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Preliminary Exploration for Function of Cotton GhCDPK1 Gene under Drought Stress

陆地棉GhCDPK1基因响应干旱胁迫的功能初探



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,2016,36(8):1515-1521
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TIANXiaohan,ZHANGMengdan,PANGXuebing,ZHUJianbo,ZHUXinxia
(ColegeofLifeScience,ShiheziUniversity,Shihezi,Xinjiang832000,China)
犃犫狊狋狉犪犮狋:CDPKs(calciumdependentproteinkinases)areimportantcalciumsignalreceptorsandresponse
proteins,whichplayimportantrolesinresponsetovariousabioticstressesinplants,suchasdrought,low
temperatureandsalinity.Tostudytheeffectofcotton犌犺犆犇犘犓1genetodroughtstress,thisstudyused
quantitativerealtimePCRtoanalyzethegeneexpressionunderPEGstress.Theresultsshowedthat犌犺
犆犇犘犓1genewasupregulatedbydroughtstress.Moreover,plantexpressionvectorwasconstructedand
犌犺犆犇犘犓1genewasintroducedintotobaccoby犃犵狉狅犫犪犮狋狌狉犻狌犿mediatedleafdisktransformationmethod
toidentifythegenefunction.Theresultsshowedthatafterdroughtstress,thewaterretentioncapacityof
transgenicplantswassignificantlyhigherthanthatofwildtypeplants;thecontentsofchlorophyl,pro
line,solubleproteinandtheactivitiesofPODandSODintransgenictobaccoplantswerehigherthanthat
ofwildtype,whereasthecontentofMDAwaslowerthanthatofwildtypeplants.Theseresultsindicate
that犌犺犆犇犘犓1gene,asapositiveregulator,wasinducedbydroughtstress,andoverexpressionof犌犺
犆犇犘犓1genecanenhancetheabilityofplantstoresistdroughtstressbyaccumulatingmoreosmoticad
justmentsubstances,enhancingtheactivityoftheantioxidantsystemandmaintainingthestabilityofthe
celmembrane.
犓犲狔狑狅狉犱狊:cotton;犌犺犆犇犘犓1;droughtstress;expressionanalysis;transgenosis
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Function
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犌犺犆犇犘犓1qR GCCTTGCTCGCTTTCAG
78Ÿ a›
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犝犅犙7R GCTTGATCTTCTTGGGCTTG
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Referencegene
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Fig.1 Analysisof犌犺犆犇犘犓1geneexpression
underdroughtstress
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Line1andLine2.Twodifferentlinesoftransgenictobacco
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[1] LOBELLDB,SCHLENKERW,COSTAROBERTSJ.Cli
matetrendsandglobalcropproductionsince1980[J].犛犮犻
犲狀犮犲,2011,333(6042):616620.
[2] BOUDSOCQM,SHEENJ.CDPKsinimmuneandstresssig
naling[J].犜狉犲狀犱狊犻狀犘犾犪狀狋犛犮犻犲狀犮犲,2013,18(1):3040.
[3] BOUDSOCQ M,WILLMANN M R,MCCORMACK M,犲狋
0251 ! " # $ % &                   36á
al.DifferentialinnateimmunesignalingviaCa2+sensorpro
teinkinases[J].犖犪狋狌狉犲,2010,464(7287):418422.
[4] ®¯°.žno犃狋犆犇犘犓1Z=…*$†‡{|^ABAWx
ƒ„w±À
[D].²^³5:ôµef)%,2009.
[5] ZOUJJ,WEIFJ,WANGC,犲狋犪犾.犃狉犪犫犻犱狅狆狊犻狊calciumde
pendentproteinkinase犆犘犓10functionsinabscisicacidand
Ca2+mediatedstomatalregulationinresponsetodrought
stress[J].犘犾犪狀狋犘犺狔狊犻狅犾.,2010,154(3):12321243.
