全 文 ：农业生物技术学报 JournalofAgriculturalBiotechnology 2006,14 (5): 736~741
*Foundationitem:TheNationalNaturalScienceFoundationofChina(No.30571192).
WUYing:bornin1978,woman,Ph.Dcandidate.
**Authorforcorrespondence.E-mail:.
Received:2005-12-15 Accepted:2006-03-07
·研究论文·
Ultraviolet-B Induced Chalcone Synthase (CHS) Gene
Expression and Its Signaling Pathways in Arabidopsis thaliana *
WUYing,LIANGYue-rong**,DONGJun-jie,LUJian-liang
(CollegeofAgricultureandBiotechnology,ZhejiangUniversity,Hangzhou310029,China)
Abstract:Transcriptsofchalcone synthase (CHS)gene in six linesofArabidopsis thaliana in response to UV-B radiation and
treatmentofhydroxylperoxide(H2O2),D/L-NG-mon methyl-arginine(D-NAME,L-NAME)and2-phenyl-4,4,4,4-tetramethylimidazo-
lin-1-oxyl-3-oxide (PTIO) were studied by RT-PCR method. Mutant Fah1-2 was the most sensitive and wild WS2 was the most
tolerant toUV-Bdamage inthesixlinesofArabidopsis.AccumulationoftranscriptsofCHSgenewasmarkedlyelevatedatsecond
hourbutdisappearedafter24hof15μmol/m2 sUV-B.H2OhadnoteffectonthetranscriptionofCHSgene.D-NAMEandL-NAME
suppressed the increase of CHS transcription in response to UV-B radiation while the effect of PTIO on CHS was very weak. It
appearsthatCHStranscriptioninresponsetoUV-Bradiationisnotregulatedbynitricoxide(NO)andH2O2.
Keywords:Arabidopsisthaliana;ultraviolet-B;chalconesynthase;geneexpression;signalingpathways
UV-B诱导拟南芥查耳酮合成酶 ( CHS) 基因表达及其信号传导途径
吴 颖,梁月荣**,董俊杰,陆建良
(浙江大学农业与生物技术学院,杭州310029)
摘要:应用 RT-PCR技术研究了 6个拟南芥(Arabidopsisthaliana)株系的查耳酮合成酶(CHS)基因转录对 UV-B照射以
及过氧化氢(H2O2)、D/L-NG 一甲基精氨酸(D-NAME、L-NAME)和 2-苯基 -l-4,4,4,4-四甲基咪唑啉 -1-烃氧基 -3-氧化物
(PTIO)等化合物处理的响应。结果表明,在6个株系中,Fah1-2突变株UV-B伤害最为敏感,而野生型WS2对UV-B伤害耐受
力最强;在 15μmol/m2 sUV-B照射 2h,CHS基因转录产物明显增加,但 24h后消失。H2O2对 CHS基因转录没有显著作用,
D-NAME和L-NAME能抑制CHS在UV-B照射下表达转录的增加,而 PTIO对 CHS基因表达的抑制作用很弱。据此认为,
CHS基因对UV-B照射响应的表达和转录不受NO和H2O2的调节。
关键词:拟南芥;UV-B;查耳酮合成酶;基因表达;信号传导途径
中图分类号:S188文献标识码:A 文章编号:1006-1304(2006)05-0736-06
Ultraviolet-B (UV-B,wavelengths280～320nm
of light) is harmful to living organisms and it is
increasingly concerned that the amount of UV-B
radiation reaching the Earths surface is increasing
because of depletion of the stratospheric ozone layer
(Frederick et al., 1989; Russel et al., 1996). The
harmfulness ofUV-B to the living organisms includes
its damage to DNA, proteins and cell membranes as
well as formation of reactive oxygen species(ROS)
such as superoxide and hydroxyl peroxide(Stapleton
andWalbot,1994;Landryetal.,1995). The excessive
UV-B compromises plant growth and its productivity
because UV-B radiation leads to reduction in
expression and synthesis of key photosynthetic
proteins in luding th chlorophyll a/b-binding protein
(Lhcb) and the D1 polypeptide of photosystem II
(psbA)(St pleton, 1992; Mackerness et al., 1997). In
responses to the UV-B radiation, however, plant
increases expression of genes encoding a number of
