Table Of Content云 南 植 物 研 究 , ():
暋2010 324 339~346
ActaBotanicaYunnanica : /
暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋DOI10灡3724SP灡J灡1143灡2010灡09252
蓟马诱导的鹅掌柴叶片虫瘿组织的生理生化响应
*
杨明挚,张汉波 ,李程晨,马双敏
**
云南大学生命科学学院植物科学研究所 云南 昆明
( , 暋650091)
摘要:以蓟马诱导的鹅掌柴叶片管状虫瘿为材料 分析了虫瘿和非虫瘿组织的抗氧化活性及其它生理响应
,
以及营养物质的含量 分配等 结果发现虫瘿组织中许多生理指标如过氧化氢浓度 过氧化物酶和苯丙氨
、 。 、
酸解氨酶活性以及丙二醛含量都比非虫瘿组织低 虫瘿组织比非虫瘿组织光合速率下降而呼吸速率增强
。 ,
虫瘿组织转变为一个营养库能够富集较多的营养物质 仅仅使用植物激素和蓟马萃取物不能诱导虫瘿的形
。
成 但是 细胞分裂素类植物生长调节物可以引起类似蓟马诱导虫瘿的早期反应
, , 。
关键词:虫瘿诱导 鹅掌柴叶片虫瘿 生理生化响应
, ;
中图分类号: 文献标识码: 文章编号:
Q945暋暋暋暋暋暋 A暋暋暋暋暋暋暋暋 0253灢2700(2010)04灢339灢08
PhysiologicalResponsesofGallTissuesonIvy
TreeLeavesInducedbyThrip
**
YANGMing灢Zhi,ZHANGHan灢Bo ,LIChen灢Cheng,MAShuang灢Min
(PlantScienceInstituteofLifeScienceSchool,YunnanUniversity,Kunming650091,China)
Abstract
:Inthisresearch,tuber灢likegallswhichinducedbythriponivytreeleaveswerechosenasmaterials,
andthephysiologicalresponsesaswellaspartitioningofmineralnutrition,solubleproteinandsaccharide
contentsingalledandun灢galledtissueswereassayed.Resultsindicatedthatcomparedtotheun灢galledtis灢
sues,gallingtissuesdecreasedinmanyphysiologicalindexesincludinghydrogenperoxideconcentration,Per灢
oxidaseandphenylnineamonialaseactivitiesandthecontentsofmalondialdehyde.Gallingtissuesdecreasedin
photosyntheticratebutincreasedinrespiratoryrate.Gallingtissuesactedasanutritivesinkthataccumulate
morenutritionthansurroundingun灢galledtissues.Phytohormonesandinsectsextractsalonecannotinduce
theformationofgalls,butcytokininssuchaskinetin,6灢BAandZeatincaninduceresponseswhicharesimilar
totheearlierresponseofthripfeeding.
