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GuardcellWikipedia,thefreeencyclopedia

Guardcell
FromWikipedia,thefreeencyclopedia

Guardcellsarespecializedcellsintheepidermisofleaves,
stemsandotherorgansthatareusedtocontrolgas
exchange.
Theguardcellsareproducedinpairswithagapbetween
themthatformsastomatalpore(Figure1).Thestomatal
poresarelargestwhenwaterisfreelyavailableandthe
guardcellsturgid,andclosedwhenwateravailabilityis
criticallylowandtheguardcellsbecomeflaccid(Figure1).
PhotosynthesisdependsonthediffusionofCarbondioxide
(CO2)fromtheairthroughthestomataintothemesophyll

OpeningandClosingofStoma.

tissues.Oxygen(O2),producedasabyproductof
photosynthesis,exitstheplantviathestomata.Whenthe
stomataareopen,waterislostbyevaporationandmustbe
replacedviathetranspirationstream,withwatertakenup
bytheroots.PlantsmustbalancetheamountofCO2
absorbedfromtheairwiththewaterlossthroughthe
stomatalpores,andthisisachievedbybothactiveand
passivecontrolofguardcellturgorandstomatalpore
size.[1][2][3][4]

Contents
1
2
3
4
5
6
7

Guardcellfunction
Waterlossandwateruseefficiency
Ionuptakeandrelease
Vacuolariontransport
Signaltransduction
Development
References

Figure1.Astomatalporeinthesurface(epidermis)
ofaleafasviewedthroughamicroscope.The
centralstomatalporeisformedbyapairofguard
cells.Thestomatalporecaneitheropen(left)or
close(right)dependingontheenvironmental
conditions.

Guardcellfunction
Openingandclosureofthestomatalpore(Figure1)ismediatedbychangesintheturgorpressureofthetwo
guardcells.Theturgorpressureofguardcellsiscontrolledbymovementsoflargequantitiesofionsandsugars
intoandoutoftheguardcells.Whenguardcellstakeupthesesolutes,thewaterpotential()insidethecells
increases(creatingahypertonicsolution),causingosmoticwaterflowintotheguardcells.Thisleadstoaturgor
pressureincreasecausingswellingoftheguardcellsandthestomatalporesopen(Figure2).Theionsthatare
takenupbyguardcellsaremainlypotassium(K+)ions[5][6][7]andchloride(Cl)ions.[8]Inadditionguardcells
takeupsugarsthatalsocontributetoopeningofthestomatalpores.

Waterlossandwateruseefficiency

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Waterstress(droughtandsaltstress)isoneofthemajorenvironmentalproblemscausingseverelossesin
agricultureandinnature.Droughttoleranceofplantsismediatedbyseveralmechanismsthatworktogether,
includingstabilizingandprotectingtheplantfromdamagecausedbydesiccationandalsocontrollinghow
muchwaterplantslosethroughthestomatalporesduringdrought.Aplanthormone,abscisicacid(ABA),is
producedinresponsetodrought.AmajortypeofABAreceptorhasbeenidentified.[9][10]Futureresearchis
neededtotestifthesereceptorscanbeusedtoengineerdroughttoleranceinplants.TheplanthormoneABA
causesthestomatalporestocloseinresponsetodrought,whichreducesplantwaterlossviatranspirationtothe
atmosphereandallowsplantstoavoidorslowdownwaterlossduringdroughts.Theuseofdroughttolerant
cropplantswouldleadtoareductionincroplossesduringdroughts.Sinceguardcellscontrolwaterlossof
plants,theinvestigationonhowstomatalopeningandclosureareregulatedcouldleadtothedevelopmentof
plantswithimprovedavoidanceorslowingofdesiccationandbetterwateruseefficiency.[1]