defense mechanisms such as enzymes involved in
synthesis of pro ective phenolic sunscreens and
pathogenesis-related proteins (Li et al., 1993; Surplus
第5期 WUYingetal:Ultraviolet-BInducedCHSGeneExpressionandItsSignalingPathwaysinArabidopsisthaliana
etal.,1998;BiezaandLois,2001).Thesignal
transductionmechanismsbywhichUV-Bradiation
regulatesgeneexpression arepoorlyunderstood
(Surplusetal.,1998).Flavonoidsareagroupof
importantsunscreensandchalconesynthase(CHS)is
thefirstenzymeintheflavonoid-specificbranchofthe
phenolpropanoidbiosynthesispathway (Dixonand
paiva,1995;WeisshaarandJenkins,1998;Jenkins,
1997;Jenkinsetal.,2001).RevealingtheUV-B
regulationoftranscriptionoftheCHSgenewilbe
helpfultolearnaboutthemechanismsthatprotect
plantsagainstUV-Binduceddamage.BiezaandLois
(2001)showedthatuvt1,amutantofArabidopsis
toleranttoiradiation,displayselevatedlevelsof
expressionofCHSmRNA andaccumulationof
flavonoidsandsinapatesbeforeexposuretoUV
thoughCHSmRNAlevelsaresignificantlyelevated
afterUV-Bexposure.Christieetal.(1996)showedthat
UV-Bphototransductionprocessesinvolvecalcium
amdcalmodulinwhereasUV-A/bluelightinductionof
CHS expression does not appear to involve
calmodulin.Fuglevandetal.(1996)confirmedthat
thereareseveraldistinctinductiveandsynergistic
UV-B, UV-A/blue phototransduction pathways
regulatingCHS genetranscriptionandtranscript
accumulationinArabidopsisleaftissueandonlythe
UV-A/blue lightinduction of CHS expression
involvedtheCRY1photoreceptor.UV-Bexposure
leadstothegenerationofreactiveoxygenspecies
(ROS)andnitricoxygen(NO),givingrisetoparalel
signalingpathwaysmediatingresponsesofspecific
genestoUV-Bradiation (Mackernessetal.,1997).
However,theUV-Binductiondoesnotappeartoin-
volveoxidativestresssignalingincelculturesystem
(Jenkins,1997;Jenkinsetal.,2001;Weisshaarand
Jenkins,1998),butwasreducedbyinhibitorsofnitric
oxidesynthase(NOS)orNOscavengers(Mackernesset
al.,1997).Thepresentstudywassettoinvestigatethe
efectofUV-BonthetranscriptionofCHSgeneinin-
tactplantsofvariousArabidopsismutantsandtherole
ofhydroxylperoxideandsomefreeradicalscavengers
intheUV-BinducedexpressionofCHSgeneinAra-
bidopsis.
1 MaterialsandMethods
1.1 Materials
ThreelinesofArabidopsisthalianawildtype
plantsandthreemutantsisolatedfromthecorespond-
ingwildtypeparentallineswereusedinthisstudy,i.e.,
aproteinkinasegeneknock-outmutantOX1isolated
fromArabidopsisthalianawildtypeWS2,theferulic
acidhydroxylasemutantFah1-2isolatedfrom wild
typeCol3andthetransparenttestaCHSmutantt4
fromwildtypeWTler.Seedsweretreatedat4℃ for
3daysaftersowingoncompost(Sinclairgrowerspot-
ting&beddingcompost,WiliamSinclairHorticulture
Ltd,FirthRoad,Lincoln,EnglandLN67AH)andthen
transferedtogrowthroomwithcontinuouslowwhite
light(fluencerate20μmol/m2/s)at19~22℃.On21st
dayaftertransferedtothegrowthroom,theplants
weregivenUV-Bradiation(fluencerate3.0μmol/m2/s
)fromtwoUV-B-313lamps(O-PanelLabCo.USA).
