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Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 resistance tocornborer, several partsoftheworld,for example,cornvarietieswith been deployedasprincipalmethodofinsectcontrolin resistance toinsectpests.Theresistantvarietieshave renewed interestinthedevelopmentofcropcultivarswith awareness ofenvironmentconservation,therehasbeena effects ofinsecticidesonnaturalenemies,and public development ofinsectresistancetoinsecticides,adverse traits duetocomplexandpolygenicinheritance.With the for resistancescreeningandslowtransferofinsect difficulties involvedinensuringadequateinsectinfestation developing -resistantcultivarsasaresultofthe achieved throughinsecticideuse,slowprogressin because oftherelativeeasewithwhichinsectcontrolis has notbeenassuccessfulbreedingfordiseaseresistance pest management(IPM),breedingforresistancetoinsects of theimportanceHPRasanintegralpartintegrated increasing demandforfood,feed,fodder, andfuel.Inspite pro-duction andproductivityofthecrops,tomeet the backboneofpestmanagementinordertoincrease Host plantresist-ance(HPR)assumesacentralrole,andis Introduction corn earworm, Keywords: Sharma C H and Dhillon K M paper Full Abstract $ Division ofEntomology, IndianAgriculturalResearchInstitute(IARI),NewDelhi-110 012,India. Pests Insect to Resistance Plant Host on Research in Shifts Paradigm E mail:[email protected] International CropsResearchInstitutefortheSemi-AridTropics (ICRISAT), Patancheru-502324,AndhraPradesh,India. to insectpests,inadditionthemore strategicresearch thatfeedsintotheseendeavors. systems willhavesubstantialimpactonbothconventionalandbiotechnologicalapproaches tobreed forresistance of phenotypingforresistance toinsectpestsremains acriticalconstraintinmanycrops. Improved phenotyping A good beginninghasbeenmadeindevelopinggenetic linkagemapsofmanycrops, buttheaccuracyandprecision of geneswithintheirgenomicenvironment, andwiththeenvironment inwhichtheirconferred phenotypeinteract. and adoptionofbiotechnology-derivedproducts willdependondevelopingafullunderstanding ontheinteraction resistance, understandnature ofgeneactionandmetabolicpathways,butrapidcosteffectivedevelopment diverse sources todevelopplantswithresistance toinsectpests,deploymentofmoleculartechniques forinsect . Although,considerableprogress hasbeenmadeoverthepasttwodecadesinmanipulatinggenesfrom to reduce pestassociatedcrop losses,andacceleratetheprogress indevelopingcultivarswithresistance to Conventional breeding inconjunctionwithmoleculartechniquesandtransgenicapproaches haveagreat promise Hostplantresistance,insectpests,molecularmarkers,genetictransformation,genepyramiding,RNAi Helicoverpa zea Ostrinia nubilalis (Boddie),sorghum togreen $ (Hubner)and Indian JournalofPlantProtection Vol. 40.No.1,2012(1-11) bug, of moleculartechniquesandmarker systemsinagricultural Last twodecadeshaveseena rapidprogressindeployment 2005, Dhillon developing insect-resistanthybrids(KameshwaraRao transfer ofinsectresistancegenesinparentallinesfor germplasm, closelyrelatedwildrelativesofcrops,and bases ofresistancethroughgenepyramidingfromcultivated advocacy anddeploymentoftechniquestodiversifythe therefore recentyearshaveobservedaparadigmshiftin of therecentlyreleasedvarieties/hybridsareinadequate,and cultivars. Moreover, thelevelsofinsectresistanceinmost has dilutedtheemphasisondevelopmentofinsect-resistant productivity offieldcropsthroughusehybridtechnology break yieldplateauforsustainableincreaseincrop in severalpartsoftheworld(Stoner, 1996).Thequestto destructor virescens Nilapavata lugens (Coquillett) inIndia;andricetobrownplanthopper, in Africa;sorghum tomidge, (Harris] inUSA;cottontojassid, - Therioaphis maculate Schizaphis graminum (Distant),andwheattoHessianfly, (Say)andwheatstemfly, et al. (Stal)andgreenleafhopper, , 2008,Sharma,2009). (Buckton)and (Rondani),andalfalfatoaphids Jacobiella facialis Stenodiplosis sorghicola Cephus cinctus Acyrthosiphon pisum Acyrthosiphon Nephotettix Mayetiola (Jacobi) Norton et al. , 1 www.IndianJournals.com Members Copy, Not for Commercial Sale

Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 2 nutritional qualityandyield(Sharma stresses (insectpests,diseases,andweeds),improve modify cropstoincreaseyields,impartresistancebiotic foreign genesintoplants,andprovidedopportunitiesto been madeinoverthepasttwodecadesintroducing new chaptertocropimprovement.Significantprogresshas to environmentalconditionsinaparticularregionhasadded into cultivarswithdesirableagronomictraits,andadaptation technology togetherwiththeabilitytoinsertspecificgenes and manipulatesinglegenesthroughrecombinantDNA improved cultivars(Sharma,2009).Theabilitytoisolate accelerate theprocessoftransferringinsectresistanceinto expressing resistancetoinsectpests,whichcanbeused these cropshavealsobeenfoundtobelinkedgenes have beendeveloped,andmolecularmarkersinmanyof , potato,rye,sorghum, soybean,tomato,andwheat density geneticlinkagemapsofseveralcropssuchasbarley, the genesassociatedwithresistancetoinsectpests.High- to manipulatetraitscontributingcropproductivity, and function ofcropplants,andrevolutionizedtheunderstanding sciences tounderstandgeneandgenomeorganization and been identifiedandsupplemented totheexistingresistance India (Table 1).Severalnewsources ofinsectresistancehave cultivars havebeenreleasedfor cultivationinmanycrops Smith, 2005,Sharma,2009), Quisenberry, 1999,Sharma two decades(PandaandKhush,1995,Clement against severalinsectspeciesindifferent cropsin thelast evaluated fortheidentificationofaccessionswithresistance Several thousandsofgermplasmcollectionshavebeen sources resistance Identification andutilizationofinsect components ofHPRtoinsectpestsoverthelasttwodecades. paper overviewstheparadigmshiftsinresearchondifferent technology forproducinginsect-resistantcropplants.This now lotsofemphasisisbeinggivenondeploymentthis genetically modifiedcropsthatproducetoxicproteins,and silencing response(RNAi),areastepaheadofexisting the world(James,2009).Thecrops,whichinitiateagene- deployed forcommercialcultivationinseveralcountriesof alone orstackedwithherbicideresistancegenes,havebeen and potatoforthemanagementofinsectpestswith plants, geneticallytransformedcotton,maize,rice,tomato, resistance toinsectshavebeeninsertedintoseveralcrop pests (Sharma,2009).Although,the develop cropcultivarsthatsuffer lessdamagefrominsect combination withconventionalhostplantresistanceto and smallRNAviruseshavebeenusedaloneorin ribosome inactivatingproteins,secondaryplantmetabolites, herbicide resistancegenes,proteaseinhibitors,plantlectins, from bacterium, Host Plant Resistance to Insect Pests Insect to Resistance Plant Host Bacillus thuringiensis et al. and severalinsect-resistant , 2003, Bt et al. Dhaliwal genesconferring , non-selective , 2004).Genes et al.,2004, Bt genes M KDhillonandHCSharma Mason), andstemborers, planthopper, Chilo suppressalis Chilo partellus Spodoptera frugiperda weevil, (Sharma borer, chickpea andpigeonpea(Sharma biguttula armigera also beenidentifiedagainstcottonbollworm, millet (SharmaandSullivan,2000).Sourcesofresistancehave Oriental armyworm, in sorghum (Sharma S. sorghicola, andheadbug, (Rondani), spottedstemborer, sorwheat (Smith,2005); Hessian fly, sugarcane borer, resistance tobrownplanthopper, greenleafhoppers, Many geneshavebeenidentifiedinricethatcontributefor incorporating resistancegenesintotheimprovedcultivars. indicates thedegreeofeaseordifficulty involvedin insect pestsisimportantforcropimprovement,which Information ongeneticsandinheritanceofresistanceto pests insect to resistance of Genetics crop improvementprograms. requires aconcertedeffort fromthescientistsinvolvedin demand infutureforsustainablecropproduction,andthis cultivars withmultipleresistancetostresseswillbeingreater backgrounds havebeendevelopedinseveralcrops,and Insect-resistant cultivarswithdesirableagronomic sources againstcornearworm, 2001). Resistanceto noxia control resistancetotheRussian wheataphid, al. control theexpressionofresistancetogreenbug(Zhu to evolutionofHessianflybiotypes.Elevendominantgenes and severalofthesegeneshavebeendeployedinresponse h4 resistance tothisinsect(Smith,2005). borer 1991, Katiyar white backedplanthopper, andgallmidge (KhushandBrar, control theinheritanceofresistancetobrownplanthopper, (Khush andBrar, 1991).Bothdominantandrecessivegenes midge resistance toHessianflyhas (Khush andBrar, 1991).Evaluationofgermplasmfor ) areinheritedasdominantorpartiallytraits, , 2004),whiletwelvegenes(both dominantandrecessive)

(Kurdj.)inwheat(Saidiand Quick, 1996,Liu is polygenic,andexistsinmanygenotypesofrice , Maruca vitrata white backedplanthopper, andyellowstemborer Bruchus pisorum et al. Ishida incotton;legumepodborer, H.armigera (Hubner),andleafhopper N. lugens M. destructor , 1999,SharmaandFranzmann,2000);pea (Swinhoe)inmaize( et al. Diatraea grandiosella , 2001),whileresistancetoyellowstem (Walker) inrice(Smith et al. separata , gallmidge (J.E. Smith),andspottedstemborer, (Geyer) inpigeonpeaandcowpea D. noxia Scirpophaga incertulas ghum shootfly, L.inpea(Clement , 2003,Sharma and greenbug, Calocoris angustatus

reported 29genesthatcontrol H. zea C. partellus et al. Kanta Orseolea oryzae Orseolea oryzae biotype Iinbarleyis , cornborer, (Dyar),fallarmyworm,

, 2005a);spottedpod All thegenes(except Amrasca biguttula et al. Atherigona soccata et al. (Walker) inpearl , sorghum midge, S. graminum , 1997); et al. et al. , 2005c);and (Walker) and Helicoverpa O. nubilalis Diuraphis (Lethiery) , 1994); , 1994). (Wood- brown et al.