Ionuptakeandrelease
Ionuptakeintoguardcellscausesstomatalopening:The
openingofgasexchangeporesrequirestheuptakeof
potassiumionsintoguardcells.Potassiumchannelsand
pumpshavebeenidentifiedandshowntofunctioninthe
uptakeofionsandopeningofstomatalapertures(Figure
2).[7][11][12][13][14][15][16][17]Ionreleasefromguardcells
causesstomatalporeclosing:Otherionchannelshavebeen
Figure2.Ionchannelsandpumpsregulating
identifiedthatmediatereleaseofionsfromguardcells,
stomatalopeningandclosure.
whichresultsinosmoticwatereffluxfromguardcellsdue
toosmosis,shrinkingoftheguardcells,andclosingof
stomatalpores(Figures1and2).Specializedpotassiumeffluxchannelsparticipateinmediatingreleaseof
potassiumfromguardcells.[13][18][19][20][21]Anionchannelswereidentifiedasimportantcontrollersof
stomatalclosing.[22][23][24][25][26][27][28]Anionchannelshaveseveralmajorfunctionsincontrollingstomatal
closing:[23](a)Theyallowreleaseofanions,suchaschlorideandmalatefromguardcells,whichisneededfor
stomatalclosing.(b)Anionchannelsareactivatedbysignalsthatcausestomatalclosing,forexampleby
intracellularcalciumandABA.[23][26][29]Theresultingreleaseofnegativelychargedanionsfromguardcells
resultsinanelectricalshiftofthemembranetomorepositivevoltages(depolarization)attheintracellular
surfaceoftheguardcellplasmamembrane.Thiselectricaldepolarizationofguardcellsleadstoactivationof
theoutwardpotassiumchannelsandthereleaseofpotassiumthroughthesechannels(Figure2).Atleasttwo
majortypesofanionchannelshavebeencharacterizedintheplasmamembrane:StypeanionchannelsandR
typeanionchannels.[22][23][25][30]

Vacuolariontransport
Vacuolesarelargeintracellularstorageorganellesinplantscells.Inadditiontotheionchannelsintheplasma
membrane,vacuolarionchannelshaveimportantfunctionsinregulationofstomatalopeningandclosure
becausevacuolescanoccupyupto90%ofguardcellsvolume.Therefore,amajorityofionsarereleasedfrom
vacuoleswhenstomataareclosed.[31]VacuolarK+(VK)channelsandfastvacuolarchannelscanmediateK+
releasefromvacuoles.[32][33][34]VacuolarK+(VK)channelsareactivatedbyelevationintheintracellular
calciumconcentration.[32]Anothertypeofcalciumactivatedchannel,istheslowvacuolar(SV)channel.[35]SV
channelshavebeenshowntofunctionascationchannelsthatarepermeabletoCa2+ions,[32]buttheirexact
functionsarenotyetknowninplants.[36]

Signaltransduction
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Guardcellsperceiveandprocessenvironmentalandendogenousstimulisuchaslight,humidity,CO2
concentration,temperature,drought,andplanthormonestotriggercellularresponsesresultinginstomatal
openingorclosure.Thesesignaltransductionpathwaysdetermineforexamplehowquicklyaplantwilllose
waterduringadroughtperiod.Guardcellshavebecomeamodelforsinglecellsignaling.UsingArabidopsis
thaliana,theinvestigationofsignalprocessinginsingleguardcellshasbecomeopentothepowerof
genetics.[26]Cytosolicandnuclearproteinsandchemicalmessengersthatfunctioninstomatalmovementshave
beenidentifiedthatmediatethetransductionofenvironmentalsignalsthuscontrollingCO2intakeintoplants
andplantwaterloss.[1][2][3][4]Researchonguardcellsignaltransductionmechanismsisproducingan
understandingofhowplantscanimprovetheirresponsetodroughtstressbyreducingplantwaterloss.[1][37][38]
Guardcellsalsoprovideanexcellentmodelforbasicstudiesonhowacellintegratesnumerouskindsofinput
signalstoproducearesponse(stomatalopeningorclosing).Theseresponsesrequirecoordinationofnumerous
cellbiologicalprocessesinguardcells,includingsignalreception,ionchannelandpumpregulation,membrane
trafficking,transcription,cytoskeletalrearrangementsandmore.Achallengeforfutureresearchistoassignthe
functionsofsomeoftheidentifiedproteinstothesediversecellbiologicalprocesses.

Development
Duringthedevelopmentofplantleaves,thespecializedguardcellsdifferentiatefromguardmother
cells.[39][40]Thedensityofthestomatalporesinleavesisregulatedbyenvironmentalsignals,including
increasingatmosphericCO2concentration,whichreducesthedensityofstomatalporesinthesurfaceofleaves
inmanyplantspeciesbypresentlyunknownmechanisms.Thegeneticsofstomataldevelopmentcanbedirectly
studiedbyimagingoftheleafepidermisusingamicroscope(Figure1).Severalmajorcontrolproteinsthat
functioninapathwaymediatingthedevelopmentofguardcellsandthestomatalporeshavebeen
identified.[39][40]