TheUVlightwasfilteredthrougha0.12-mm-thick
clearDiacetatefilm(Diacelproduct)andtheDiacetate
filmwaschangeddailytoachievedesiredUV-Bflu-
encerate.TheUV-Bfluencewasmonitoredbya
SR9910PCMacamspectroradiometre(MacamPhoto-
metrics,Livingston,Scotland).Thelightspectraofthe
UV-BwasshowninFig.1.Rosetes(100mg)fortotal
RNA extraction and reverse-transcription PCR
(rt-PCR)weresampledat2nd,4th,6thand24thhour
afterUV-Biradiation.Thesampleswerefrozenin
liquidnitrogenandstoredat-80℃tiltotalRNAex-
traction.
Fig.1.LightspectraofUV-B
1.2 TotalRNAextraction
TotalRNAwasextractedusingPURESCRIPTTM
R-5110AtotalRNAIsolationKit(Gentrasystems,
Minneapolis,MN55441,USA)andtheextractionpro-
cedurewascariedoutaccordingtotheproductin-
structions.TheRNAconcentrationwasdeterminedby
737
农 业 生 物 技 术 学 报 2006年
photospectrometricmethodusingBio-RadSmart-
specTM 3000spectrophotometer.ThetotalRNAob-
tainedwaspurifiedusingDNA-freeTMDNasetreatment
&RemovalReagents(AmbionInc.,2130Woodward
St.,Austin,TX78744,USA).
1.3Reversetranscription-polymerasechainreaction
(RT-PCR)
Reversetranscriptionwascariedoutin25μLre-
actionsolutionincluding5μL5×amplificationbufer;
1μL25mmol/LdNTPs,0.6μL10μmol/LOligod
(T),0.6μLRNaseinhibitor,1μLAMVreversetran-
scriptase,10μL0.25μg/μLtotalRNAand6.8μL
H2O.
Thereversetranscription reaction waspro-
grammedat48℃ for45min,95℃ for5minand
thenstoredat4℃.Thereactionproductswerethe
firststrandcDNAtobeusedforthefolowingPCR.
AmplificationofthefirststrandcDNAwasper-
formedusingcombinationoftwopairs(100bpse-
quenceCHS1and500bpsequenceCHS2)upper
primersandlowerprimersdesignedfromtheknown
CHSgeneofArabidopsisthaliana.Thereactionwas
cariedoutinafinalvolumeof25μL,including2.5
μL10×amplificationbufer,2.5μL25mmol/LMg++;
0.2μL25mmol/LdNTPs;2.0μL10mmol/Lupper
primer;2.0μL10mmollowerprimer;2.5μLabove
reversetranscriptionproduct;0.1 μL5U/μLTaq
DNApolymeraseand13.2μLH2O.
Thethermalcyclingwasprogrammedinthree
phasesinDYADTMPeltierThermalCyclermachine,i.
e.,initialdenaturation(at94℃for5min);25cyclesof
denaturation(at94℃for30s),annealing(at50℃for
30s)andextension(at72℃for1min);andfinalex-
tensionstepwas5minat72℃.ThePCRproductwas
checkedbyelectrophoresison2%(W/V)agarosegel.
1.4 Treatmentofhydrogenperoxide
Twenty-one-dayoldplantsofArabidopsiswild
typeWS2anditsmutantOX1grownundertheearlier
conditionsweresprayedwithhydrogenperoxide(H2O2,
0.9,9and90mmol/L)andH2O(control)untilrunof.
Thetreatedplantswereplacedinthesamegrowth
conditionsasbefore,i.e.continuouslowwhitelight
(fluencerate20μmol/m2/s)at19~22℃.
RosetesforRT-PCRweresampledatsecond,
fourth,sixthandtwenty-fourthhourafterthesprayand
RT-PCRwascariedoutasinsection1.3.