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Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 (Williams resistance tofallarmyworm amounts ofvariationindifferent maizepopulationsfor al. epistatic aswelladditive-dominanceeffects (Warnock in sweetcorninvolvesmultiplegenes,andiscontrolledby al. controlled bydominantallelesattwoloci(Momhinweg Pod borer Pod Pigeonpea borer Pod Chickpea Leaf miner Soybean ICGV86031,FDRS10 aphid Mustard Rapeseed andmustard Tobaccocaterpillar Thrips Leaf miner Groundnut E302,303,ICSV700,705,SPV101 Head bug Midge borer stem Spotted Ganga4,5,7and9,safed2,Deccan101,103,Ageti,Kanchan,Kundan Shoot fly Sorghum borer stem Spotted borer stem Pink Maize Yellow stemborer Leaf folder leafhopper Green Manasarowar, Bhadra,Jyoti,Co42,MTU5249,46,Shyraksha,andDhanya TangnerGall midge and Amelbero White backedplanthopper planthopper Brown Rice Insect pests Table 1.Insect-resistant cultivarsofmajorfoodcrops released forcultivationinIndia Source: Dhaliwaletal.(2004) Pod fly , 1998).BothGCAandSCA effects explainsignificant 1995).ResistancetoeardamagebyEuropeancornborer et al. , 1995,1998).Inheritanceofmaysin content DJ 6514,AF28,ICSV197,745,88032 M 13,Robut33-1 KM 7 KM Sharaksha, andKavya CSM 388,Chencholam,ICSV239 Nimsoy ASD 20 IR 20andVani M 35-1,Swati,SPV491,ICSV700,705,PhuleYashoda, SPV1015 C 235,PantG114, Anupam,JG74,ICCV10,Dulia,Pusa261,Vijay, Vishal Deccan 101and103 Ratana, Sasyasree,Saket,andMTU5849 T 21,Bori,BDN2,ICPL332,MABahar, ICPL84060, PantA1,BSMR1 PBR 91,PusaJaiKisan,C294,Laha101,Kalyani,Regent,Tora, Sariahi ICGV 86031,IGCS156(M13),FDRS10 IR 36,Kakatiya,Surekha,Phalguna,Kunti,Shakti,Shamlei,Asha,Rajendradhan, Cultivars

and southwesterncornborer et et Indian JournalofPlantProtection Vol. 40.No.1,2012(1-11) corn leafaphid, effects explain the variationinexpressionofresistanceto action (Cartea additive, dominantandepistatic effects controlthe gene nonagrioides Snook, 2001).Stalkresistancetothestemborer, by thepresenceofamajormodifiergene(Widstrom and in maizeimpartsresistancetocornearworm (Lefebvre) isquantitativelyinherited,and et al.,2001).Bothadditiveanddominant Rhopalosiphum maidis (Fitch.) (Bingand , andisgoverned Sesamia 3 www.IndianJournals.com Members Copy, Not for Commercial Sale

Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 4 with resistancetoleafhoppers,isgovernedbyfivegenes Trichome density bollworms incotton,isdueto containing glands,whichareassociatedwithresistanceto into locallyadaptedcultivars.Inheritanceofgossypol cotton aregovernedbyoligogenes,andcanbetransferred the Azukibeanweevil, wild weevil, epistatic geneeffects (Park complex, andiscontrolledbyadditive,dominant, kraemeri Inheritance ofhookedtrichomesresponsiblefor to antixenosis mechanismofresistancein indicated theinvolvementofasingledominantgenein in effects, whilenon-additivetypeofgeneactionwasobserved to podborerin is controlledbymultiplegenes,andinheritanceofresistance t cytoplasm inChina(Shan bug, (Sharma recessive trait,andiscontrolledbyadditivegeneeffects et al. govern resistancetospottedstemborerinsorghum(Sharma al. to shootflyisinheritedbyadditivegeneaction(Dhillon Guthrie, 1991),andto characters associatedwithresistanceto type ofgeneeffects(Ratnadass Eurytylus oldi (Sharma controlled bybothadditiveandnon-additivegeneaction et al. to insectpests(Sharma,2001, Dhillon based hybridshavebeenreported tobehighlysusceptible the cowpeaweevil, Cardona, 1991).Resistanceinmungbean, complementary effect of tworecessivegenes(Kornegayand inherited asasimpledominanttrait(Tomaka wild mungbean, on pests. Mostofthesorghum hybridsgrowntodate arebased on theinfluenceofCMSexpressionresistancetoinsect physiological characteristicsindifferent cropplants, and of cytoplasmicmale-sterility(CMS)onmorphologicaland Considerable informationhasbeengeneratedontheeffects resistance insect for systems male-sterility cytoplasmic of Exploitation 5 (LacapeandNguyen,2005) kabuli , 2006e),whileadditiveandnon-additivegeneeffects H. armigera Milo Host Plant Resistance to Insect Pests Insect to Resistance Plant Host , 2008).Theexpressionofnonpreference andantibiosis , 2007).Resistancetosorghummidgeisinheritedasa C. angustatus Phaseolus (A Acanthoscelides obtectus types(Gowda et al. et al. RossandMooreresistanceinLimabeanis 1 ) cytoplasm,exceptafewhybridsbasedonA , 2000),whileresistancetoAfricanheadbug, 1996).However, resistancetosorghum head (Poppius)islargelycontrolledbyadditive desi and in accession,andisinheritedasa Vigna radiata is inheritedasapartiallydominanttrait chickpeaisgovernedbyadditivegene

Gossypium C. maculatus C. partellus Melanagromyza obtusa et al. Callosobruchus chinensis Callosobruchus et al. et al. , 2005).Verulkar G 13 . species,whichisassociated Resistance to , 2000).TheA , 1994).Resistancetobean et al.,2002).Mostofthe (Pathak, 1991).Resistance (Say),isderivedfroma var. (F.) is derivedfromthe allele(Calhoun,1997). et al. sublobata C. scarabaeoides Vigna radiata H. armigera , 2005,Dhillon et al. et al. H. armigera 1 cytoplasm Malloch. Empoasca , andis L.,and , 1992). , (1997)

