Aphid Vectors and Grass Hosts of Barley Yellow Dwarf Virus and Cereal Yellow Dwarf Virus in Alabama and Western Florida by Buyun
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AphidVectorsandGrassHostsofBarleyYellowDwarfVirusandCerealYellow DwarfVirusinAlabamaandWesternFlorida by BuyungAsmaraRatnaHadi AdissertationsubmittedtotheGraduateFacultyof AuburnUniversity inpartialfulfillmentofthe requirementsfortheDegreeof DoctorofPhilosophy Auburn,Alabama December18,2009 Keywords:barleyyellowdwarf,cerealyellowdwarf,aphids,virusvectors,virushosts, Rhopalosiphumpadi , Rhopalosiphumrufiabdominale Copyright2009byBuyungAsmaraRatnaHadi Approvedby KathyFlanders,Co-Chair,AssociateProfessorofEntomologyandPlantPathology KiraBowen,Co-Chair,ProfessorofEntomologyandPlantPathology JohnMurphy,ProfessorofEntomologyandPlantPathology Abstract Yellow Dwarf (YD) is a major disease problem of wheat in Alabama and is estimated to cause yield loss of 21-42 bushels per acre. The disease is caused by a complex of luteoviruses comprising two species and several strains, including Barley yellowdwarfvirus (BYDV),strainPAV,and Cerealyellowdwarfvirus (CYDV),strain RPV. The viruses are exclusively transmitted by aphids. Suction trap data collected between1996and1999inNorthAlabamarecordedthe presence of several species of aphidsthatareknowntobeB/CYDVvectors. Aphidsweresurveyedinthebeginningofplantingseasonsinseveralwheatplots throughout Alabama and western Florida for four consecutive years. Collected aphids wereidentifiedandbioassayedfortheirB/CYDV-infectivity.Thissurveyprogramwas designedtoidentifytheaphid(Hemiptera:Aphididae)speciesthatserveasfallvectorsof B/CYDVintowheatplanting.From2005to2008,birdcherry-oataphid, Rhopalosiphum padi (L.), rice root aphid, Rhopalosiphum rufiabdominale (Sasaki), and greenbug, Schizaphis graminum (Rondani), were consistently found between October and December. The species of aphids and their timing of appearance in wheat plots were consistentwithflightdatacollectedinNorthAlabamabetween1996and1999.Both R. padi and R.rufiabdominale werefoundtocarryandtransmitBYDV-PAVandCYDV- RPV. Low overall numbers of collected aphids and low proportion of infective aphid madeitdifficulttoconclusivelyidentifytheprimaryvectorofB/CYDVinAlabama. ii The source of summer/fall infection of BYDV and CYDV is not known in Alabama.Pasturegrassesmayprovideameansofsurvivalduringsummermonthswhen wheatisnotavailableinAlabama.Varietyplotsofthreepasturegrassesweresampledin summerbetween2007and2009forB/CYDVandaphids.Ofthethreepasturegrasses surveyed, bahiagrass was found to consistently harbor BYDV-PAV and CYDV-RPV, whilelimpograsswasfoundtoharborBYDV-PAVandCYDV-RPVirregularly.Thisis thefirstreportofBYDVonbahiagrassandlimpograss.Noaphidswerefoundduringthis studyonlimpograssandgamagrasswhiletheaphid Siphaflava (Forbes)wasfoundin twooutofthreeyearsofsurveyonbahiagrass. iii Acknowledgments Itisapleasuretoacknowledgetheencouragement,forbearanceandexpertiseof manythatcarriedthisworktoitscompletion.Iamdeeplyindebtedtotheco-chairofmy advisory committee, Kathy Flanders, for her patience and expertise in guiding me throughthelongandwindingroad,literally,offieldresearchandacademicjourney.Her genuine friendship and mentorship are invaluable for me. Co-chair Kira Bowen and committeememberJohnMurphybothtrainedmeinacquiringnewskillscrucialforthis work.Theirexamplesofcriticalandbroadthinking,andtotakelifewithapinchofsalt (andanextrahelpingofdarkhumor),willsurelyhavetheirprintsinmycareerandlife. SusanHalbertfromFloridaDepartmentofPlantIndustrysetmeoffonmypathinaphid identification,ofwhichIreallyamgrateful.TheworkisfundedbytheAlabamawheat andfeedgraincheck-offcommittee. AspecialthankstoThomasWilliamsoftheNavigators.Hisfriendshipandthe exemplary life that he lived before my eyes are precious forever. Ryan Schumacher, NirmalAndrews,PaulDawson,TrafinaJadhaf,FranklinQuarcoo,MosesNtam,Michael Madan Mohan, Saisree Uppala and Dayton Cook have surrounded me with encouragementsandjoyofsharedlives. MyfamilyinIndonesia,asalways,hasbeenaconstantsourceofstrength and wisdom.Theyespeciallyknowhowtoteachmehumilityandputmyimportantworkin iv perspective,“So,youaregoingtogetadegreeforcrawlingaroundfieldsandcounting insects?” Andasforadedication,I’mtryingstilltoheedanitinerantpreacher’swordslong ago, “ …whether you eat or drink or whatever you do (e.g. do research and write a dissertation),doitallforthegloryofGod.” v TableofContents Abstract...............................................................................................................................ii Acknowledgments..............................................................................................................iv ListofTables.....................................................................................................................ix