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?. . . . . . . . . . . . 203 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 AlinaMosblech,IvoFeussner,andIngoHeilmann Contents xiii FattyAcidAmideHydrolaseandtheMetabolismof N-AcylethanolamineLipidMediatorsinPlants. . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 KentD. ChapmanandElisonB. Blanca?or SphingolipidSignalinginPlants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307 LouiseV. MichaelsonandJohnathanA. Napier Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 Contributors Steven A. Arisz Section Plant Physiology, Swammerdam Institute for Life Sciences,UniversityofAmsterdam,SciencePark904,NL-1098XH,Amsterdam, TheNetherlands ElisonB. Blanca?or SamuelRobertsNobleFoundation,PlantBiologyDivision, Ardmore,OK73401,USA,eblanca?or@noble.