1.5 TreatmentofNOscavengers
Plantsoftheabove6linesofArabidopsisthaliana
(21-dayold)weresprayedwithD/L-NG-monomethyl-1
-argininesolution (L-NAME andD-NAME,100
mmol/L),2-phenyl-4,4,5,5-tetramethylimidazolin-1-
oxyl-3-oxidesolution(PTIO,10μmo/Ll)andH2O
(control)untilrunof.Thecompoundswereapplied1
hbeforethebeginningof3.0μmol/m2/sUV-Btreat-
ment.
RosetesfortotalRNAextractionandRT-PCR
weresampledat4thand24thhoursafterUV-Biradi-
ation.TheRT-PCRwascariedoutasearliermethods
describedinsection1.3.
1.6 Determinationofmorphologicaldamageandde-
hydrationindex(DI)
Seedsweretreatedat4℃for3daysaftersowing
ontheearlierdescribedcompostandthentransfered
togrowthroomwithcontinuoushighwhitelight(flu-
encerate200μmol/m2/s)at19~22℃.On14thday
aftertransferedtothegrowthroom,theplantswere
givenUV-Bradiation(fluencerate15μmol/m2/s)
fromsixUV-B-313lamps(O-PanelLabCo.USA)
supplementedwith40μmol/m2/swhitelightbySylva-
niaGRO-LuxF58Wlamps(GRO,Germany).Morpho-
logicaldamageoftheArabidopsisplantsiradiatedby
UV-Bwasobservedat6th,24thand72ndhoursafter
UV-Bradiation.
Plantsfordryweight(DW)measurementwerecut
ofabovetheroot,weighed,placedinglassdishesand
driedat45℃ for72hbeforere-weighing.Because
theinitialmoisturecontentsandmorphologicalchar-
actersofvariousgenotypesarediferent,whichafects
thedehydrationrateofvariousgenotypes,DIofthe
mutantswascalculatedbasedonmoisturecontentof
theirparentalplantssoastodiminishtheefectof
genotypediferenceandidentifytheefectofvarious
mutationsontheplantdehydrationduringUV-Bradia-
tion,i.e.,
DI(%)=(MMC1-MMC2)/MMC1
(PMC1-PMC2)/PMC1
×100
MMC1,mutantmoisturecontent(mg/g)beforeUV-B
iradiation;MMC2,mutantmoisturecontent(mg/g)af-
terUV-Biradiation;PMC1,parentallinemoisture
content(mg/g)beforeUV-B iradiation;PMC2,
parentallinemoisturecontent(mg/g)afterUV-Bira-
diation.
738
第5期 WUYingetal:Ultraviolet-BInducedCHSGeneExpressionandItsSignalingPathwaysinArabidopsisthaliana
2 Results
2.1TranscriptionofCHSgeneinducedbyUV-B
Accordingtothegeneexpressionprofilesdeter-
minedbyRT-PCR,therewasaveryweakexpression
oftheCHSgeneinthe6testedlinesofArabidopsis
thalianabeforeUV-Btreatment(Fig.2).TheCHStran-
scriptsmarkedlyincreasedwhentheplantswereex-
posedtoUV-Bfor2handthehighestaccumulationof
theCHStranscriptsappearedat4thhourexceptfor
OX1whichwasfoundtobeat6thh.At24thhourof
exposuretoUV-B,theCHStranscriptsreducedtothe
levelbeforeUV-Biradiation.Itsuggestedthattheex-
pressionofCHSgenewasinducedbyshorttermsof
UV-Biradiation.However,itsexpressionwouldbe
suppressedbylongtimeexposuretoUV-B.
Fig.2.EfectofUV-BradiationdurationonthetranscriptionofCHSgene
Plantsgrewunder20μmol/m2/sofwhitelightand19~22℃for21daysbeforeUV-Biradiation.UV-Bfluence:3.0μmol/m2/s.A.CHSgene
expressionprofilesofArabidopsisthalianalinesWS2anditsmutantOX1;B.CHSgeneexpressionprofilesofArabidopsisthalianalinesWTlerand
itsmutantt4;C.CHSgeneexpressionprofilesofArabidopsisthalianalinesCol3anditsmutantFah1-2;CHS1.RT-PCRresultswithfirstpairof
primerofCHSgene;CHS2.RT-PCRresultswithsecondpairofprimerofCHSgene;ACT,RT-PCRresultswithprimerofactinascontrol.
