DISCUSSION Open Access No scientific consensus on GMO - PDF document

safetyofallGMOsispossibleandthattheymustbe assessedona ‘ case-by-case ’ basis.Moreover,theclaimthat itdoesexist-whichcontinuestobepushedintheabove listedcircles-ismisleadingandmisrepresentsoroutright ignoresthecurrentlyavailablescientificevidenceandthe broaddiversityofscientificopinionsamongscientistson thisissue.Theclaimfurtherencouragesaclimateof complacencythatcouldleadtoalackofregulatoryand scientificrigourandappropriatecaution,potentially endangeringthehealthofhumans,animals,andthe environment. Scienceandsocietydonotproceedonthebasisofa constructedconsensus,ascurrentknowledgeisalways opentowell-foundedchallengeanddisagreement.We endorsetheneedforfurtherindependentscientific inquiryandinformedpublicdiscussiononGMproduct safety. Someofourobjectionstotheclaimofascientific consensusarelistedinthefollowingdiscussion.The originalversionendorsedby300scientistsworldwide canbefoundatthewebsiteoftheEuropeanNetworkof ScientistsforSocialandEnvironmentalResponsibility[7]. Discussion 1 ThereisnoconsensusonGMfoodsafety RegardingthesafetyofGMcropsandfoodsforhuman andanimalhealth,acomprehensivereviewofanimal feedingstudiesofGMcropsfound ‘ Anequilibriuminthe number[of]researchgroupssuggesting,onthebasisof theirstudies,thatanumberofvarietiesofGMproducts (mainlymaizeandsoybeans)areassafeandnutritiousas therespectiveconventionalnon-GMplant,andthoserais- ingstillseriousconcerns ’ .Thereviewalsofoundthatmost studiesconcludingthatGMfoodswereassafeandnutri- tiousasthoseobtainedbyconventionalbreedingwere ‘ performedbybiotechnologycompaniesorassociates, whicharealsoresponsible[for]commercializingthese GMplants ’ [8]. Aseparatereviewofanimalfeedingstudiesthatis oftencitedasshowingthatGMfoodsaresafeincluded studiesthatfoundsignificantdifferencesintheGM-fed animals.Whilethereviewauthorsdismissedthesefind- ingsasnotbiologicallysignificant[9],theinterpretation ofthesedifferencesisthesubjectofcontinuingscientific debate[8,10-12]andnoconsensusexistsonthetopic. RigorousstudiesinvestigatingthesafetyofGMcrops andfoodswouldnormallyinvolve, interalia ,animal feedingstudiesinwhichonegroupofanimalsisfedGM foodandanothergroupisfedanequivalentnon-GM diet.Independentstudiesofthistypearerare,butwhen suchstudieshavebeenperformed,somehaverevealed toxiceffectsorsignsoftoxicityintheGM-fedanimals [2,8,11-13].Theconcernsraisedbythesestudieshave notbeenfollowedupbytargetedresearchthatcould confirmorrefutetheinitialfindings. ThelackofscientificconsensusonthesafetyofGM foodsandcropsisunderlinedbytherecentresearch callsoftheEuropeanUnionandtheFrenchgovernment toinvestigatethelong-termhealthimpactsofGMfood consumptioninthelightofuncertaintiesraisedbyani- malfeedingstudies[14,15].Theseofficialcallsimply recognitionoftheinadequacyoftherelevantexisting scientificresearchprotocols.Theycallintoquestionthe claimthatexistingresearchcanbedeemedconclusive andthescientificdebateonbiosafetyclosed. 2 Therearenoepidemiologicalstudiesinvestigating potentialeffectsofGMfoodconsumptiononhuman health Itisoftenclaimedthat ‘ trillionsofGMmeals ’ have beeneatenintheUSwithnoilleffects.However,no epidemiologicalstudiesinhumanpopulationshave beencarriedouttoestablishwhetherthereareany healtheffectsassociatedwithGMfoodconsumption. AsGMfoodsandotherproductsarenotmonitoredor labelledafterreleaseinNorthAmerica,amajorproducer andconsumerofGMcrops,itisscientificallyimpossible totrace,letalonestudy,patternsofconsumptionand theirimpacts.Therefore,claimsthatGMfoodsaresafe forhumanhealthbasedontheexperienceofNorth Americanpopulationshavenoscientificbasis. 