[6] ZOUJJ,LIXD,RATNASEKERAD,犲狋犪犾.犃狉犪犫犻犱狅狆狊犻狊
CALCIUMDEPENDENTPROTEIN KINASE8andCATA
LASE3functioninabscisicacidmediatedsignalingandH2O2
homeostasisinstomatalguardcelsunderdroughtstress[J].
犘犾犪狀狋犆犲犾犾,2015,27(5):14451460.
[7] ZHUSY,YUXC,WANGXJ,犲狋犪犾.Twocalciumdepend
entproteinkinases,犆犘犓4and犆犘犓11,regulateabscisicacid
signaltransductionin犃狉犪犫犻犱狅狆狊犻狊[J].犘犾犪狀狋犆犲犾犾,2007,19:
30193036.
[8] MASY,WU W H.犃狋犆犘犓23functionsin犃狉犪犫犻犱狅狆狊犻狊re
sponsestodroughtandsaltstresses[J].犘犾犪狀狋犕狅犾.犅犻狅犾.,
2007,65(4):511518.
[9] XUJ,TIANYS,PENGRH,犲狋犪犾.犃狋犆犘犓6,afunctionaly
redundantand positiveregulatorinvolvedin salt/drought
stresstolerancein犃狉犪犫犻犱狅狆狊犻狊[J].犘犾犪狀狋犪,2010,231(6):
12511260.
[10] WEIS,HU W,DENGX,犲狋犪犾.Aricecalciumdependent
proteinkinase犗狊犆犘犓9positivelyregulatesdroughtstress
toleranceandspikeletfertility[J].犅犕犆犘犾犪狀狋犅犻狅犾.,2014,
14(1):113.
[11] CAMPOS,BALDRICHP,MESSEGUERJ,犲狋犪犾.Overex
pressionofacalciumdependentproteinkinaseconferssalt
anddroughttoleranceinricebypreventingmembranelipid
peroxidation[J].犘犾犪狀狋犘犺狔狊犻狅犾.,2014,165(2):688704.
[12] YUXC,ZHUSY,GAOGF,犲狋犪犾.Expressionofagrape
calciumdependentprotein kinase 犃犆犘犓犾in 犃狉犪犫犻犱狅狆狊犻狊
狋犺犪犾犻犪狀犪promotesplantgrowthandconfersabscisicacidhy
persensitivityingermination,postgerminationgrowth,and
stomatalmovement[J].犘犾犪狀狋犕狅犾.犅犻狅犾.,2007,64(5):
531538.
[13] HUANGQS,WANG H Y,GAOP,犲狋犪犾.Cloningand
characterizationofacalciumdependentproteinkinasegene
associatedwithcottonfiberdevelopment[J].犘犾犪狀狋犆犲犾犾
犚犲狆.,2008,27(12):18691875.
[14] WANG H,MEIW,QIN Y,犲狋犪犾.1Aminocyclopropane
carboxylicacidsynthaseisphosphorylatedbycalciumde
pendentproteinkinase1duringcottonfiberelongation[J].
犃犮狋犪犅犻狅犮犺犻犿.犅犻狅狆犺狔狊.犛犻狀.(Shanghai),2011,43(8):654
661.
[15] ¶·¸,D ¹,º»¼,„._`‚†‡|ˆ78犌犺犆犘犓5
wÈÈäa›
[J].‰ef)%%&,2011,16(3):
814.
ZHENJB,ZHANGX,WANGYM,犲狋犪犾..Isolationand
sequenceanalysisofsaltresponsivegene犌犺犆犘犓5from犌狅狊
狊狔狆犻狌犿犺犻狊狌狋狌犿L[J].犑狅狌狉狀犪犾狅犳犆犺犻狀犪犃犵狉犻犮狌犾狋狌狉犪犾犝狀犻
狏犲狉狊犻狋狔,2011,16(3):814.
[16] DE1,½¾¿.#$*h%ijH¤[M]."¼:¼„ÀD
#ÁÂ
,2003:7073.
[17] ÃÄÄ,Å»¿,ÆÇÈ,„.#$ôS†‡Xpqrsw
;<—f
[J].!"#$%&,2012,32(5):10521061.
PEILL,GUOYH,XUZS,犲狋犪犾..Researchprogresson
Stresrelatedproteinkinasesinplants[J].犃犮狋犪犅狅狋.犅狅狉犲犪犾.
狅犮犮犻犱犲狀狋.犛犻狀.,2012,32(5):10521061..
[18] TAKAHASHIY,ITOT.StructureandFunctionofCDPK:
ASensorResponderofCalciumSignalingandCommunication
inPlants[M].2011,129146.
[19] SAIJOY,HATAS,KYOZUKAJ,犲狋犪犾.Overexpression
ofasingleCa2+dependentproteinkinaseconfersbothcold
andalt/droughttoleranceonriceplants[J].犘犾犪狀狋犑.,2000,
23(3):319327.
[20] ASANOT,HAYASHIN,KOBAYASHIM,犲狋犪犾.Arice
calciumdependentproteinkinase犗狊犆犘犓12oppositelymodu
latessaltstresstoleranceandblastdiseaseresistance[J].
犘犾犪狀狋犑.,2012,69(1):2636.
[21] ASANOT,HAYASHIN,KOBAYASHIiM,犲狋犪犾.Func
tionalcharacterisation of犗狊犆犘犓21,acalciumdependent
proteinkinasethatconferssalttoleranceinrice[J].犘犾犪狀狋
犕狅犾.犅犻狅犾.,2011,75(2):179191.

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