Keywords
:Gallinginduction;Leafygall;Plantcellphysiologicalresponse
Introduction
havebeendemonstrated (Stoyanovaand Ham灢
Gallsarethespecialstructuresinducedbygall灢 burge,2003;GeurtsandBisseling,2002;Lim灢
ingmakersandcouldbeformedinplantleaves, pensetal灡,2003).Insectgallsarethemost
stemsandroots.Gallingmakersincludebacterial, commonphenomenonandmanyspeciesofin灢
fungi,mites,nematodesandinsects.Bacteria灢in灢 sectshaveprovedtobegallingmakers.Differ灢
ducedgallssuchascrowngallsandrootnodulesare entspeciesofgallinginsectsinducetheproduc灢
paidmuchattentionandtheformationmechanisms tionofdifferenttypesandmorphologicalgalls
* Foundationitems:DepartmentofScienceandTechnologyofYunnanProvince,China(No.2007PY01灢24),thenationalscience
foundationofChina(No.30560033,No.30960077)
** ZhangHan灢BoandYangMing灢Zhiareequallycontributedtothispaper
Receiveddate:2009灢12灢23,Accepteddate:2010灢04灢07
作者简介 杨明挚 男 博士 副教授 主要从事植物生理与分子生物学方面的研究
: (1970-) , , , 。E灢mail:mzh灢yang@163灡com
云 南 植 物 研 究 第 卷
3暋40暋暋暋暋暋暋暋暋暋暋暋 暋暋暋暋暋暋暋 暋 暋 暋 暋 暋 暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋 32
(Rohfritsch,1992;Stoneetal灡,2003;Dorchin opentuber灢likegallsarefrequentlyseenonwildivytree
etal灡,2002).Asplantpests,biologyandge灢 leavesfromspringtoautumn.Forstudythebehaviorof
neticsofgallinginsectswerereported(Abraha灢 thriponivytreeandthemorphogenesisofgalls,every20
gallandinsect灢freeivytrees(purchasedfromflowermar灢
msonetal灡,2003;Crespietal灡,1998;Fenton
ket)wereplantedandculturedintwodifferentmanualat灢
etal灡,2000;Bjorkman,2000;Fayetal灡,1996),
mosphereroom,respectively,bybranchescuttingmeth灢
andgallsareprovedtoaffectthereallocationof
odat25曟 with12h/12hlight/darkcycles.Ripethrip
nutrientsandgallsbecomevegetativesinkin insectsinivytreeleafgallswerecollectedfromcampusof
plants (Bagattoetal灡,1996;Rehilletal灡, YunnanUniversity.Inonemanualatmosphereroom,ev灢
2003;Florentineetal灡,2005;Koyamaetal灡, erythripwasputintooneivytreeononeyoungleaflet.
2002).However,littleistoknowaboutthefor灢 Mainstemsofeveryivytreewerecyclingpaintedwith
mationmechanism,thegallinginductionsub灢 vaselinetopreventinsectsfromescaping.Theactivitiesof
stances,andtheresponsesofplanttogalling thethripintheivytreewerecarefullyobservedandrecor灢
ded.After1,2,3,4,5days,thripsineverythreeivy
process.Leafygallsonthegardenplantivytree
treeweretakeout,respectively,andtheleftivytrees
werefrequentlyseenalongstreetinKunming,
wereletcontinuouslyco灢culturewiththrips.Gallingre灢
China.Gallingthripscaneasily,stablyand
sponseswereobservedandrecordedeveryday.
higherefficientlyinducetheproductionofgalls,
1灡2灡2暋GallinductionexperimentsbyPhytohormoneand
bymeansoffeeding.Thetuber灢likegallscanbe insectextract暋Foranalysisoftherolesofphytohormones
seenrapidlygrowwithinfewdays.Tostudythe ingallinduction,auxins(3mg·L灢1,10mg·L灢1of
inductionmechanismofgallformation,experi灢 IAA),cytokines(3mg·L灢1,10mg·L灢1ofKT,6灢BA,
mentsofgallinduction,physiologicalresponses orzetin),GA(10mg·L灢1GA3)andtheirmixtureswere
andnutritionallocationingalledandun灢galled usedtotreatyoungleavesofivytreesbymicro灢injuryin灢
jection.Insectextracts(extractedfrom50thripsbyphos灢
tissueswereassayedinthispaper.
phatebuffer)werealsousedinthisexperiment.Respon灢
Materialsandmethods seswereobservedandrecordedeveryday.