in t to 1 et – 2 M KDhillonandHCSharma fly andRussianwheataphid (Table 2). striped ricestemborer;andwheat forresistancetoHessian hopper, yellowstemborer, and and sorghum midge;riceforbrownplanthopper, greenleaf and podsuckingbug;sorghum forshootfly, stemborer, armyworms; groundnut forleafminer, leafhoppers,podborer, and pod fly, andpodwasp;chickpeaforborerbruchids; leafhoppers; tomatoforfruitborer;pigeonpeapodborer, to pinkbollworm,cottonwhitefly, and to insects,forexample,wildrelativesofcottonresistance species ofseveralcropsassourcesgenesforresistance a paradigmshiftinidentificationanddeploymentofwild resistance tothetarget insects. hybridization toincreasethelevelsanddiversifybasisof showing highlevelsofresistancetoinsectscanbeusedinwide- resistance tobioticandabioticconstraints. Wild speciesofcropsareimportantsourcesgenesfor crops of relatives wild from genes resistance insect of Search pests forsustainablecropproduction R-lines todevelophybridswith pests, andtransferinsectresistancegenesintoA-,B-, cytoplasms fortheireffects oncultivarsusceptibilitytoinsect therefore, thereisacontinuingneedtoevaluatedifferent depending onthecropandpestspeciesinvolved; expression ofresistancetoinsectpestsanddiseases and therestorationabilitiesofrestorersinfluences the cytoplasmofmaintainerlineswithnucleargenes CMS, cytoplasmicfactors,theinteractionsoffactorsin systems incerealssuggeststhatthegeneticbackgroundof The analysesofliteratureavailableondifferent CMS maintainer linesofsorghum (Dhillon found tobesignificantlylowerinCMSascomparedthe to shootfly, midge, different mechanismsandtraitsassociatedwithresistance than theinbreds(KumarandMihm,1996).Expressionof saccharalis the A comparatively lesssusceptibleto (Sharma needed inboththeparentstoproduceinsect-resistanthybrids the geneticbackgroundofCMSlines,and expression ofresistancetotheseinsectsisinfluencedby resistant orsusceptiblerestorerlines,suggestingthat damage thanthehybridsbasedonsusceptibleCMSand shoot fly-resistantCMSandrestorerlinessuffered less Hybrids basedonshootbug,sugarcaneaphid,midge,and 2006). Furthermore,theA components ofresistanceto 1 , A 2 et al. , A washigherinresistantinbredlinesbasedhybrids 3 pea forbruchid;cowpeaspottedpodborer , A , 1996,Dhillon 4 G, A shoot bug,andsugarcaneaphid 4 VzM cytoplasms(Dhillon 4 M cytoplasmwasfoundtobe Last twodecadeshaveobserved et al.,2006c,Sharma multiple resistancetoinsect D. grandiosella A. soccata (Dhillon et al. , 2006b,c,d). damagethan et al. et al. resistance is Genotypes have been , 2008). and , 2005). et al., D. www.IndianJournals.com Members Copy, Not for Commercial Sale

Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 mapping inmanycrops(Mohan recombinant inbredlines(RILs)canbeusedforgene F acceptable combinationoftraits.Nearisogeniclines(NILs), sufficiently large scaletoidentifystableprogenywithan to thecomplexityofachievinginterspecifichybridsona from wildrelativesmaytakeconsiderablylongertimedue cultivars throughconventionalbreeding,whilegenetransfer to transferinsectresistancetraitsintothehigh-yielding in agriculture(Sharma,2009).Ittakesfivetosixgenerations wheat aresomeoftherecentbiotechnologicaldevelopments rice, potato,rye,sorghum, soybean,cowpea,tomato,and genetic linkagemapsofseveralcropssuchasbarley, maize, molecular markersforthedevelopmentofhigh-density model organisms anddeploymentofdifferent typesof diagnostics. Wholegenomesequencingofanumber population genetics,, plantbreeding,and genome mapping,genelocalization,evolution, characterization ofgeneticdiversity, genomefingerprinting, past fewyearstodetectDNAsequencepolymorphism, A widevarietyoftechniqueshavebeendevelopedinthe insects to resistance plant host for markers molecular of Deployment the wildrelativesintocultigen.Furthermore,marker valuable toolsforintrogressionofgenesinterestfrom However, markers developedfromsuchmapsmaybe to different recombinationpatterns(Fulton may havelimitedrelevanceincropbreedingprogramsdue polymorphism, butlinkagemapsderivedfromsuchcrosses genetic linkagestudiesduetohighlevelofdetectable populations frominterspecificcrossesareoftenusedfor Rice Chickpea Source: Sharma(2009) Wild species Tomato Crop Table 2.Wild speciesofdifferent crops identifiedassources ofgenesforresistance todifferent insectpests Cotton Sorghum Pigeonpea Groundnut 2 andbackcrosspopulations,doubledhaploids, Oryza officinalis Cicer bijugum C. acutifolius,albicans S. versicolor, S.matarankense S. extans,intrans,interjectum, Lycopersicon esculantum Gossypium thurberi,G.somalense and G. capitisviridis,raimondaii,trilobum,sinense,latifolium Sorghum australiense, S. purpureosericeum, S. brevicallosum, S. timorense, Atherigona soccata Atherigona Sorghum australiense,S.purpureosericeum, S.brevicallosum, S.timorense, Rhynchosia aurea A. stenosperma,paraguariensis,pusilla, Arachis cardenasii, A.duranensis,kempffmercadoi, A.monticola, and G.barbosanum .bahatalugens O. brachyantha

C. cuneatum, , , O. minuta R. bracteata, Cajanus scarabaeoides, C. sericeus, Helicoverpa armigera Helicoverpa R.bracteata,Cajanusscarabaeoides,C.sericeus, et al. , L. hirsutum, and , , 1997).Mapping O.latifolia

C. pinnatifidum,andjudaicu , Flemingia bracteata S. nitidum et al.,1997). , G. armourianum and and , S. stipodeum O.australiensis , S. angustum,ecarinatum, L. hirsutum and A. triseminata Indian JournalofPlantProtection Vol. 40.No.1,2012(1-11)