3 Claimsthatscientificandgovernmentalbodiesendorse GMOsafetyareexaggeratedorinaccurate Claimsthatthereisaconsensusamongscientificand governmentalbodiesthatGMfoodsaresafe,orthat theyarenomoreriskythannon-GMfoods[16,17],are false.Forinstance,anexpertpaneloftheRoyalSociety ofCanadaissuedareportthatwashighlycriticalofthe regulatorysystemforGMfoodsandcropsinthatcoun- try.Thereportdeclaredthatitis ‘ scientificallyunjustifi- able ’ topresumethatGMfoodsaresafewithoutrigorous scientifictestingandthatthe ‘ defaultprediction ’ forevery GMfoodshouldbethattheintroductionofanewgene willcause ‘ unanticipatedchanges ’ intheexpressionof othergenes,thepatternofproteinsproduced,and/or metabolicactivities.Possibleoutcomesofthesechanges identifiedinthereportincludedthepresenceofnewor unexpectedallergens[18]. AreportbytheBritishMedicalAssociationcon- cludedthatwithregardtothelong-termeffectsofGM foodsonhumanhealthandtheenvironment, ‘ manyun- answeredquestionsremain ’ andthat ‘ safetyconcerns cannot,asyet,bedismissedcompletelyonthebasisof Hilbeck etal.EnvironmentalSciencesEurope (2015) 27:4 Page2of6 informationcurrentlyavailable ’ .Thereportcalledfor moreresearch,especiallyonpotentialimpactsonhuman healthandtheenvironment[19]. Moreover,thepositionstakenbyotherorganizations havefrequentlybeenhighlyqualified,acknowledging datagapsandpotentialrisks,aswellaspotentialbene- fits,ofGMtechnology.Forexample,astatementbythe AmericanMedicalAssociation ’ sCouncilonScienceand PublicHealthacknowledged ‘ asmallpotentialforadverse events … duemainlytohorizontalgenetransfer,allergen- icity,andtoxicity ’ andrecommendedthatthecurrent voluntarynotificationprocedurepractisedintheUSprior tomarketreleaseofGMcropsbemademandatory[20].It shouldbenotedthatevena ‘ smallpotentialforadverse events ’ mayturnouttobesignificant,giventhewidespread exposureofhumanandanimalpopulationstoGMcrops. AstatementbytheboardofdirectorsoftheAmerican AssociationfortheAdvancementofScience(AAAS) affirmingthesafetyofGMcropsandopposinglabelling [21]cannotbeassumedtorepresenttheviewofAAAS membersasawholeandwaschallengedinanopenletter byagroupof21scientists,includingmanylong-standing membersoftheAAAS[22].Thisepisodeunderlinedthe lackofconsensusamongscientistsaboutGMOsafety. 4 EUresearchprojectdoesnotprovidereliableevidence ofGMfoodsafety AnEUresearchproject[23]hasbeencitedinter- nationallyasprovidingevidenceforGMcropandfood safety.However,thereportbasedonthisproject, ‘ A DecadeofEU-FundedGMOResearch ’ ,presentsnodata thatcouldprovidesuchevidencefromlong-termfeeding studiesinanimals. Indeed,theprojectwasnotdesignedtotestthesafetyof anysingleGMfoodbuttofocuson ‘ thedevelopmentof safetyassessmentapproaches ’ [24].Onlyfivepublished animalfeedingstudiesarereferencedintheSAFOTEST sectionofthereport,whichisdedicatedtoGMfood safety[25].Noneofthesestudiestestedacommercialized GMfood;nonetestedtheGMfoodforlong-termeffects beyondthesubchronicperiodof90days;allfounddiffer- encesintheGM-fedanimals,whichinsomecaseswere statisticallysignificant;andnoneconcludedonthesafety oftheGMfoodtested,letaloneonthesafetyofGMfoods ingeneral.Therefore,theEUresearchprojectprovidesno evidenceforsweepingclaimsaboutthesafetyofanysingle GMfoodorofGMcropsingeneral. 5 ListofseveralhundredstudiesdoesnotshowGMfood safety AfrequentlycitedclaimpublishedonanInternetweb- sitethatseveralhundredstudies ‘ documentthegeneral safetyandnutritionalwholesomenessofGMfoodsand feeds ’ [26]ismisleading.Examinationofthestudies listedrevealsthatmanydonotprovideevidenceofGM foodsafetyand,infact,someprovideevidenceofalack ofsafety.Forexample:
Manyofthestudiesarenottoxicologicalanimal feedingstudiesofthetypethatcanprovideuseful informationabouthealtheffectsofGMfood consumption.Thelistincludesanimalproduction studiesthatexamineparametersofinteresttothe foodandagricultureindustry,suchasmilkyieldand weightgain[ 27 , 28 ];studiesonenvironmentaleffects ofGMcrops;andanalyticalstudiesofthe compositionorgeneticmakeupofthecrop.