1暋 1灡2灡3暋Physiologicalresponsesassaytoplanttissues
Materials
1灡1暋 (1)Peroxidase(POX)activityassay.POXactivitywas
Ivytree(ScheffleraheptaphyllaHarms)andgalling assayedbyusingaspectrophotometricprocedure.20毺lofex灢
thrip(Phlaeothripidaespp)werechosenasgallingfor灢 tractwereaddedto1980毺lof88灡2mmol·L灢1 H2O2and
mationstudymodelinthisresearchformanyadvantages. 4灡6mmol·L灢1guaiacolin0灡1mol·L灢1KSuc,pH5灡5,and
1)Ivytreeisacommongardenplantsandcanrapidly vortexedfor5sec.Theincreaseinabsorbanceduetothe
propagatebyvegetablemethods;2)Tuber灢likegallsin灢 formationoftetraguaiacolwasmonitoredfrom0灡5minto
ducedbythripsonleavesbelongtotheopentypeofgall; 3灡5minat470nm.,andoneunit(U)ofPOXactivity
3)theactivityofthegallandgallmakerareeasilyob灢 wasdefinedas0灡1changeofabsorbanceperminute.
served;4)Gallformationisstableandefficient.Bothivy (2)Hydrogenperoxidecontentassay.Theconcen灢
treeplantsandgall灢madeinsect(thrips)werecollected trationofhydrogenperoxide(H2O2)wasdeterminedas
fromgardensandstreetsinKunming,China.Plantmate灢 describedbyMukherjeeandChoudhuri(Mukherjeeetal灡,
th
rialslistedbelowwerecollectedduringJuly21-25 in 1983).Briefly,freshleaves(0灡3g)werehomogenizedin
2007andusedintheexperiments:normalyoungleaves; 3mlofrefrigerated10%acetone,themixturewascentri灢
normalmatureleaves,normaloldleaves,younggalled fugedat10000暳gfor10min,and1mlsupernatantwas
leafgalledtissues,younggalledleafun灢galledtissues, mixedwith0灡1ml5% Ti(SO4)2and0灡2ml35%am灢
maturegalledleafgalledtissues,maturegalledleafun灢 monia.Aftertheprecipitatewasformed,thereaction
galledtissues,oldgalledleafgalledtissuesandoldgalled mixturewascentrifugedat10000暳gfor10min.There灢
leafun灢galledtissues.Eachsamplecollectedfromdiffer灢 sultingpelletwaswashedwithacetonethreetimesand
entsitehad8-12repeats. thendissolvedin2mol·L灢1H2SO4,andtheabsorbance
Methods
1灡2暋 wasrecordedat415nm.TheH2O2levelwascalculated
1灡2灡1暋Gallinductionexperiments暋Thripsandactive accordingtoastandardcurveofH2O2.Theconcentration
期
3 暋暋YANGMing灢Zhietal灡:PhysiologicalResponsesofGallTissuesonIvyTreeLeavesInducedbyThrip暋3暋41暋
ofH2O2wasexpressedasmmol·g灢1FW. contentswereexpressedas毺g/gormg/gaccordingtothe
(3)Phenylalnineammonialyase(PAL)detection. values.
ThePALassaywasconductedbyfollowingtheprocedure (2)Solvablesugardetection:Solublesugarwas
ofKoukolandConn(1961).About1gofleaftissueswas measuredasdescribedbyLehneretal灡(2006).Thecon灢
removedatrandom,andhomogenizedin10mlofcoldso灢 centrationofsolublesugarwasexpressedasmg·g灢1DW.
lutionof0灡05mol·L灢1sodiumboratebuffer,pH8灡5. (3)Solvableproteincontentanalysis:solubleprotein
Thehomogenatewasfilteredthroughcheeseclothand wasmeasuredineachsampleasdescribedbyBradford
centrifugedat14500暳gfor30min.Thecrudeextract (1976)andtheconcentrationwasexpressedasmg·g灢1
wasassayedimmediatelyafterhomogenization.ThePAL FW.
activitywasexpressedastheamount(毺gpergramtissue (4)C,H,Nelementsdetection:ThecontentsofC,
dryweight)ofcinnamicacidformed.Thereactionmix灢 H,andNindryingmaterialsweredetectedbyanorganic
tureconsistedof40mmolofL灢phenylalanine,200mmol elementsanalyzer(Elementar,zarioELIII,Germany)
ofsodiumboratebuffer(pH8灡8)andtheenzymesolu灢 accordingtothemanualsprotocol.
