生命科学概论论文.docx
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生命科学概论论文
Anintroductiontolifescience
Courseassignments
BtandMonarch
Despitewidespreadadoptionofgeneticallymodifiedcropsinmanycountries,heatedcontroversiesabouttheiradvantagesanddisadvantagescontinue.Especiallyfordevelopingcountries,thereareconcernsthatgeneticallymodifiedcropsfailtobenefitsmallholderfarmersandcontributetosocialandeconomichard-ship.Manyeconomicstudiescontradictthisview,butmostofthemlookatshort-termimpactsonly,sothatuncertaintyaboutlonger-termeffectsprevails.So,BtornotBt,thatisaquestion.
Plantresistancetoinsectpestshasevolvednaturallyovermanymillionsofyearsandinvolves(i)bothconstitutiveandinduciblephytochemicalandmorphologicalmechanismsinplants,(ii)counteradaptationstoplantdefensesbytheherbivores,and(iii)bioticinteractionsofthemultitrophiclevelcommunitiesofinsectpathogens,parasites,andpredators(1–4).Theoutcomesofsuchcomplexbioticinteractionsaresometimesdeterminedbylocalmosaicsofabioticenvironmentalconditionsorregionalclimatechangesthatdirectlyinfluencethecomponentparticipantsandtheirbehavioral,physiological,andgeneticadaptations(5–7).Theintentionalselectionandbreedingofinsectandyorpathogen-resistantplantgenotypessuchascorn(Zeamays)hasresultedinslow,butsteady,progressagainststalk-boringlarvaesuchastheEuropeancornborerandothersuchspecies,andhostplantresistanceingeneralhassignificantlyreducedtheneedforbroad-spectruminsecticidesacrossagro-ecosystemsandforests(8,9).Ithasrecentlybeenpossibletogeneticallyengineerplants,suchasZ.mays,toavoidproblemswithinsecticidalspraydriftandmoreefficientlyexpresstoxinsinparticularplanttissuesagainstlepidopterouspests.Suchefficient,effective,andrelativelysafetoolsforpestmanagementhaveeludedclassicalplantbreedersandintegratedpestmanagementpractitionersformanydecades(10).However,eventhoughsafeformostothertaxonomicgroupsofinsectsotherthanLepidoptera,thenontargetimpactsofBtrequirecarefulandthoroughevaluationcombinedwithextensivepublicizing.Ifnotdone,therealrisks,whichmaybeminimal,canbesignificantlyamplifiedinthepublicmindsbyperceivedrisksanddistrustofbiotechcorporationsandtheregulatoryfederalagenciesandsimilargoal-affiliatedagriculturalresearchinstitutionsanduniversities(11,12).AdditionalrisksforcroppestmanagementsuchastherapiddevelopmentofinsectresistancetotheBtinsecticidesalsoneedseriousattentionandconsiderationinlong-termagroecosystemdesignaswell(13,14)
Recentpublicconcernsoverthetrans-genic(geneticallymodified)plantsandnontargetimpactssuchasthosefromBt-toxinexpressingcornpollenonthemonarchbutterflypopulationshaveescalated,despitegoodpestmanagementintentionsandgoodscience(15–23).Asinglelaboratorystudyonmonarchbutterflieshaspromptedwidespreadconcernthatcornpollen,engineeredtoexpressBacillusthuringiensis(Bt)endotoxin,mighttravelbeyondcornfieldsandcausemortalityinnontargetlepidopterans.Amongthelepidopteransathighpotentialriskfromthistechnologyistheblackswallowtailbutterfly,Papiliopolyxenes,whosehostplantsinthemidwesternU.S.arelocatedprincipallyinnarrowstripsbetweenroadsandcropfields.AfieldstudywasperformedtoassesswhethermortalityofearlyinstarblackswallowtailswasassociatedeitherwithproximitytoafieldofBtcornorbylevelsofBtpollendepositiononhostplants.PottedhostplantswereinfestedwithfirstinstarblackswallowtailsandplacedatintervalsfromtheedgeofafieldofBtcorn(Pioneer34R07containingMonsantoevent810)atthebeginningofanthesis.TheyconfirmedbyELISAthatpollenfromtheseplantscontainedCry1Abendotoxin(2.12560.289ngyg).Althoughmanyofthelarvaediedduringthe7daysthattheexperimentswererun,therewasnorelationshipbetweenmortalityandproximitytothefieldorpollendepositiononhostplants.Moreover,pollenfromthesesameplantsfailedtocausemortalityinthelaboratoryatthehighestpollendosetested(10,000grainsycm2),alevelthatfarexceededthehighestpollendensityobservedinthefield(200grainsycm2).theyconcludethatBtpollenofthevarietytestedisunlikelytoaffectwildpopulationsofblackswallowtails.Thus,theirresultssuggestthatatleastsomepotentialnontargeteffectsoftheuseoftransgenicplantsmaybemanageable.
