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. Distefano, M. LucianaLanteri, ArjentenHave, CarlosGarc?a-Mata, LorenzoLamattina, andAnaM. Laxalt 3-Phosphoinositide-DependentProteinKinaseisaSwitchboard fromSignalingLipidstoProteinPhosphorylationCascades. . . . . . . . . . . . . . 243 ChristineZalejskiandLaszloBo]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, OK