References
1.SchroederJI,KwakJM,&AllenGJ(2001)Guard
cellabscisicacidsignallingandengineeringdrought
hardinessinplants.Nature410:327330.
2.HetheringtonAM&WoodwardFI(2003)Theroleof
stomatainsensinganddrivingenvironmentalchange.
Nature424:901908.
3.ShimazakiK,DoiM,AssmannSM,&KinoshitaT
(2007)Lightregulationofstomatalmovement.Annu
RevPlantBiol58:219247.
4.KwakJM,MserP,&SchroederJI(2008)The
clickableguardcell,versionII:Interactivemodelof
guardcellsignaltransductionmechanismsand
pathway.TheArabidopsisBook,edsLastR,Chang
C,GrahamI,LeyserO,McClungR,&WeinigC
(AmericanSocietyofPlantBiologists,Rockville),pp
117.
5.ImamuraS(1943)Untersuchungenuberden
mechanismusderturgorschwankungder
spaltoffnungsschliesszellen.Jap.J.Bot.12:251346.
6.HumbleGD&RaschkeK(1971)Stomatalopening
quantitativelyrelatedtopotassiumtransport.
Evidencefromelectronprobeanalysis.PlantPhysiol.
48:447453.
7.SchroederJI,HedrichR,&FernandezJM(1984)
Potassiumselectivesinglechannelsinguardcell
protoplastsofViciafaba.Nature312:361362.
8.MacRobbieEAC(1981)Ionfluxesin'isolated'guard
cellsofCommelinacommunisL.J.Exp.Bot.32:535
https://en.wikipedia.org/wiki/Guard_cell

cellsofCommelinacommunisL.J.Exp.Bot.32:535
562.
9.MaY,SzostkiewiczI,KorteA,MoesD,YangY,
ChristmannA,&GrillE(2009)RegulatorsofPP2C
phosphataseactivityfunctionasabscisicacidsensors.
Science324:10641068.
10.ParkSY,FungP,NishimuraN,JensenDR,FujiiH,
ZhaoY,LumbaS,SantiagoJ,RodriguesA,Chow
TF,AlfredSE,BonettaD,FinkelsteinR,ProvartNJ,
DesveauxD,RodriguezPL,McCourtP,ZhuJK,
SchroederJI,VolkmanBF,&CutlerSR(2009)
Abscisicacidinhibitstype2Cproteinphosphatases
viathePYR/PYLfamilyofSTARTproteins.Science
324:10681071.
11.AssmannSM,SimonciniL,&SchroederJI(1985)
Bluelightactivateselectrogenicionpumpingin
guardcellprotoplastsofViciafaba.Nature318:285
287.
12.ShimazakiK,IinoM,&ZeigerE(1986)Bluelight
dependentprotonextrusionbyguardcellprotoplasts
ofViciafaba.Nature319:324326.
13.SchroederJI,RaschkeK,&NeherE(1987)Voltage
dependenceofK+channelsinguardcellprotoplasts.
Proc.Natl.Acad.Sci.USA84:41084112.
14.BlattMR,ThielG,&TrenthamDR(1990)
ReversibleinactivationofK+channelsofVicia
stomatalguardcellsfollowingthephotolysisofcaged
1,4,5trisphosphate.Nature346:766769.
15.ThielG,MacRobbieEAC,&BlattMR(1992)
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15.ThielG,MacRobbieEAC,&BlattMR(1992)
Membranetransportinstomatalguardcells:The
importanceofvoltagecontrol.J.Memb.Biol.126:1
18.
16.KwakJM,MurataY,BaizabalAguirreVM,Merrill
J,WangM,KemperA,HawkeSD,TallmanG,&
SchroederJI(2001)DominantnegativeguardcellK+
channelmutantsreduceinwardrectifyingK+currents
andlightinducedstomatalopeninginArabidopsis.
PlantPhysiol.127:473485.
17.LebaudyA,VavasseurA,HosyE,DreyerI,
LeonhardtN,ThibaudJB,VeryAA,SimonneauT,&
SentenacH(2008)Plantadaptationtofluctuating
environmentandbiomassproductionarestrongly
dependentonguardcellpotassiumchannels.Proc.
Natl.Acad.Sci.USA105:52715276.
18.SchroederJI(1988)K+transportpropertiesofK+
channelsintheplasmamembraneofViciafabaguard
cells.J.Gen.Physiol.92:667683.
19.BlattMR&ArmstrongF(1993)K+channelsof
stomatalguardcells:Abscisicacidevokedcontrolof
theoutwardrectifiermediatedbycytoplasmicpH.
Planta191:330341.
20.AcheP,BeckerD,IvashikinaN,DietrichP,
RoelfsemaMR,&HedrichR(2000)GORK,a
delayedoutwardrectifierexpressedinguardcellsof
Arabidopsisthaliana,isaK+selective,K+sensing
ionchannel.FEBSLett486:9398.
21.HosyE,VavasseurA,MoulineK,DreyerI,Gaymard
F,PoreeF,BoucherezJ,LebaudyA,BouchezD,
VeryAA,SimonneauT,ThibaudJB,&SentenacH
(2003)TheArabidopsisoutwardK+channelGORK
isinvolvedinregulationofstomatalmovementsand
planttranspiration.ProcNatlAcadSciUSA
100:55495554.
22.KellerBU,HedrichR,&RaschkeK(1989)Voltage
dependentanionchannelsintheplasmamembraneof
guardcells.Nature341:450453.
23.SchroederJI&HagiwaraS(1989)Cytosoliccalcium
regulatesionchannelsintheplasmamembraneof
Viciafabaguardcells.Nature338:427430.
24.HedrichR,BuschH,&RaschkeK(1990)Ca2+and
nucleotidedependentregulationofvoltagedependent
anionchannelsintheplasmamembraneofguard
cells.EMBOJ.9:38893892.
25.SchroederJI&KellerBU(1992)Twotypesofanion
channelcurrentsinguardcellswithdistinctvoltage
regulation.Proc.Natl.Acad.Sci.USA89:50255029.
26.PeiZM,KuchitsuK,WardJM,SchwarzM,&
SchroederJI(1997)Differentialabscisicacid
regulationofguardcellslowanionchannelsin
Arabdiopsiswildtypeandabi1andabi2mutants.
PlantCell9:409423.
27.NegiJ,MatsudaO,NagasawaT,ObaY,TakahashiH,
KawaiYamadaM,UchimiyaH,HashimotoM,&Iba
K(2008)CO2regulatorSLAC1anditshomologues
areessentialforanionhomeostasisinplantcells.
Nature452:483486.
28.VahisaluT,KollistH,WangYF,NishimuraN,Chan
WY,ValerioG,LamminmakiA,BroscheM,Moldau
H,DesikanR,SchroederJI,&KangasjarviJ(2008)
SLAC1isrequiredforplantguardcellStypeanion
https://en.wikipedia.org/wiki/Guard_cell