G. gossypiodies 1994, Biradar genomic regions,andplayamajorroleforresistancein D. grandiosella (Wang resistance toricegallmidgeanditsbiotypes(Mohan resistance (Selvi insects. Molecularmarkerslinkedtoyellowstemborer quantitative traitloci(QTLs)associatedwithresistanceto genetic linkagemapsofdifferent crops,toidentify used toevaluateDNApolymorphism,andfordeveloping (Smith, 2005).Severaltypesofmolecularmarkershavebeen resistance traitsinseveralcropstodifferent insectpests identifying genomicregionsandgenesassociatedwith Considerable progresshasbeenmadeintherecentpast with minimalamountoflinkagedragfromthewildrelatives. identify individualswithappropriatecombinationofgenes, of generationsandthesizepopulationsrequiredto dramatically speeduptheprocessbyreducingnumber assisted selection(MAS)hasshownthepotentialto responsible forresistancetostemboringby have beenfoundforresistancetoborers,theQTLs been reportedbyByrne depicting changeinmaysinconcentrationmaizesilkhave insect resistance.ThecornearwormresistanceQTLs 2005), havebeenmappedandusedinwheatbreedingfor wheat aphid(Ma biotypes (Dweikat responsible forresistancetoHessianflyanditsdifferent resistant rice.Severalpolymorphicmarkersandgenes mapped anddeployedinMASfordevelopinginsect- hopper (Sharma , O.granulata et al.,2004,Fujita et al.,2004,Jain , , and et al. et al. et al. et al. 2002),andQTLsgenesconferring D. saccharalis , 2003,Jena Insect pests Helicoverpa armigera Helicoverpa Sogatella furcifera Helicoverpa armigera Helicoverpa and bollworms andsuckinginsectpests Spodoptera litura, Spodoptera leafminer, , 1998),andgreenbug(Zhu et al. , 2004,Wang Stenodiplosis sorghicola et al. , and , (1996).Although,afewQTLs Helicoverpaarmigera, et al. , 2006),andbrownplant Scirpophaga et al. sharesomecommon et al. , 2004),leafhoppers , and Nilaparvata , , 2006),havebeen Chilo partellus , 2006),Russian Empoasca kerri

incertulas O. nubilalis et al. et al. , , , , 5 www.IndianJournals.com Members Copy, Not for Commercial Sale

Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 6 et al. with resistancetogreenbuganditsIKbiotypes(Agrama et al. stalk, andstemtunnelingbyEuropeancornborer(Krakowsky reported inMesoamerican major QTLfor have beenreportedincommonbean(Murrayetal. four markersto resistance toleafminer, responsible forresistancetothisinsect.TheQTLs BAC contigconstructionandmap-basedcloningofgenes plants suchasmaize,rice, wheat, sorghum,sugarcane, resistance toinsectshavebeen insertedintoseveralcrop and widenthepoolofusefulgenes. The genes intocropplantstoimpartresistanceinsectpests in isolation,cloning,andintroductionofforeigninsecticidal Significant progresshasbeenmadeoverthepasttwodecades isolate andmanipulategenesforcropimprovement. Recombinant DNAtechnologyhasopenedupnewvistasto resistance insect for Genetic transformation and gene pyramiding recessive gene(Herselman breeding lineICG12991,whichiscontrolledbyasingle Aphis craccivora includens mechanism ofresistanceto QTLs havebeenidentifiedinsoybeanforantibiosis sheath aciddetergentlignin,starchconcentrationin detergent fiber, aciddetergent fiberinleaf-sheaths,leaf- to maize (Khairallah resistance topotatoleafhopper, et al. components ofresistanceto mapping populationstoidentifyQTLslinkedvarious and theeffortsareunderwaytoevaluateinter-specific resistance topodborerinchickpeaisonlyjustbeginning, resistance inpigeonpea(Sharma,2009).Mappingfor to has beendeveloped,andisunderevaluationforresistance mapping populationbasedon also beenidentifiedintomato(Moreira resistance (Chen for Coloradopotatobeetle, where fifteenBACclonesharboredpolyphenoloxidaseloci potato, et al. in sorghum(Folkertsma responsible forresistancetoshootflyhavebeenidentified et al. of resistancetomidgehavebeenmappedinsorghum (Tao O. nubilalis H. armigera Host Plant Resistance to Insect Pests Insect to Resistance Plant Host , 2002,Nagaraj , 2007).SeveralQTLshavealsobeenfoundassociated , 2005b,Sharma,2009).Eightmarkersassociatedwith , 2006a).Two BAClibrariesofwildMexicandiploid , 2003).TheQTLsassociatedwithphenotypictraits Solanum pinnatisectum (Walker)(Terry appearstobecontrolledbyQTLsforneutral E. kraemeri Thrips palmi andidentifyQTLslinkedtopodborer Kochhasbeenidentifiedinthegroundnut et al. et al. et al. , 1997,Butron , 2004),whichmightbeusefulfor et al. 2005),andtwodifferentmechanisms

Liriomyza trifolii bean (Freietal. et al. Leptinotarsadecemlineata(Say) , andthreemarkerstobothspecies H. armigera et al. , 2003,Hash Karny resistancehasalsobeen C. cajan H. zea, , 2000).Resistancetoaphid, , 2004). Empoasca fabae , havebeenconstructed et al. and inchickpea(Sharma xC.scarabaeoides Bt et al. , 2005).Two major , 2005).Resistance genesconferring et al. (Burgess) have Pseudoplusia , 2003,Dhillon , 1999).A , 2004).A (Harris),