ListofFigures.....................................................................................................................x ListofAbbreviations.........................................................................................................xi 1 Introduction.................................................................................................................1 2 LiteratureReview........................................................................................................4 2.1 Thehosts..............................................................................................................5 2.2 Theviruses...........................................................................................................5 2.3 Thevectors...........................................................................................................8 2.3.1 Taxonomyandlifecycle...............................................................................8 1.1.1 Aphidmovement.........................................................................................10 1.1.2 BYDVandCYDVvectorsinAlabama......................................................12 2.4 Virus-hostrelationships......................................................................................19 2.4.1 Symptomformation....................................................................................19 2.4.2 Physiologyofyieldreductionininfectedplants.........................................23 2.4.3 Virussystemicmovementinsidetheplant.................................................24 2.5 Host-vectorrelationships....................................................................................25 2.5.1 Aphidreproductiononhealthyhosts..........................................................25 vi 2.5.2 Aphiddispersalandhostfindingstrategy...................................................26 2.6 Virus-vectorrelationships..................................................................................28 2.7 Virus-vector-hostrelationships..........................................................................30 2.7.1 Aphidpopulationdynamicsandalataeproductiononinfectedhost..........30 2.7.2 Aphidattractiontoinfectedhosts...............................................................31 2.7.3 Aphidfeedingbehavioroninfectedhosts..................................................32 2.8 Environmentalrelationships...............................................................................33 2.8.1 Temperatureandrainfall.............................................................................33 2.8.2 Cropmanagementpractices........................................................................35 2.8.3 Otherfieldcharacteristics...........................................................................38 2.8.4 Naturalenemies..........................................................................................39 2.8.5 Plantresistanceandtolerance.....................................................................40 2.8.6 Chemicalcontrolandforecastingschemes.................................................46 3 Long-TermGoal,ObjectivesandHypothesis...........................................................60 4 Species Composition of the Aphid Vectors of Barley Yellow Dwarf Virus and CerealYellowDwarfVirusinAlabamaandWesternFlorida.................................61 4.1 Introduction........................................................................................................61 4.2 Methodology......................................................................................................62 4.3 Results................................................................................................................65 4.4 Discussion..........................................................................................................67 4.5 Conclusion..........................................................................................................75 5 Perennial Pasture Grasses as Hosts of Barley Yellow Dwarf Virus and Cereal YellowDwarfVirus..................................................................................................81 vii 5.1 Introduction........................................................................................................81 5.2 Methodology......................................................................................................83 5.3 Results................................................................................................................87 5.4 Discussion..........................................................................................................88 5.5 Conclusion..........................................................................................................92 References.........................................................................................................................95