tioninatotalvolumeof5ml.About0灡1mlof6mol·
灢1
L HCIwasusedtostopthereaction.Theacidifiedmix灢 Results
2暋
turewasextractedwithethyletherandtheresiduedis灢
Gallinductionandformation
solvedin0灡05mol·L灢1sodiumhydroxideandabsorbance 2灡1暋
wasmeasuredat268nm. Onivytreeleaf,tuber灢likegallswerestably
(4)malondialdehyde(MDA)contentdetection:Ox灢 inducedbythrips,andthripsanditslarvaecan
idativedamagetolipidswasexpressedasequivalentsof befoundwithinanactivegall(Fig灡1:A,B).
malondialdehyde(MDA).MDAcontentwasdetermined Obviousresponses wereseenin1 dayafter
asdescribedbyDuanetal灡 (2005).Freshleavesortis灢 thrips曚“treatment暠.Theearlierresponsepro灢
sues(about0灡5g)werehomogenizedin10mlof10%
duceslightencoloredspots(about6mmdiame灢
trichloroaceticacid(TCA),andcentrifugedat12000暳g
ter)aroundtheinductionsites(Fig灡1:C,D).
for10min.Then,2ml0灡6%thiobarbituricacid(TBA)
Enationsusuallyappearedonleaves曚oppositeof
in10%TCAwasaddedtoanaliquotof2mlofthesuper灢
natant.Themixturewasheatedinboilingwaterfor30 thetreatedsiteswithintwodays.Onethripcan
min,andthenquicklycooledinanicebath.Aftercentrif灢 inducemanyleafygallswithinoneday曚s.Galls
ugationat1000暳gfor10min,theabsorbanceofthesu灢 cancontinuouslygrow andforminto mature
pernatantat450,532and600nmwasdeterminedwitha gallsevenwhentheinsectsweretakenawayaf灢
spectrometer(LibraS22,England).TheMDAconcen灢 terfewhourstreatment(Fig灡1:E,F,G).
trationwasestimatedusingtheformulaC(毺mol·L灢1)= 2灡2暋Phytohormonetreatmentresults
6灡45(A532灢A600)灢0灡56A450.TheMDAconcentration
Comparedtothecontrol,noobviousre灢
wasexpressedas毺mol·g灢1FW.
sponsewasinducedwhenIAAorGAwasused
(5)Photosyntheticandrespiratoryratemeasurement:
O2releasingratein500毺mol·m灢2redlightanddecreasing alone(Fig灡2:f,g).However,symbolssimilar
rateindarkconditionsweremeasuredbyaoxygenelectrode totheearlierresponse(lightencoloredspotsa灢
equipment(Oxy灢Lab,Hansatech,Egland)togalledand roundtheinductionsite)ofthripinductionwere
un灢galledleaftissuesaccordingtothemanualsprotocol. foundwhencytokinessuchasZeatin,6灢BAand
PhotosynthesisratewasexpressedbyO2increasingrated KTwereused(Fig灡2:b,c,d).Contrarytothe
inunitoftimeandunitofleafarea(毺mol·min灢1·m灢2). obviousboundarylightencoloredspotsinduced
AndrespiratoryratewasexpressedbyO2uptakeamount byinsects,lightencoloredspotsinducedbycy灢
inunitoftimeandunitmassoftissues.
tokinesdisplayedavagueboundary.Nogallwas
1灡2灡4暋Detectingnutrientcontent.
inducedorseenanygrowthofleaftissueswhen
(1)Mineralnutrientsanalysis:Fordetectionofthe
treatedbydifferenttypeofphytohormones,dif灢
mineralelementsofevytree,tissuesasdescribedinmate灢
rials(2灡1)werecollectedanddetectedbyaequipmentof ferenthormonecompositionaswellasinsects
ICP(ICP灢AES,ICPS灢1000II,Japan).Mineralelements extracts(Fig灡2).