the M KDhillonandHCSharma to controlinsectpestsviainplanta expressionofadsRNA damage, suggestingthattheRNAi pathwaycanbeexploited and cottonaphid, cotton with for aviableinsectcontrolstrategy. Transgenic plantsof transgene andQTL-mediatedresistancecanbecombined corn earworm(Walker rootworm, have shownasignificantreductioninwesterncorn Transgenic cornplantsengineeredtoexpress WCRdsRNAs expression ofthetarget geneissuppressedbyRNAi. the plant,andconductRNAiagainsttarget gene,thereby insects' bodiesafterbeingingestedbytheinsectthateats formed intheplants.TheinterferingRNAsthenenterinto transgenic technique,andthentheinterferingRNAsare the dsRNAofinsect'sgeneisexpressedinplantsbyusing pests (GordonandWaterhouse, 2007).InRNAitechnology, genomics ofvaluablecroptraitsforresistanceagainstinsect demonstrated tobehelpfulinunderstandingthefunctional (Hannon, 2002).Recently, theRNAitechnologyhasbeen acids, andregulatestheexpressionofprotein-codinggenes endogenous parasiticandexogenouspathogenicnucleic transcriptional genesilencing,mediatesresistancetoboth oftenly knownasRNAinterference(RNAi)orpost- A conservedbiologicalresponsetodouble-strandedRNA, resistance plant host in technology (RNAI) interference RNA of Deployment Zilberman, 2003,DhillonandSharma,2009). and foodproducts,easiercropmanagement(Qaim organisms, reducedamounts ofinsecticideresiduesinfood use, reductioninharmfuleffects ofinsecticidesonnon-target have beenhigheryield,lowerinputcostsintermsofpesticide world. Thebenefitsofgrowingtransgeniccropstogrowers under commercialcultivationinseveralcountriesofthe resistance genes,andthesegeneticallymodifiedcropsare insect pests(Table 3),aloneorstackedwithherbicide potato forthemanagementoflepidopteranandcoleopteran three eventsofrice,oneeventtomato,and28 31 eventsofgeneticallymodifiedcotton,40maize, transgene, for Till date,atotalof14 develop aseriesofBC QTLs duringandafterthetwobackcrossesinsoybeanto been usedtotracktheantibiosis/antixenosisresistancelinked durable resistancetoinsectpests.Forexample,MAShas transformation canprovideagermplasmbasetoachieve insertion ofinsectresistancegenesthroughgenetic conventional/marker assistedhostplantresistancewith (Hilder andBoulter, 1999,Sharma pigeonpea, cowpea,groundnut,tomato,brinjal,andsoybean cotton, potato,tobacco,broccoli,cabbage,chickpea, Diabrotica virgifera virgifera Bt +GNA Bt andQTLconditioningresistanceagainst Aphis gossypii Bt conferredresistanceto 2 genes havebeendeployedthrough F et al.,2002),indicatingthat 3 plantswithorwithout Glover (Liu et al. , 2004).Combining LeContefeeding et al. H. armigera , 2003). cry1Ac Bt - www.IndianJournals.com Members Copy, Not for Commercial Sale

Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 and withtheenvironment inwhichtheirconferred the interactionofgeneswithin theirgenomicenvironment, products willdependondeveloping afullunderstandingon development, andadoption ofbiotechnology-derived action andmetabolicpathways,butrapidcosteffective techniques forinsectresistance,understandnatureofgene with resistancetoinsectpests,deploymentofmolecular manipulating genesfromdiversesourcestodevelopplants progress hasbeenmadeoverthepasttwodecadesin and increasecropproductivity. Although, considerable the progressindevelopingcultivarswithresistancetoinsects promise toreducepestassociatedcroplosses,andaccelerate molecular techniquesandtransgenicapproacheshaveagreat Augmentation ofconventionalbreedingwiththeuse Conclusions gossypol. When directly involvedintheabilityofcottonbollwormtotolerate the firstgossypol-inducibleP450genefrombollworms,is (Baum to down-regulate gossypol isgreatlyreduced(Mao achieved byplant-mediatedRNAi,thelarvaltoleranceto genetically modifiedplants. be saferandlesslikelytohaveunintendedeffects thanother crops target particulargenesininsects,theywill crops suchascorn,cottonandsoybeans,thenew the currentstrategyofexpressing action forthecontrolofinsectpeststhatcouldcomplement potent RNAiinthepest.providesauniquemodeof would notprovidesufficient levelsofintactdsRNAtotrigger hairpin RNAinplantmaterialtobeingestedbyaninsect using thisapproachseemedtosuggestthatsimplyexpressing However, thepassageofyearswithoutreportssuccess approach forgene-silencinginherbivorousinsects. leaves, suggeststhatplant-mediatedRNAimaybeageneral midgut byfeedingcottonbollwormsdsRNA-producing oao1 3 Tomato Rice az 40 Maize oao28 Source: CBD(2010);ILSIResearchFoundation(2010) Potato 31 Cotton No.ofevents Crop insect pests Table 3.Commercially deployed et al. , 2007).CytochromeP450gene( CYP6AE14 CYP6AE14 expressionissuppressed,as Lepidopteran andColeopteran and Bt et al. GST1 Bacillus thuringiensis insecticidalproteinsin Lepidopteran Lepidopteran Lepidopteran Coleopteran Insect pests Insect , 2007).Theability expressioninthe CYP6AE14 ) genesthrough genetically modified crops for resistance to Indian JournalofPlantProtection Vol. 40.No.1,2012(1-11) cry1Ac cry1Ac cry1A.105 cry1Ab flcry1Ab cry3A cry1Ac Gene(s) Agrama HA,Widle GE,ReeseJC,CampbellLRandTuinstra References Bing JWandGuthrieD1991. T,Baum JA,Bogaert Clinton W, HeckGR,FeldmannP, use ofbiotechnologicaltools. based onre-engineeredbreedingprogramstomakebest insects, anddevelopmentofnewparadigmsinbreeding precise mappingoftheQTLsassociatedwithresistanceto crops forothercomponentsofplantresistancethrough resistance toinsectpests,andstrengthen molecular techniquestodevelopcultivarswithimproved pests andabioticstresses.Therefore,thereisaneedtouse case ofmorecomplextraitssuchasresistancetoinsect readily identified.Thesamepotentialmaybeachievedin quality traitswheresimplyinheritedcomponentscouldbe the privatesector, inbreedingfordisease-resistanceand assisted selectionhashadadramaticimpact,particularlyin strategic researchthatfeedsintotheseendeavors.Marker- breed forresistancetoinsectpests,inadditionthemore on bothconventionalandbiotechnologicalapproachesto Improved phenotypingsystemswillhavesubstantialimpact insect pestsremainsacriticalconstraintinmanycrops. accuracy andprecisionofphenotypingforresistanceto developing geneticlinkagemapsofmanycrops,butthe phenotype interact.A good beginninghasbeenmadein Biradar SK,SundaramRM, ThirumuruganT, BenturJS, Applied Genetics resistance andtoleranceinSorghum bicolor. M R2002. Aphididae). resistance inmaizetothecorn leafaphid(Homoptera: 1322-1326. pests throughRNAinterference. Vaughn TandRobertsJ2007. Ilagan O,JohnsonS,PlaetinckG,MunyikwaT, PleauM, , , cry2Ab2 and cry1Ac , cry2Ab2 cry1Ab GeneticmappingofQTLsassociatedwithgreenbug , Journal ofEconomicEntomology cry9C , cry1Ab ,