SLAC1isrequiredforplantguardcellStypeanion
channelfunctioninstomatalsignalling.Nature
452:487491.
29.GrabovA,LeungJ,GiraudatJ,&BlattMR(1997)
Alterationofanionchannelkineticsinwildtypeand
abi11transgenicNicotianabenthamianaguardcells
byabscisicacid.PlantJ.12:203213.
30.LinderB&RaschkeK(1992)Aslowanionchannel
inguardcells,activationatlargehyperpolarization,
maybeprincipalforstomatalclosing.FEBSLett.
131:2730.
31.MacRobbieEAC(1998)Signaltransductionandion
channelsinguardcells.Phil.Trans.Roy.Soc.
London1374:14751488.
32.WardJM&SchroederJI(1994)Calciumactivated
K++channelsandcalciuminducedcalciumrelease
byslowvacuolarionchannelsinguardcellvacuoles
implicatedinthecontrolofstomatalclosure.Plant
Cell6:669683.
33.AllenGJ&SandersD(1996)Controlofionic
currentsguardcellvacuolesbycytosolicandluminal
calcium.PlantJ.10:10551069.
34.GobertA,IsayenkovS,VoelkerC,CzempinskiK,&
MaathuisFJ(2007)ThetwoporechannelTPK1gene
encodesthevacuolarK+conductanceandplaysarole
inK+homeostasis.ProcNatlAcadSciUSA
104:1072610731.
35.HedrichR&NeherE(1987)Cytoplasmiccalcium
regulatesvoltagedependentionchannelsinplant
vacuoles.Nature329:833836.
36.PeiterE,MaathuisFJ,MillsLN,KnightH,PellouxJ,
HetheringtonAM,&SandersD(2005)Thevacuolar
Ca2+activatedchannelTPC1regulatesgermination
andstomatalmovement.Nature434(7031):404408.
37.PeiZM,GhassemianM,KwakCM,McCourtP,&
SchroederJI(1998)Roleoffarnesyltransferasein
ABAregulationofguardcellanionchannelsand
plantwaterloss.Science282:287290.
38.WangY,YingJ,KuzmaM,ChalifouxM,SampleA,
McArthurC,UchaczT,SarvasC,WanJ,DennisDT,
McCourtP,&HuangY(2005)Moleculartailoringof
farnesylationforplantdroughttoleranceandyield
protection.PlantJ43:413424.
39.BergmannDC&SackFD(2007)Stomatal
development.AnnuRevPlantBiol58:163181.
40.PillitteriLJ&ToriiKU(2007)Breakingthesilence:
threebHLHproteinsdirectcellfatedecisionsduring
stomataldevelopment.Bioessays29:861870.

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