Phytohormones and Abiotic Stress Tolerance in Plants . Nafees A. Khan Rahat Nazar Noushina Iqbal l l Naser A. Anjum Editors Phytohormones and Abiotic Stress Tolerance in Plants Editors NafeesA.Khan NaserA.Anjum RahatNazar CentreforEnvironmentaland NoushinaIqbal MarineStud AligarhMuslimUniversity DepartmentofChemistry DepartmentofBotany Aveiro Aligarh Portugal India [email protected] [email protected] [email protected] [email protected] ISBN978-3-642-25828-2 e-ISBN978-3-642-25829-9 DOI10.1007/978-3-642-25829-9 SpringerHeidelbergDordrechtLondonNewYork LibraryofCongressControlNumber:2012933369 #Springer-VerlagBerlinHeidelberg2012 Thisworkissubjecttocopyright.Allrightsarereserved,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting, reproductiononmicrofilmsorinotherways,andstorageindatabanks.Duplicationofthispublicationor partsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9,1965,in itscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violationsareliableto prosecutionundertheGermanCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnot imply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotective lawsandregulationsandthereforefreeforgeneraluse. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface Plants are exposed to rapid and various unpredicted disturbances in the environ- ment resulting in stressful conditions. Abiotic stress is the negative impact of nonlivingfactorsonthelivingorganismsinaspecificenvironmentandconstitutes amajorlimitationtoagriculturalproduction.Theadverseenvironmentalconditions that plants encounter during their life cycle disturb metabolic reactions and ad- versely affect growth and development at cellular and whole plant level. Under abioticstress,plantsintegratemultipleexternalstresscuestobringaboutacoordi- nated response and establish mechanism to mitigate the stress by triggering a cascadeofeventsleadingtoenhancedtolerance.Responsestostressarecomplicat- ed integrated circuits involving multiple pathways and specific cellular compart- ments, and the interaction of additional cofactors and/or signaling molecules coordinatesaspecifiedresponsetoagivenstimulus.Stresssignalisfirstperceived bythereceptorspresentonthemembraneoftheplantcells.Thesignalinformation isthentransduceddownstreamresultingintheactivationofvariousstress-responsive genes.Theproductsofthesestressgenesultimatelyleadtostresstoleranceresponse or plant adaptation and help the plant to survive and surpass the unfavorable conditions. Abiotic stress conditions lead to production of signaling molecule(s) thatinducethesynthesisofseveralmetabolites,includingphytohormonesforstress tolerance.Phytohormonesarechemicalcompoundsproducedinonepartandexert effect in another part and influence physiological and biochemical processes. Phytohormonesarecriticalforplantgrowthanddevelopmentandplayanimportant role in integrating various stress signals and controlling downstream stress responsesandinteractincoordinationwitheachotherfordefensesignalnetwork- ingtofine-tunedefense.Theadaptiveprocessofplantsresponseimposedbyabiotic stresses such as salt, cold, drought, and wounding is mainly controlled by the phytohormones. Stress conditions activate phytohormones signaling pathways that are thought to mediate adaptive responses at extremely low concentration. Thus,anunderstandingofthephytohormoneshomeostasisandsignalingisessen- tialforimprovingplantperformanceunderoptimalandstressfulenvironments. v vi Preface Traditionallyfivemajorclassesofplanthormoneshavebeenrecognized:auxins, cytokinins, gibberellins, abscisic acid, and ethylene. Recently, other signaling molecules that play roles in plant metabolism and abiotic stress tolerance have also been identified, including brassinosteroids, jasmonic acid, salicylic acid, and nitric oxide. Besides, more active molecules are being found and new families of regulatorsareemergingsuchaspolyamines,plantpeptides,andkarrikins.Several biologicaleffects ofphytohormones areinducedbycooperation