cry3A 104 : , cry34Ab1 , cry1C , 1373-1378.

and vip3Aa20 , cry1A , cry35Ab1 Generationmeananalysisfor Control ofcoleopteraninsect Nature Biotechnology , vip3A(a) , cry1F Theoretical and 84 : , Bt cry1F , cry3Bb1 1080-1082. transgenic , and

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Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 8 Byrne PF, McMullenMD,SnookE,MusketTA,TheuriJ Butron A,Tarrio R,RevillaP, OrdasAandMalvarR2005. Chen Q,SunS,Ye Q,McCuine S,HuffEandZhangHB Cartea ME,MalvarRA,Vales MJ,BurtonAandOrdas Calhoun DS1997. Dhillon MKandSharmaHC 2009. Dhaliwal GS,SinghRandJindalV2004. Crouch JH,GaurPM,BuhariwallaHK,Barmanand Clement SL,SharafEl-DinN,Weigand SandLateef1994. Convention onBiodiversity(CBD)2010. Clement SLandQuisenberryS.(Eds.)1999. Journal ofAgriculturalSciences (SpecialIssue) different trophic levels,andseedcottonyield. cotton ontarget andnontarget ,toxinflowthrough Sciences, USA factor, inmaizesilks. of theconcentrationmaysin,acornearwormresistance Quantitative traitlociandmetabolicpathways:Geneticcontrol M, Widstrom NW, Wiseman BRandCoeEH1996. Applied Genetics selection forresistancetopinkstemborer. Molecular changesinthemaizecompositeEPS12during Theoretical andAppliedGenetics rice gallmidgeresistancegene P N2004. Amudhan S,ShenoyVV, MishraB,BennetJandSharma Mexican diploidpotato, 2004. Journal ofEconomicEntomology Sesamia nonagrioides 2001. resistance traitincotton. New Delhi,India.Pp517-558. Dhaliwal GSandRamSingh),PanimaPublishingCorporation, Resistance toInsects:ConceptsandApplications(eds. and insectpestmanagement:Progresspotential.HostPlant New Delhi,India.Pp307-328. Research (ed.SharmaHC)OxfordandIBHPublishingCo, Management: Emerging Trends andStrategiesforFuture Helicoverpa Sharma HC2005. Portal. http://bch.cbd.int/database/lmo-registry. Organism (LMO)Registry-BiosafetyClearing-HouseCentral cool seasonfoodlegumes. Research achievementsinplantresistancetoinsectpestsof Florida, USA:CRCPress.295pp. Genetic ResourcesforInsect-ResistantCrops.BocaRaton, 1002-1009. beetle resistanceloci. identification ofclonesnearthelateblightandColoradopotato Host Plant Resistance to Insect Pests Insect to Resistance Plant Host Inheritanceofresistancetoeardamagecausedby ConstructionoftwoBAClibrariesfromthewild IdentificationofflankingSSRmarkersforamajor resistanceingrainlegumes.

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108 : M KDhillonandHCSharma Fulton T, Beck-BunnT, EmmattyD,Eshed Y, LopezJ,Petiard Dhillon MK,SharmaHC,RamSinghandNaresh JS2006d. Dhillon MK,SharmaHC,FolkertsmaRTandChandraS Dhillon MK,SharmaHCandSmith2008. Dhillon MK,SharmaHC,ReddyBVS,RamSinghand Dhillon MK,SharmaHC,PampapathyGandReddyBVS Dhillon MK,SharmaHC,Naresh JS,RamSinghand Fujita D,DoiK,Yoshimura AandYashi H2006.Molecular Dweikat I,OhmH,PattersonF, andCambron S.2004. Dhillon MK,SharmaHC,ReddyBVS,RamSingh,Naresh Frei A,BlairMW, Cardona C, BeebeSE,GuHandDorn RT,Folkertsma SajjanarGM,ReddyBVS,SharmaHC rice leafhopper(Nephotettixcincticeps and Genetics shoot fly, physico-chemical traitsassociatedwithresistancetosorghum Influence ofcytoplasmicmale-sterilityonexpression 66-68. partellus ( Euphytica shoot fly-resistantand-susceptibleparentstheirhybrids. 2006a. Natural Resources Prospectives inAgriculture, Veterinary Science,Nutritionand deployment ofpestresistanthybridsincereals. of cytoplasmicmale-sterilitysystemsfordevelopmentand fly, Naresh JS2006e. resistance toshootbug( 2006c. 99 (4): Atherigona soccata on different mechanismsofresistanceinsorghum toshootfly, Pampapathy G2006b. sativa Euphytica mapping ofanovelgene, genes inwheat. Identification ofRAPDmarkersfor11 Hessianflyresistance sterile cytoplasmsinsorghum toshootfly, J SandKaiZ2005.Relativesusceptibilityofdifferent male- common bean. 2005. 42 pp. Genome XI,11-15 January, 2003.SanDiego,California,USA. ( sorghum shootfly( and HashCT2003. 423. Melanaphis sacchari Atherigona soccata L. QTL Mappingofresistanceto Geneticdivergence andmolecularcharacterizationof Cytoplasmicmale-sterilityaffects expressionof 1452-1461. ). Theoretical andAppliedGenetics

