underopenpollinationcon—
dition，fruitsetofL-morphwas9．11％±0．01％
(n=50)，rangingfrom2．33％tO11．24％，while
showed8．93％±O．06％(n=50)，rangingfrom
1．90％to11．69％intheS-morph．Thereisno
significantdifferenceinfruitsetbetweentwofloraI
morphs(F(1．39)=2．47，P=0．25)．Controlledpolli—
nationtreatmentsindicatedthatfruitsetsintheL-
LandS—Scombinationsare2．71％±0．02％(n=
20)and2．58％4-0．01％(n=20)，respectively：
whileshowed36．41％士0．07％(n一20)and
36．27％4-0．03％(n=20)intheL—SandS-Lcorn—
binations，respectively，andtherwasnosignificant
differenceinfruitsetsbetweentheL—SandS-L
combinations(F(1．3。)一1．38，P=O．14)．
3 Discussion
Forsythiasuspensaexhibitsthetypicalfloral
morphologicalsyndromeofdistylousspecies，the
L-morphflowerhasa relativelyhighstigmaand
lowanthers，whiletheS-morphflowerhashighan—
thersanda10wstigma．Thefloraltraitdifferences
betweentheS-andL-morphsintheF．suspen5口is
inagreementwithwhathasbeenobservedinthe
otherdistylousspecies(Ganders，1979；Riveroset
口Z．，1995；Li&Johnston，2001)．
Thelowerf uitsetunderopenpollinationin
thenaturalpopulationsmaybefromthelackof
pollenavailabilityandviability，pollinatoractivity，
andecologicalandphysiologicalfactorsrelatedo
pollinationmechanism(Kangeta1．，2000)．Polli—
natorsvi itflow rstocollectpollenor ectar，at
thesametimeplayinga importantroleaspollen—
vectors，andtheyinfluencefruitsetrates．The
fruitsetofartificialross—pollination(36．34％of
theaverageoftheL-Sand孓Lcombinations)inF．
suspensaisgreaterhanthatofopenpollination
(9．02％)，whichindicatesthatpollinatorlimitation
万方数据
146 广西植物 28卷
potentiallyoccursinthePaotaipopulation．Inthe
PaotaipopulationofF．suspensa，pollinationap—
pearstobeaffectedprimarilybyclimatefactors
duringthebloomingt me，suchastemperaturend
wind．Ifclimateconditionsarewindyandcloudy
withlowtemperature，largesizepollinators’activi—
tiesreduceandtheirvisitationratearealsolow．
Duringover5daysobservations，onlyfourspecies
ofinsectsarefoundedn aroronF．suspensa，inclu-
dingtwospeciesofbee，onespeciesofbeetle，and
onespeciesofspider．
Heterostylyhasevolvedtopreventself-pollination
andentrustpollinationbetweenindividualsofdifferent
types，andthereisaneffectiveincompatibilitysystemin
thedistylousspecies(B；arretet 1．，2000)．Omduff
(1966)suggestedthathe rostylousspeciesarein-
tramorph-incompatibleoriginally，butthedioecousevolu—
donmaybefromtheselectionopromoteutcrossing
becauseofthebreakdownofincompatibility．Theoccur-
renceofunusualincompatibilitysystemsindistylous
Villarsia(Ornduff，1988)andtristylousNarcissus(Sage
甜a1．。1999)indicatethathemorphologicalcompo—
nentsofheterostylyarenotalwaysas ociatedwithin-
tramorphmatingbarrierstypicalofspecieswithdiallel—
icincompatibility(Barrette a1．，2000)．Indeed，as
moreheterostylousspeciesarestudiedexperimentally
itisevidentthatmanyareself-compatible(Barrettet
a1．，2000)．The10w rintramorphfe tilizationofcon-
trolledintramorphpollinationsinF．$1Aspeflsaindicated
thathisspeciesisintramorph-incompatible，andthe
resultissimilarwiththeresultbyRyueta1．(1976)on
heterostyly-incompatibilityofA．d st chum(1．3％
fruitset)．
Kohn&Barrett(1992)reportedthatintheheter-
ostylousspecies，individualsoflong-styledmorphactas
femaleandshort-styledtypeasmale．Thefruitsetof
theS-morphflowerinthedistylousSarcothecacelebica
is10werthanthatoftheL-morphflower(Lack&Ke—
van，1987)．InsomeflowersofF．$lKspensa，anthersofL-
(Fig．1：D)andS(Fig．1：c)-morphsbothappearinflated
andpersistentlyu -dehiscentduringtheflowering，but
thefrequencyofflowerswithinflatedantherinS-morph
(2．84％4-0．01％，n=1000)isinorethanthatofL-mor-
ph(O．12％士O．01％，n=i00O)inthePaotaipopulation
(【，=O．000，W=237，PberelationwiththenaturalconditionsofthePaotain
population，butitisnecessarytofurtherstudythevo—
lutiona dadaptivesignificanceofi flatedantheroe-
cttrringinF．suspert．衄．
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2期 阮成江：连翘的二型花柱 147
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连翘的二型花柱
阮成江
(大连民族学院生物技术与资源利用国家民委教育部重点实验室，辽宁大连116600)
摘要：报道了连翘中的二型花柱(长花柱和短花柱)。长花柱的柱头和雄蕊高度分别为6．12士o．05ITllrn和
2．35士0．04rnnl，短花柱则为2．23士0．04rlLrn和6．02士0．06rnlTl。短花柱花的花冠大小明显超过长花柱。开放
授粉条件下，长短花柱花的座果率分别为9．11％士0．04％和8．93％士0．07％。人工异交的座果率在长一短
(36．8％士0．04％)与短一长(36．2％士0．07％)组合间无明显差异(F(1，39)一1．38，P一0．14)。人工异交实验表
明，传粉者限制可能发生在生长于中国东北部的人工连翘种群中，这可能是因为该地区早春的低温和多风气候
条件影响传粉者的种类和活动。
关键词：二型花柱；连翘；花性状
掌言，暑，三，；，；■i-i●Ii-鲁-善_星I毒罢，昌重善，言j●看，暑，i，暑，jIiIj■a奢i-鲁-昙●I毒l奢宅重量目●三，三j●ajj眚●i●叠暮-j重量署亭寻
(上接第272页Continuefrompag272)
需要进一步通过拟南芥p亚基和7亚基缺失突变体
细胞中提取蛋白质试验进行确证。
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万方数据
连翘的二型花柱
作者： 阮成江， RUAN Cheng-Jiang
作者单位： 大连民族学院,生物技术与资源利用国家民委教育部重点实验室,辽宁,大连,116600
刊名： 广西植物
英文刊名： GUIHAIA
年，卷(期)： 2008,28(2)
被引用次数： 1次

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