云 南 植 物 研 究 第 卷
3暋42暋暋暋暋暋暋暋暋暋暋暋 暋暋暋暋暋暋暋 暋 暋 暋 暋 暋 暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋 32
Fig灡1暋Thripinducedtuber灢likeopengallsonivytreeleaves
A.Thripinducedtuber灢likeopengalls;B.Thripsanditslarvaecanfindinanaturallyproducedactivegalls;
C,D.lightencoloredspotsaroundtheinductionsiteswastheearlierresponseofgallinduction;E.obviously
enationsappearedwithintwodaysafterthrips曚treatment;F,G.onedaystayingofonethripcaninduce
morethan15gallsinonecomplexleaf;G.gallscangrowmaturelywithoutthecontinuouslyinduction
Fig灡2暋Phytohormonesandthripextractsinductionresults
a:Theearlierresponseoflifethripinduction;b,c,d:Responsesof6灢BA,zeatinandkatininistreatment,respectively;
e.f,g.Resultsofthripextracts,GAandIAAtreatment,respectively
Nutrientsallocationingalledandungelledtissues
2灡3暋 table1.Galledtissuescontainedhighernutri灢
Solvableprotein,soluablesugar,organic entscontentthanthatoftheun灢galledtissues,
elementsandinorganicelementsingalledand especiallythecontentofsolvableprotein,solva灢
un灢galledtissueswereanalyzedandillustratedin blesugar,totalNandtherecycledmineralele灢
期
3 暋暋YANGMing灢Zhietal灡:PhysiologicalResponsesofGallTissuesonIvyTreeLeavesInducedbyThrip暋3暋43暋
ments,suchasphosphate,potassium,Zincand
Boron.GalledtissuesdisplayedrelativelylowC/
Nratiocomparedtothatoftheun灢galledtis灢
sues.
Physiologicalmetabolismofgallingtissues
2灡4暋
Photosyntheticandrespiratoryrateofgal灢
ledandun灢galledtissuesweredetectedbyanox灢
ygenelectrodeinstrument.Asexpectednormal
matureleafhasmorephotosynthesisratethan
thoseofnormalyoungandolderleaves.Theac灢
Fig灡3暋Photosyntheticandrespiratoryrates
tivegalledtissuesalmosthavenotanyphotosyn灢
ofgalledandun灢galledtissues
theticabilitybutthetissuesaroundthegalled PR:photosynthesisrate;RR:respiratoryrate.A灢I:different
tissuesonthesameleafshowedhigherphoto灢 tissuesamplesasindicatedinthenotesoftable1
syntheticratethanthecorrespondingage曚snor灢
malleaf.Respiratoryratesdecreasedalongwith peroxidethanun灢galledtissues(Fig灡4:a).Per灢
theleafage.Therefore,galledtissueshave oxidaseactivityincreasesfrom youngtoold
higherintensityofrespiratoryratesthanitssur灢 leaves.However,onallkindsofleavesgalled
roundingtissuesandtheactivegalltissueshave tissuesalwaysdisplaylowerperoxidaseactivities
similarintensityofrespiratoryratesasthenor灢 thanun灢galledtissues(Fig灡4:b).Activitiesof
malyoungleaves(Fig灡3). phenylnineamonialaseandthecontentsofma灢
Stressphysiologicallyresponseofgallingtissues
2灡5暋 londialdehydedecreasedalongwiththephysio灢
Normalyoungandoldleaftissuescontain logicalageinnormalleaves.Thesametendency
morehydrogenperoxidethanthatofthemature wasalsofoundingalledandun灢galledtissuesas
leaf,asexpected.However,galledtissuescon灢 peroxidaseactivityandhydrogenperoxidecon灢
tainrelativelylow concentration ofhydrogen tent(Fig灡4:b,d).