ofmorethanone phytohormone. Substantial progress has been made in understanding individual aspects of phytohormones perception, signal transduction, homeostasis, or influ- enceongeneexpression.However,thephysiological,biochemical,andmolecular mechanisms induced by phytohormones through which plants integrate adaptive responsesunderabioticstressarelargelyunknown.Thisbookupdatesthecurrent knowledge on the role of phytohormones in the control of plant growth and development, explores the mechanism responsible for the perception and signal transduction of phytohormones, and also provides a further understanding of the complexityofsignalcrosstalkandcontrollingdownstreamstressresponses.There is next to none any book that provides update information on the phytohormones significanceintolerancetoabioticstressinplants. Weextendourgratitudetoallthosewhohavecontributedinmakingthisbook possible.Simultaneously,wewouldliketoapologizeunreservedlyforanymistakes orfailuretoacknowledgefully. Aligarh,India NafeesA.Khan,RahatNazar,NoushinaIqbal Aveiro,Portugal NaserA.Anjum Contents 1 SignalTransductionofPhytohormonesUnderAbioticStresses ....... 1 F.Eyidogan,M.T.Oz,M.Yucel,andH.A.Oktem 2 Cross-TalkBetweenPhytohormoneSignalingPathwaysUnder BothOptimalandStressfulEnvironmentalConditions ............... 49 MarciaA.Harrison 3 PhytohormonesinSalinityTolerance:EthyleneandGibberellins CrossTalk ................................................................. 77 NoushinaIqbal,AsimMasood,andNafeesA.Khan 4 FunctionofNitricOxideUnderEnvironmentalStressConditions ... 99 MarinaLeterrier,RaquelValderrama,MouniraChaki, MorakAiraki,Jose´ M.Palma,JuanB.Barroso,andFranciscoJ.Corpas 5 AuxinasPartoftheWoundingResponseinPlants ................... 115 ClaudiaA.Casalongue´,DiegoF.Fiol,RamiroPar´ıs, AndreaV.Godoy,Sebastia´nD‘Ippo´lito,andMar´ıaC.Terrile 6 HowDoLettuceSeedlingsAdapttoLow-pHStressConditions? AMechanismforLow-pH-InducedRootHairFormation inLettuceSeedlings ...................................................... 125 HidenoriTakahashi 7 CytokininMetabolism ................................................... 157 SomyaDwivedi-Burks 8 OriginofBrassinosteroidsandTheirRoleinOxidative StressinPlants ........................................................... 169 AndrzejBajguz vii viii Contents 9 HormonalIntermediatesintheProtectiveActionofExogenous PhytohormonesinWheatPlantsUnderSalinity ..................... 185 FaridaM.Shakirova,AzamatM.Avalbaev,MarinaV.Bezrukova, RimmaA.Fatkhutdinova,DilaraR.Maslennikova,RuslanA.Yuldashev, ChulpanR.Allagulova,andOksanaV.Lastochkina 10 TheRoleofPhytohormonesintheControlofPlantAdaptation toOxygenDepletion .................................................... 229 VladislavV.YemelyanovandMariaF.Shishova 11 StressHormoneLevelsAssociatedwithDroughtTolerancevs. SensitivityinSunflower(HelianthusannuusL.) ..................... 249 CristianFerna´ndez,SergioAlemano,AnaVigliocco, AndreaAndrade,andGuillerminaAbdala 12 AnInsightintotheRoleofSalicylicAcidandJasmonic AcidinSaltStressTolerance .......................................... 277 M.IqbalR.Khan,ShabinaSyeed,RahatNazar,andNaserA.Anjum Index .......................................................................... 301 Chapter 1 Signal Transduction of Phytohormones Under Abiotic Stresses F.Eyidogan,M.T.Oz,M.Yucel,andH.A.Oktem Abstract Growth and productivity of higher plants are adversely affected by various environmental stresses which are of two main types, biotic and abiotic, dependingonthesourceofstress.Broadrangeofabioticstressesincludesosmotic stresscausedbydrought,salinity,highorlowtemperatures,freezing,orflooding, aswellasionic,nutrient,ormetalstresses,andotherscausedbymechanicalfactors, light, or radiation. Plants contrary to animals cannot escape from these environ- mental constraints, and over the course of evolution, they have developed some physiological,biochemical,ormolecularmechanismstoovercomeeffectsofstress. Phytohormones such as