Amongtheanimalfeedingstudiesandreviewsof suchstudiesinthelist,asubstantialnumberfound toxiceffectsandsignsoftoxicityinGM-fedanimals comparedwithcontrols[ 29 - 34 ].Concernsraisedby thesestudieshavenotbeensatisfactorilyaddressed andtheclaimthatthebodyofresearchshowsa consensusoverthesafetyofGMcropsandfoodsis falseandirresponsible.
Manyofthestudieswereconductedovershort periodscomparedwiththeanimal ’ stotallifespan andcannotdetectlong-termhealtheffects[ 35 , 36 ]. Weconcludethatthesestudies,takenasawhole,are misrepresentedontheInternetwebsiteastheydonot ‘ documentthegeneralsafetyandnutritionalwholesome- nessofGMfoodsandfeeds ’ .Rather,someofthestudies giveseriouscauseforconcernandshouldbefollowed upbymoredetailedinvestigationsoveranextended periodoftime. 6 Thereisnoconsensusontheenvironmentalrisksof GMcrops EnvironmentalrisksposedbyGMcropsincludethe effectsofinsecticidalBt(abacterialtoxinfrom Bacillus thuringiensis engineeredintocrops)cropsonnon-target organismsandtheeffectsoftheherbicidesusedintan- demwithherbicide-tolerantGMcrops. AswithGMfoodsafety,noscientificconsensusexists regardingtheenvironmentalrisksofGMcrops.Are- viewofenvironmentalriskassessmentapproachesfor GMcropsidentifiedshortcomingsintheprocedures usedandfound ‘ noconsensus ’ globallyonthemethod- ologiesthatshouldbeapplied,letaloneonstandardized testingprocedures[37].Somereviewsofthepublished dataonBtcropshavefoundthattheycanhaveadverse effectsonnon-targetandbeneficialorganisms[38-41]- effectsthatarewidelyneglectedinregulatoryassess- mentsandbysomescientificcommentators.Resistance Hilbeck etal.EnvironmentalSciencesEurope (2015) 27:4 Page3of6 toBttoxinshasemergedintargetpests[42],andprob- lemswithsecondary(non-target)pestshavebeennoted, forexample,inBtcottoninChina[43,44]. Herbicide-tolerantGMcropshaveprovedequallycon- troversial.Somereviewsandindividualstudieshaveasso- ciatedthemwithincreasedherbicideuse[45,46],therapid spreadofherbicide-resistantweeds[47],andadverse healtheffectsinhumanandanimalpopulationsexposed toRoundup,theherbicideusedonthemajorityofGM crops[48-50]. AswithGMfoodsafety,disagreementamongscien- tistsontheenvironmentalrisksofGMcropsmaybe correlatedwithfundingsources.Apeer-reviewedsurvey oftheviewsof62lifescientistsontheenvironmental risksofGMcropsfoundthatfundinganddisciplinary traininghadasignificanteffectonattitudes.Scientists withindustryfundingand/orthosetrainedinmolecular biologywereverylikelytohaveapositiveattitudeto GMcropsandtoholdthattheydonotrepresentany uniquerisks,whilepublicly-fundedscientistsworking independentlyofGMcropdevelopercompaniesand/or thosetrainedinecologyweremorelikelytoholda ‘ moderatelynegative ’ attitudetoGMcropsafetyandto emphasizetheuncertaintyandignoranceinvolved.The reviewauthorsconcluded ‘ Thestrongeffectsoftrain- ingandfundingmightjustifycertaininstitutionalchanges concerninghowweorganizescienceandhowwe makepublicdecisionswhennewtechnologiesareto beevaluated ’ [51]. 