2.2 EfectsofH2O2andNOscavengersontranscrip-
tionofCHSgene
Fromthemorphologicalbehaviorsoftheplants,it
wasfoundthattheefectofH2O2ontheinjuryofthe
Arabidopsisthalianaplantsdependedonitsconcentra-
tion.Plantssprayedwith90mmol/LH2O2showedob-
viousmorphologicalinjuryontheleavesafter4hof
treatmentandtheplantswerefulywitheredafter24h.
Duringthe24haftertreatment,theplantssprayed
with0.9mmol/LH2O2lookedashealthasthecontrol
plantssprayedwithH2O,whiletheplantssprayedwith
9.0mmol/LH2O2wereslightlyinjured.TheRT-PCR
resultsshowedthattherewerenosignificantdifer-
encesintheprofilesofCHSgeneexpressionbetween
plantstreatedwithvariousconcentrationofH2O2and
wateriftheplantswerenotiradiatedbyUV-B(Fig.
3).ThissuggestedthatH2O2hadnoefectonthetran-
scriptionofCHSgene.
Plantsgrewunder20μmol/m2/sofwhitelight
and19~22℃for21daysbeforeH2O2treatment.After
H2O2 treatment,theplantswereplacedinthesame
growthconditionsasbefore.
Whentheplantsweresprayedwith(L-NAME,
100mmol/L),ascavengerofNO,anditsinactiveiso-
merD-NAME(100mmol/L)beforeUV-Bradiation,
theexpressionofCHSgenewasreduced,especialyon
lineOX1(Fig.4).However,PTIO(10μmol/L),anin-
hibitorofNOsynthase(NOS),hadaveryweaksup-
pressiveefectontheexpressionofCHSgeneinOX1
butonefectonWS2.
Fig.3.TranscriptionofCHSgeneunderstressofvariousconcentrationsofhydrogenperoxide
739
农 业 生 物 技 术 学 报 2006年
Fig.4.EfectofNOscavengerandNOSinhibitorontheCHS
geneexpressionArabidopsisthalianalinesWS2andOX1
A.CHSgeneexpressionprofilesafter3.0μmol/m2/sUV-Biradiation
for4h;B.TotalRNAusedfortheRT-PCRtest;M,DNAmolecular
marks;0,beforeUV-Biradiation;H2O,plantssprayedwithwateras
control;1,plantssprayedwith100mmol/LD-NAME;2,plantssprayed
with100mmol/LL-NAME;3,plantssprayedwith10μmol/LPTIO.
2.3 UV-Bsensitivityofvariousmutants
WhentheArabidopsisthalianaplantswereiradi-
atedwith15μmol/m2/sUV-Bfor6h,thediferences
inleafinjurywereobservedandtheirtoleranceto
UV-Bwasdiferentiatedwithiradiationtime.Ac-
cordingtothepercentageofinjuredleavesafter48h
UV-Biradiation(15μmol/m2/s),mutantFah1-2was
themostsensitivewhileWS2andOX1werethemost
toleranttotheUV-Biradiationamongthe6lines
(Table1).After72hiradiation,alleavesonthe6
lineswerewithered.However,recoveryofgrowthdif-
feredwhentheplantsweretransferedtogrowthroom
with200μmol/m2/swhitelightfor5d(Fig.5).WS2
andOX1recoveredgrowthmorequicklythanthatof
theotherlinesandtheyhadmoregreenleaves.The
Fah1-2lookedtheworstandt4thenextworst.
Fig.6showedthatDIwerethehighestonFah1-2
andthelowestonOX1,withthoseoft4inbetween.
ThissuggestedthatOX1dehydratedmoreslowly
duringUV-BiradiationthanthatofFah1-2andt4,
anditwasmoretoleranttoUV-B.Theresultswere
agreedwiththemorphologicalobservations(Fig.5).