auxin, cytokinin, abscisic acid, jasmonic acid, ethylene, salicylic acid, gibberellic acid, and few others, besides their functions during germination, growth, development, and flowering, play key roles and coordinate various signal transduction pathways in plants during responses to environmental stresses. Complex networks of gene regulation by these phytohormones under abioticstressesinvolvevariouscis-ortrans-actingelements.Someofthetranscrip- tionfactorsregulatedbyphytohormonesincludeARF,AREB/ABF,DREB,MYC/ MYB, NAC,andothers. Changes ingene expression, proteinsynthesis,modifica- tion, ordegradation initiatedby or coupled tothese transcription factors andtheir correspondingcis-actingelementsarebrieflysummarizedinthiswork.Moreover, crosstalk between signal transduction pathways involving phytohormones is explained in regard to transcriptional or translational regulation under abiotic stresses. F.Eyidogan(*) BaskentUniversity,Ankara,Turkey e-mail:[email protected] M.T.Oz•M.Yucel•H.A.Oktem DepartmentofBiologicalSciences,MiddleEastTechnicalUniversity,Ankara,Turkey N.A.Khanetal.(eds.),PhytohormonesandAbioticStressToleranceinPlants, 1 DOI10.1007/978-3-642-25829-9_1,#Springer-VerlagBerlinHeidelberg2012 2 F.Eyidoganetal. 1.1 Introduction Plants have successfully evolved to integrate diverse environmental cues into theirdevelopmentalprograms.Sincetheycannotescapefromadverseconstraints, theyhavebeenforcedtocounteractbyelicitingvariousphysiological,biochemi- cal,andmolecularresponses.Theseresponsesincludeorleadtochangesingene expression,regulationofproteinamountoractivity,alterationofcellularmetab- olite levels, and changes in homeostasis of ions. Gene regulation at the level of transcription is one of the major control points in biological processes, and transcriptionfactorsandregulatorsplaykeyrolesinthisprocess.Phytohormones are a collection of trace amount growth regulators, comprising auxin, cytokinin, gibberellic acid (GA), abscisic acid (ABA), jasmonic acid (JA), ethylene, salicylicacid(SA),andfewothers(TutejaandSopory2008).Hormoneresponses are fundamental to the development and plastic growth of plants. Besides their regulatory functions during development, they play key roles and coordinate varioussignaltransductionpathways during responsestoenvironmentalstresses (WoltersandJ€urgens2009). A range of stress signaling pathways have been elucidated through molecular genetic studies. Research on mutants, particularly of Arabidopsis, with defects in theseandotherprocesseshavecontributedsubstantiallytothecurrentunderstand- ingofhormoneperceptionandsignaltransduction.Planthormones,suchasABA, JA,ethylene,andSA,mediatevariousabioticandbioticstressresponses.Although auxins, GAs, and cytokinins have been implicated primarily in developmental processes in plants, they regulate responses to stress or coordinate growth under stress conditions. The list of phytohormones is growing and now includes brassinosteroids (BR), nitric oxide (NO), polyamines, and the recently identified branchinghormonestrigolactone(Gray2004). Treatment of plants with exogenous hormones rapidly and transiently alters genome-wide transcript profiles (Chapman and Estelle 2009). In Arabidopsis, hormone treatment for short periods (<1 h) alters expression of 10–300 genes, with roughly equal numbers of genes repressed and activated (Goda et al. 2008; Nemhauseretal.2006;Paponovetal.2008).Notsurprisingly,longerexposureto most hormones ((cid:1)1 h) alters expression of larger numbers of genes. Complex networks of gene regulation by phytohormones under abiotic stresses involve variouscis-ortrans-actingelements.Someofthetranscriptionfactors,regulators, and key components functioning in signaling pathways of phytohormones under abioticstressesaredescribedinthiswork.Moreover,changesingeneexpression, protein synthesis, modification, or degradation initiated by or coupled to plant hormonesarebrieflysummarized.
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