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Downloaded From IP - 220.227.242.220 on dated 23-Aug-2012 Gordon KHJandWaterhouse PM2007. Jena KK,JeungJU,LeeH,ChoiHCandBrarDS2006. Hilder VAandBoulterD1999. Kameshwara RaoN,MallikarjunaSharmaKK,ReddyL James C2009. Jain A,AriyadasaR,KumarSrivastavaMN,Mohan ILSI Research Foundation(2010).GMCropDatabase,Center Gowda CLL,RameshS,ChandraSand Upadhyaya H D Herselman L,ThwaitesR,KimminsFM,CourtoisB,vander Hash CT, FolkerstmaRT, RamuP, ReddyBVS, Hannon GJ2002.RNAinterference. Applied Genetics BPH resistanceinrice( wild species. cultivated tomatoandcomparisonswithQTLsfoundinother of anadvancedbackcross V, UhligJ,ZamirDandTanksley SD1997. proof plants. resistance gene, High resolutionmappingofanewbrownplanthopper(BPH) Acquisition ofAgri-BiotechApplications,Ithaca,NY, USA. crops: 2009. 18 : plants forinsectresistance-acriticalreview. Theoretical andAppliedGenetics resistance totheaphidvectorofgroundnutrosettedisease. Future Research(ed.SharmaHC) Helicoverpa of cropsforresistanceto J, SinghPandGotmare V2005. Applied Genetics marker-aided selectionandgenepyramiding. resistance gene, and NairS2004. DC [email protected] Sciences Institute1156 15thStreet,NWSuite200Washington, for EnvironmentalRiskAssessment,InternationalLife 2005. AFLP markerslinkedtopeanut( Merwe PJandSealSE2004. Bolona, Italy. 82pp. the GeneRevolution,May27-31,2003.UniversityofBologna, the Wake oftheDoubleHelix:FromGreenRevolutionto tolerance andresistancetoshootfly Marker assistedbreedingacrossICRISAT forterminaldrought R, HaussmannBIG,FergusonMEandCrouch JH2003. Mahalakshmi V, SharmaHC,RattundeFW, Weltzein E 199-214. arietinum resistance andgrainyieldin 177-191. Genetic basisofpodborer( L.)underunprotectedconditions. Management:Emerging Trends andStrategiesfor ISAAA BriefNo.41 Globalstatusofcommercializedbiotech/GM Nature Biotechnology Theoretical andAppliedGenetics Gm8 Bph18(t), 112 :

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RNAiforinsect- 418 : chickpea(Cicer Theoretical and Euphytica Crop Protection Theoretical and in sorghum. In 95 : QTL analysis 244-251. . Heliothis / 881-894. tothe 145 : L.) ) Indian JournalofPlantProtection Vol. 40.No.1,2012(1-11) Katiyar SK,Tan Y, HuangB,ChandelG,XuY, ZhangY, Xie Kanta U,DhillonBDandSekhonS1997. Lacape JMandNguyenTB2005. Kumar HandMihmJA1996. Khush GSandBrarD1991. Khairallah M,HoisingtonD,GonzalezdeLeonBohn Liu Z,GuoWZhuXF, ZhuZandZhangT2003. Liu X,SmithCM,GillBSandTolmay V2001. Krakowsky MD,LeeandHollandJB2007. Kornegay JLandCardonaC1991. Mao YB,CaiWJ,Wang JW, HongGJ,Tao XY, Wang LJ, Ma ZQ,SaidiA,QuickJSandLapitanNLV1998. gall midgeinChina. which confersresistanceagainstfourbiotypesofAsianrice Z andBennettJ2001. Center, Mexico,DF. Pp246-254. (ed. MihmJA)InternationalMaizeandWheatImprovement borer. InsectResistantMaize:RecentAdvancesandUtilization development ofmaizegermplasmforresistancetospottedstem Co, NewDelhi,India.Pp243-253. of Heredity loci associatedwithleafandstempubescenceincotton. Fabricius. grandiosella inbreds tofirst-generationsouthwesterncornborer, corn borerinmaize. components andresistancetostalktunnelingbytheEuropean correlation andmultivariateQTLanalysesforcellwall crossed tocommercialbeancultivars. Acanthoscelides obtectus in cropplants. Improvement Center, Mexico,DF. Pp148-154. Utilization (ed.MihmJA)InternationalMaizeandWheat tropical maize.InsectResistantMaize:RecentAdvancesand southwestern cornborerandsugarcaneresistancein J A1997. Melchinger A,JewellDC,DeutschJMihmand 953-961. Inheritance analysisofresistancetransgenic wheat. markers linkedtosixRussianwheataphidresistancegenesin line to Biotechnology 1307-1313. larval toleranceofgossypol. P450 monooxygenasegenebyplant-mediated RNAiimpairs Huang YPandChenX2007. Dn4 mapping ofRussianwheataphidresistancegenes in wheat. Theoretical andAppliedGenetics Helicoverpa armigera Crop Protection

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