Table1暋Contentofsolvableprotein,saccharide,organicandinorganicelementsingalledandun灢galledtissues
Tissues* A B C D E F G H I
Soluableprotein
4灡52暲0灡7 3灡16暲0灡5 3灡02暲0灡3 5灡87暲0灡5 5灡14暲0灡5 3灡79暲0灡4 2灡57暲0灡3 4灡72暲0灡4 4灡45暲0灡5
(mg/g)
Soluablesugar
111灡76暲9灡4 101灡19暲11灡5 94灡46暲8灡5 68灡80暲6灡7 47灡42暲7灡5 108灡74暲6灡4 93灡77暲7灡3 74灡57暲5灡9 52灡89暲5灡7
(mg/g)
TotalN(%) 3灡85暲0灡3 3灡14暲0灡3 2灡80暲0灡3 3灡87暲0灡3 3灡67暲0灡7 2灡93暲0灡2 2灡75暲0灡3 2灡61暲0灡2 2灡63暲0灡3
TotalC(%) 46灡37暲5灡5 48灡24暲4灡3 50灡51暲5灡7 46灡32暲5灡1 46灡00暲5灡0 48灡03暲4灡7 46灡4暲4灡3 45灡19暲4灡4 44灡43暲4灡2
C/Nratio 12灡03暲0灡9 15灡36暲1灡4 18灡05暲1灡5 11灡97暲1灡1 12灡54暲0灡9 16灡41暲1灡3 16灡85暲1灡2 17灡30暲1灡1 16灡91暲1灡6
Ca(mg/gdw) 5灡46暲0灡4 9灡95暲0灡7 12灡70暲0灡3 2灡87暲0灡1 4灡91暲0灡3 3灡59暲0灡4 10灡20暲0灡7 4灡49暲0灡3 9灡64暲0灡2
P(mg/g) 1灡51暲0灡1 0灡58暲0灡1 0灡65暲0灡1 1灡54暲0灡1 1灡08暲0灡1 1灡17暲0灡1 0灡70暲0灡1 0灡91暲0灡1 0灡65暲0灡1
K(毺g/g) 7灡01暲0灡5 7灡28暲0灡9 7灡77暲0灡6 9灡03暲0灡8 4灡33暲0灡7 10灡60暲0灡6 9灡90暲0灡5 9灡78暲0灡8 8灡53暲0灡7
Fe(毺g/g) 36灡48暲1灡5 52灡47暲2灡3 50灡17暲2灡6 72灡13暲2灡7 65灡08暲4灡5 89灡13暲4灡9 105灡00暲8灡3 111灡00暲8灡2 69灡70暲3灡9
S(毺g/g) 387暲19 372暲25 625暲22 318暲31 343暲21 268暲18 278暲13 230暲21 197暲20
Mg(mg/g) 2灡25暲0灡1 3灡75暲0灡1 1灡67暲0灡1 2灡39暲0灡1 1灡73暲0灡1 1灡66暲0灡3 2灡53暲0灡2 1灡66暲0灡2 1灡63暲0灡1
Mn(毺g/g) 75暲6灡3 146暲19 54暲7 88暲7 97暲6 78暲5 168暲9 50暲4 34暲6
B(毺g/g) 34暲2 27暲3 58暲3 24暲2 22暲3 21暲2 16暲1 15暲2 15暲1
Zn(mg/g) 9灡73暲0灡5 23灡22暲0灡7 12灡24暲2灡2 11灡00暲1 7灡27暲0灡5 13灡88暲0灡8 6灡44暲0灡7 14灡67暲0灡3 11灡53暲0灡7
“*暠Tissuesamplesnotes:A:normalyoungleaf;B:normalmatureleaf;C:normaloldleaf;D:younggalledleafgalledtissues;E:
younggalledleafun灢galledtissues;F:maturegalledleafgalledtissues;G:maturegalledleafun灢galledtissues;H:oldgalledleafgalled
tissues;I:oldgalledleafun灢galledtissues
云 南 植 物 研 究 第 卷
3暋44暋暋暋暋暋暋暋暋暋暋暋 暋暋暋暋暋暋暋 暋 暋 暋 暋 暋 暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋暋 32
Fig灡4暋Physiologicalresponsesofgalledandun灢galledtissuesofivytreeleaves
A灢I:differenttissuesamplesasindicatedinthenotesoftable1
Discussion
3暋 leadtothefindingsanddeeplyunderstandingof
Traditionally manygallmakinginsectshave plantendogenousregulatorsofIAA,cytokine
beenstudiedasplantpest,(Rohfritsch,1992; andGA,respectively.Whatmaytellusthatso
Abroletal灡,2006;Protasovetal灡,2007;Lvarez manytypesofgallsinducedbyinsects?
