Lipid Signaling in Plants

Phospholipidshavelongbeenknownfortheirkeyroleinmaintainingthebilayer structureofmembranesandinphysicallyseparatingthecytosolfromorganelles andtheextracellularspace. Inthepastdecade,acompletelynovelandunexpected functionemerged,full?llingacrucialroleincellsignaling. Itwasthediscoveryin animalcells,thatagonist-activatedcellsurfacereceptorsledtotheactivationofa phospholipase C (PLC), to hydrolyze the minor lipid, phosphatidylinositol 4- bisphosphateintotwosecondmessengers,inositol1,4,5-trisphosphate(InsP)and 3 2+ diacylglycerol(DAG). WhileInsP diffusesintothecytosol,whereitreleasesCa 3 2+ from an intracellular store by activating a ligand-gated Ca -channel, DAG remainsinthemembranetorecruitandactivatemembersoftheproteinkinase Cfamily. Overtheyears,avarietyofotherlipidbased-signalingcascadesweredisc- ered. Theseinclude,phospholipaseA,generatinglyso-phospholipidsandfreefatty acids(tobeconvertedintoprostaglandinsandleukotrienes),phospholipaseD,to generatethelipidsecondmessenger,phosphatidicacid(PA),andphosphoinositide 3-kinase (PI3K), generating a distinct set of polyphosphoinositides (PPI) ph- phorylated at the D3-position of the inositol ring, all with separate signaling functions. Sphingolipids,representinganotherimportantgroupofsignalinglipids, alsocameacross. Themajorityoftheselipid-basedsignalingpathwayshavebeendiscoveredin plantcellstoo. Moreover,theyhavebeenfoundtobeactivatedinresponsetoa widevarietyofbioticandabioticstresssignals,butalsotobebasicallyinvolvedin plantgrowthanddevelopment. Whilemanyoftheenzymes,lipids,andtheirtargets involved arewell conserved, major differences with the mammalian paradigms havealsoemerged. Thisbookhighlightsthecurrentstatusofplantlipidsignaling. Allchaptershave beenwrittenbyexpertsinthe?eldandcoverinformationforbothbeginnersand advancedlipidologists. PartIincludesphospholipases(Chaps. 1-3),partII,lipid kinases (Chaps. 4-7), part III, lipid phosphatases (Chaps. 8-9), part IV, ix x Preface inositolphosphates and PPI metabolism (Chaps. 10-13), part V, PA signaling (Chaps. 14-17),andpartVI,additionallipidsignals,e. g. oxylipins,NAPEand sphingolipids(Chaps18-20). Ithasbeenagreatpleasuretobetheeditorofthis bookandtobeawitnessofthislipid-signalingadventure. Amsterdam,June2009 TeunMunnik Contents PartI Phospholipases PhospholipaseAinPlantSignalTransduction...3 Gu..ntherF. E. Scherer TheEmergingRolesofPhospholipaseCinPlantGrowth andDevelopment...23 PeterE. DowdandSimonGilroy PlantPhospholipaseD...39 WenhuaZhang,XiaoboWan,YueyunHong,WeiqiLi,andXueminWang PartII Kinases Phosphatidylinositol4-PhosphateisRequiredforTip GrowthinArabidopsisthaliana ...65 AmyL. SzumlanskiandErikNielsen PIP-KinasesasKeyRegulatorsofPlantFunction ...79 TillIschebeckandIngoHeilmann PlantPhosphatidylinositol3-Kinase...95 YureeLee,TeunMunnik,andYoungsookLee DiacylglycerolKinase...107 StevenA. AriszandTeunMunnik xi xii Contents PartIII Phosphatases SignalingandthePolyphosphoinositidePhosphatasesfromPlants ...117 GlendaE. Gillaspy PhosphatidicAcidPhosphatasesinSeedPlants...131 YukiNakamuraandHiroyukiOhta PartIV PPIMetabolism InsP inPlantCells ...145 3 YangJuIm,BrianQPhillippy,andImaraYPerera InositolPolyphosphatesandKinases...161 JillStevenson-PaulikandBrianQ. Phillippy PhosphoinositidesandPlantCellWallSynthesis ...175 RuiqinZhong,RyanL. McCarthy,andZheng-HuaYe ImagingLipidsinLivingPlants ...185 JoopE. M. VermeerandTeunMunnik PartV PASignaling PhosphatidicAcid:AnElectrostatic/Hydrogen-BondSwitch?...2 03 EdgarEduardKooijmanandChristaTesterink NitricOxideandPhosphatidicAcidSignalinginPlants...223 AyelenM. Diste'fano,M. LucianaLanteri,ArjentenHave, CarlosGarc?'a-Mata,LorenzoLamattina,andAnaM. Laxalt 3-Phosphoinositide-DependentProteinKinaseisaSwitchboard fromSignalingLipidstoProteinPhosphorylationCascades...243 ChristineZalejskiandLa'szlo'Bo..gre PartVI AdditionalLipidSignals DiacylglycerolPyrophosphate,ANovelPlantSignalingLipid...263 EmmanuelleJeannette,SophieParadis,andChristineZalejski OxylipinSignalingandPlantGrowth...277 AlinaM...