3 Discussion
3.1Mechanismofplantself-defensefromUV-B
damage
ThepresentresultsshowedthatCHS gene
expressionwasinducedbyUV-Banditincreasedup
to6hofUV-Biradiation,butsuppressedafter24h.
TheCHSwasanimportantenzymeinvolvedinthe
phenolicsbiosynthesispathwayandthephenolicswere
protectivesunscreens(Lietal.,1993;DixonandPaiva,
1995;Surplusetal.,1998;BiezaandLois,2001).The
elevationofCHSgeneexpressionduringtheearly
stagesofUV-Biradiationmightbetheself-defense
mechanismofplantfromUV-Bdamage.However,the
plantswereseriousinjuredbylongtermsiradiationof
UV-B during which CHS geneexpression was
blocked.ThatexplainedwhytheCHSgeneexpression
wasnotobservedafter24hofUV-Biradiation.The
resultssuggested thatCHS mightbe a useful
molecularmarkerforbreedingcropcultivarsofUV-B
Table1Percentageofinjuredleavesofvariouslinesof
Arabidopsisthaliana*
*Theplantswereiradiatedwith15μmol/m2/sUV-Bfor48h.
Fig.5.RecoverygrowthstatusofArabidopsisthalianalines
afterUV-Biradiation
14-dayoldplantswereiradiatedwith15μmol/m2/sUV-Bfor72h
andthentransferedtogrowthroomunder200μmol/m2/swhitelight
for5d.
Fig.6.Dehydrationindex(D1)ofvariousmutants
Lines WS2 OX1 WTler t4 Col3 Fah1-2
Injuredleaves/% 40 45 50 60 45 80
740
第5期 WUYingetal:Ultraviolet-BInducedCHSGeneExpressionandItsSignalingPathwaysinArabidopsisthaliana
toleranceandplantswithhighconcentrationsof
phenolicsmightbetheresourcefordevelopmentof
anti-violetproducts.
Fig.2showedthattherewerenodiferencesinthe
CHSgeneexpressionprofilesbetweenthewildtypeof
Arabidopsisthalianalinesandtheirmutantsduringthe
UV-Btreatment.Theremightbetworeasonsonit.
Firstly,themutantsmightbeasinglepointmutationof
theCHSgeneandthiswasnotidentifiedbythe
RT-PCRtest.Forexample,transcriptionofCHSgene
inthemutantt4wasfoundnottobediferentiated
fromitsparentalplantWTler,suggestingthatitisa
singlepointmutationinsteadofafulCHSgene
knock-outmutant.Secondly,thediferentiationof
CHSgenetranscriptionmightbetakenplaceduring
theperiodbetween6hand24hofradiation,which
wasnomonitoredduringthepresentstudy.
3.2SignalingpathwayofCHSgeneexpression
ThepresentstudyconfirmedthatH2O2hadno
efectontheexpressionprofilesofCHSgene(Fig.3)
thoughtheplantsweredamagedbyhighconcentration
ofH2O2.ItsuggestedthattheregulationofCHSgene
transcriptiondidnotrequireH2O2.Thisagreedwiththe
discoverybyJenkinsetal.(2001)thattheinductionof
CHSgenetranscriptionbyUV-B wasnottobe
throughH2O2pathwaythoughup-regulationofFR-1
(pathogenesis-related-1)anddown-regulationofLhcb
(lightharvestingcomplexbindingproteins)genesby
UV-BradiationisregulatedbyH2O2(Mackernessetal.
,1997).
Fig.4showedthattheNOscavengerL-NAME
hadaweaksuppressionontheexpressionofCHS
gene,buttherewasnodiferencebetweenL-NAME
anditsinactiveisomerD-NAME.Also,PTIO,an
inhibitorofNOsynthase,hadnosignificantefecton
theexpressionofCHSgene.Thesesuggestedthatthe
efectofL-NAMEonCHSgeneexpressionwasnot
throughNOsignaling.ThesuppressofCHSgene
transcriptionbyD-NAMEandL-NAMEmightbe
someotherphototransductionpathwayinsteadofNO.
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