LaboratorytestswereconductedbyRichardLtoestablishtherelativetoxicityofBacillusthuringiensis(Bt)toxinsandpollenfromBtcorntomonarchlarvae.ToxinstestedincludedCry1Ab,Cry1Ac,Cry9C,andCry1F.Threemethodswereused:
(i)purifiedtoxinsincorporatedintoartificialdiet,(ii)pollencollectedfromBtcornhybridsapplieddirectlytomilkweedleafdiscs,and(iii)Btpollencontaminatedwithcorntasselmaterialapplieddirectlytomilkweedleafdiscs.BioassaysofpurifiedBttoxinsindicatethatCry9CandCry1Fproteinsarerelativelynon-toxictomonarchfirstinstars,whereasfirstinstarsaresensitivetoCry1AbandCry1Acproteins.Olderinstarswere12to23timeslesssusceptibletoCry1Abtoxincomparedwithfirstinstars.Pollenbio-assayssuggestthatpollencontaminants,anartifactofpollenpro-cessing,candramaticallyinfluencelarvalsurvivalandweightgainsandproducespuriousresults.TheonlytransgeniccornpollenthatconsistentlyaffectedmonarchlarvaewasfromCry1Abevent176hybrids,currently<2%cornplantedandforwhichre-registrationhasnotbeenapplied.ResultsfromtheothertypesofBtcornsuggestthatpollenfromtheCry1Ab(eventsBt11andMon810)andCry1F,andexperimentalCry9Chybrids,willhavenoacuteeffectsonmonarchbutterflylarvaeinfieldsettings.
ToassessthelikelihoodthatmonarchlarvaewillbeexposedtoBacillusthuringiensis(Bt)pollen,KarenS.etalstudiedmilkweedandmonarchdensitiesinhabitatswhichcomprisemuchofthelandavailabletobreedingmonarchs,e.g.,cornfields,cornfieldedges,otheragriculturalfields,andnonagriculturalareas,infourregionsofthemonarchbreedingrange.Theyfoundthatmonarchsusemilkweedincornfieldsthroughouttheirbreedingseason,andthatperplantdensitiesareashighorhigherinagriculturalhabitatsasinnonagriculturalhabitats.Asaresultoftheprevalenceofagriculturalland,mostofthemonarchsproducedintheupperMidwestarelikelytooriginateincornfieldsorotheragriculturalhabitats.Therewasagreatertemporaloverlapbetweensusceptiblemonarchsandcornanthesisinthenorthernthanthesouthernpartofthesummerbreedingrange,becauseofearlierpollenshedinthesouth.Theimportanceofagriculturalhabitatstomonarchproductionsuggeststhat,regardlessoftheimpactofgeneticallymodifiedcrops,agriculturalpracticessuchasweedcontrolandfoliarinsecticideusecouldhavelargeimpactsonmonarchpopulations.
AcollaborativeresearcheffortbyscientistsinseveralstatesandinCanadahasproducedinformationtodevelopaformalriskassessmentoftheimpactofBtcornonmonarchbutterfly(Danausplexippus)populations.InformationwassoughtontheacutetoxiceffectsofBtcornpollenandthedegreetowhichmonarchlarvaewouldbeexposedtotoxicamountsofBtpollenonitshostplant,thecommonmilkweed,Asclepiassyriaca,foundinandaroundcornfields.ExpressionofCryproteins,theactivetoxicantfoundinBtcorntissues,differedamonghybrids,andespeciallysointheconcentrationsfoundinpollenofdifferentevents.Inmostcommercialhybrids,Btexpressioninpollenislow,andlaboratoryandfieldstudiesshownoacutetoxiceffectsatanypollendensitythatwouldbeencounteredinthefield.Otherfactorsmitigatingexpo-
sureoflarvaeincludethevariableandlimitedoverlapbetweenpollenshedandlarvalactivityperiods,thefactthatonlyaportionofthemonarchpopulationutilizesmilkweedstandsinandnearcornfields,andthecurrentadoptionrateofBtcornat19%ofNorthAmericancorn-growingareas.Their2-yearstudysuggeststhattheimpactofBtcornpollenfromcurrentcommercialhybridsonmonarchbutterflypopulationsisnegligible
ConceptualModel.Riskassessmentrequiresknowledgeoffouressentialcomponents:
(i)hazardidentification,(ii)natureofdoseresponsetoatoxin,(iii)probabilityofexposuretoaneffectivedose,and(iv)characterizationofrisk(24).ComponentsofariskassessmentapproachasappliedtothecaseofBtcornandmonarchbutterflyaredepictedinFig.1.Btproteinsexpressedincornplanttissuescanbringaboutspecificreactionsinthegutoflepidopteranlarvae(25),includingnontargetlarvaethatconsumeBtcornpollen.Themagnitudeofthereactionwilldependonthetypeofproteinproducedbyvarioustransgeniceventso-fhybridBtcorn,theamountofproteinexpressedinpollengrainsfromdifferentevents,theamountofpollenconsumedbylarvaeofdifferentdevelopmental
stages,andthesusceptibilityoflarvaetotheBtprotein.ThatahazardmayexistwassuggestedbyLoseyetal.(26).Characterizationoftoxiceffectsisnecessarytoestablishthefirstcomponentofrisk.LaboratoryandfieldassaysoflethalandsublethaltoxicityresultingfromexposuretodosesofBtpollenarerequiredtoestablishtoxicitythresholdsforcomparisonagainstthedoseencounteredwithintheenvironment.ThesetoxicitythresholdswillvarybasedonexpressionlevelsforindividualBtcorneventsinconjunctionwithenvironmentalfactorsdeterminingecologicalexposure.
OnlythreepapersconcerningtheimpactofBtcornpollenonnontargetLepidopterahavebeenpublished(26,33,34)before,andtheyarelimitedintheirapplicatio