etal灡,2009),andresearchworksmainlyhave Convenientlypropagationofivytreeplant,
focusedongallmorphological(StoneandSchon灢 widelyseenofgallandgallingthrip,rapidly
rogge,2003;Arduinetal灡,2005),damageof gallingrespond,higherefficiencyofinduction
galltoplant(Abroletal灡,2006)andsoon. andeasilyobservationmakeopentuber灢likeleafy
Themorphologicalofgallsareconsideredtobe gallinducedbythripagoodmodelinplant灢in灢
determinedbythegeneticcharactersofgalling sectinteractionstudy.Inthiscase,wefocused
makers (Brantneretal灡,2000;Stoneand ontheplantresponsesduringgallinginduction
Schonrogge,2003;Arduinetal灡,2005).How灢 andformation.Asdescribedinothertypeof
ever,gallformationmechanismsstillremainun灢 galls(Bagattoetal灡,1996;RehillandSchultz,
clear.Gallsmaybeinducedbyinsectsormicro灢 2003;Florentineetal灡,2005;Koyamaetal灡,
organisms.Nodulesandcrowngallsarethefre灢 2002),gallingtissuesconcentratedmorenutri灢
quentlyseenmicrobe灢inducedgalls.Nodulesare entssuchassolvableproteins,solvablesaccha灢
inducedbynitrogen灢fixingRhizobium andthe ride,Nitrogen,phosphate,potassiumthansur灢
noduleinductionfactorisidentifiedandproved roundingun灢galledtissues(Table1).Gallinge灢
tobeoneoftheplantendogenoussubstances ventsleadedtothereallocationofnutrients.But
(GeurtsandBiseeling,2002;Limpensetal灡, theaveragecontentofmanynutrientswithin
2003).Studiesonthecrowngallleadtothe galledleaveswaslowerthanthatofthenormal
findingsofplantcytokinesandthebroadlyappli灢 correspondingstageleaves.Itimpliedthatnu灢
cationoftransgenictool(T灢DNA).Itiswell trientreallocationwasmostlyrestrictedwithin
knownthat,abnormalgrowthofplantsuchas thenearingtissuesandorgans.However,one
phototropism,crowngallandBanakaeDisease, questionishowplantcellsrespondtotheaction
期
3 暋暋YANGMing灢Zhietal灡:PhysiologicalResponsesofGallTissuesonIvyTreeLeavesInducedbyThrip暋3暋45暋
ofgallinginsects.Innormalconditions,when especiallytheplantcellgrowthanddivisionen灢
diseaseorpestthreatensaplant,plantcellsre灢 hancinghormones.Itisindirectevidencethat
spondasberesistantorsusceptible.Nomatter cytokinestypesofphytohormonesuchas6灢BA,
resistantorsusceptibleresponse,plantcellsof灢 KTandZeatincaninduceresponsessimilarto
tenrespondwithincreseofROs(reactiveoxi灢 theearlierinductionbythrips.Earlierreports
dates)stress,anti灢oxidaseactivities.MDAcon灢 demonstratedthatendogenousphytohormones
tentmayincreaseforoxidativedamagetolipids. weredetectedandfound17timesmoreauxinac灢
Inthisresearch,contentofhydrogenperoxide, tivityandasmuchas21timesmoregibberellin灢
MDAcontentsandtheactivitiesofPAL,POX likeactivityingallingtissuesthannormaltis灢
wereallseenobviouslydecreasedorinhibited. sues(Byersetal灡,1976).Cytokinesisalso
Alltheseindexesabovewereseenloweringall灢 foundtoincreaseinballgalls(MapesandDa灢
ingleavesthanthatofun灢galledornormalleav灢 vies,2001).So,phytohormones mayactto
es.Theseresultsmeansthatthegallingplant downstreamthissignalpathwayduringgallin灢
hasnotseeanyevidencestoinjuryorresistance. duction.Rootnodinducingfactorsasasignal
Wehypothesizedherethatthesubstances thatacceptedbycellularreceptorsthenmediate
whichgallinginsectsusedinplantgallinginduc灢 thenoduleformation hasbeen demonstrated
tionmaybeakindofplantcellendogenouscom灢 (Geurtsetal灡,2002;Limpensetal灡,2003).
ponents,andactinthefarupstreamofgallin灢 Geneswithinhostplanthavebeenprovedaffect灢
duction.Thecontentandproductionofthese ingtheinteractionofgallmidgeandrice(Limaa
substancesareunderstrictlycontrollinginnor灢 etal灡,2007).Fornutrientreallocation,anutri灢
malun灢galledplantcells.Exogenousintroduc灢 entshypothesisisprovidedtoexplaintheforma灢
tionofthesekindsofsubstancesleadstothe tionmechanismofgalls(Koyamaetal灡,2004).
changesbothinplantcellmetabolismandphyto灢 Butgallsbecomingamobilevegetablesinkmay
hormoneequilibrium,andconsequently,thegall alsobetheresponsetothehighercontentsof
isformed.Withthishypothesis,allofthere灢 phytohormones,because many ofthe plant
sultscanbereasonablyexplained.Becausethe growth enhancing hormones such as IAA,
CTK,GAandsoonareinvolvedinthenutrients
gallinginduction substances are endogenous
compoundsandplanttreatsthesemoleculesas partitioningregulation.
endogenoussignals,noresistantresponseisde灢 Multiplemorphologicalgallsareinducedby
differenttypeofgallinginsectsanddifferent
tected.Directevidencescomefromthemecha灢
gallinginsectsmayproducedifferenttypeofgall
nismofrootnoduleandcrowngallformation.
Rhizobiumnodfactor(akindoflow molecular inductionsubstanceandcreatingdifferentpat灢
polysaccharide)whichproducedbyrootnodule ternsofcellularequilibriumofplantregulators
duringthegallinductionprocess.So,identifica灢
bacterialhasbeenprovedasoneoftheplantcell
endogenous signal (Engleretal灡,1999). tion,functioningofthesetraceexistingsub灢
stancesmaydeeplyandwidelyimproveourun灢
Agrobacteriacandirectlytransformplantcyto灢
derstandingofplantcelldivisionandgrowth
kinesgeneintoplantcellandleadtotheforma灢
tionofcrowngalls(Lynnetal灡,1987).Itis regulatingmechanism.
notanewstorythatpests,pathogensorother Acknowledgements
:WeappreciateDr.ChenJin(Presi灢
plantsymbioticorganismownplantendogenous
dentofXishuangbannaTropicalBotanicalGarden,Chi灢
regulatorasitsbenefit.Ontheotherside,stud灢
neseAcademyofScience)theencouragementtopublic
iesontheinteractionbetweenplantanditsene灢
theseinitiallystudyresults.
miesalsodramaticallybroadenedourknowledge
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