7 Internationalagreementsshowwidespreadrecognition ofrisksposedbyGMfoodsandcrops TheCartagenaProtocolonBiosafetywasnegotiated overmanyyearsandimplementedin2003.TheCartagena Protocolisaninternationalagreementratifiedby166gov- ernmentsworldwidethatseekstoprotectbiologicaldiver- sityfromtherisksposedbyGMtechnology.Itembodies thePrecautionaryPrincipleinthatitallowssignatory statestotakeprecautionarymeasurestoprotectthem- selvesagainstthreatsofdamagefromGMcropsand foods,evenincaseofalackofscientificcertainty[52]. Anotherinternationalbody,theUN ’ sCodexAlimentar- ius,workedwithscientificexpertsfor7yearstodevelop internationalguidelinesfortheassessmentofGMfoods andcropsbecauseofconcernsabouttheriskstheypose. TheseguidelineswereadoptedbytheCodexAlimentarius Commission,ofwhichover160nationsaremembers, includingmajorGMcropproducerssuchastheUnited States[53]. TheCartagenaProtocolandCodexshareaprecau- tionaryapproachtoGMcropsandfoods,inthatthey agreethatgeneticengineeringdiffersfromconventional breedingandthatsafetyassessmentsshouldberequired beforeGMorganismsareusedinfoodorreleasedinto theenvironment. Theseagreementswouldneverhavebeennegotiated, andtheimplementationprocesseselaboratinghowsuch safetyassessmentsshouldbeconductedwouldnotcur- rentlybehappening,withoutwidespreadinternational recognitionoftherisksposedbyGMcropsandfoods andtheunresolvedstateofexistingscientificunder- standing.Concernsaboutrisksarewellfounded,ashas beendemonstratedbystudiesonsomeGMcropsand foodsthathaveshownadverseeffectsonanimalhealth andnon-targetorganisms,indicatedabove.Manyofthese studieshave,infact,fedintothenegotiationand/orimple- mentationprocessesoftheCartagenaProtocolandthe Codex.WesupporttheapplicationofthePrecautionary Principlewithregardtothereleaseandtransboundary movementofGMcropsandfoods. Conclusions Inthescopeofthisdocument,wecanonlyhighlightafew examplestoillustratethatthetotalityofscientificresearch outcomesinthefieldofGMcropsafetyisnuanced;com- plex;oftencontradictoryorinconclusive;confoundedby researchers ’ choices,assumptions,andfundingsources; and,ingeneral,hasraisedmorequestionsthanithas currentlyanswered. Whethertocontinueandexpandtheintroductionof GMcropsandfoodsintothehumanfoodandanimal feedsupply,andwhethertheidentifiedrisksareaccept- ableornot,aredecisionsthatinvolvesocioeconomic considerationsbeyondthescopeofanarrowscientific debateandthecurrentlyunresolvedbiosafetyresearch agendas.Thesedecisionsmustthereforeinvolvethe broadersociety.Theyshould,however,besupportedby strongscientificevidenceonthelong-termsafetyofGM cropsandfoodsforhumanandanimalhealthandthe environment,obtainedinamannerthatishonest,eth- ical,rigorous,independent,transparent,andsufficiently diversifiedtocompensateforbias. Decisionsonthefutureofourfoodandagriculture shouldnotbebasedonmisleadingandmisrepresentative claimsbyaninternalcircleoflikemindedstakeholders thata ‘ scientificconsensus ’ existsonGMOsafety. Thisdocumentwassubsequentlyopenedforendorse- mentbyscientistsfromaroundtheworldintheirper- sonal(ratherthaninstitut ional)capacitiesreflecting theirpersonalviewsandbasedontheirpersonalexpertise. Thereisnosuggestionthattheviewsexpressedinthis statementrepresenttheviewsorpositionofanyinstitu- tionororganizationwithwhichtheindividualsareaffili- ated.Qualifyingcriteriaforsigningthestatementwere deliberatelyselectedtoincludescientists,physicians,so- cialscientists,academics,andspecialistsinlegalaspects andriskassessmentofGMcropsandfoods.Scientistand Hilbeck etal.EnvironmentalSciencesEurope (2015) 27:4 Page4of6 academicsignatorieswererequestedtohavequalifications fromaccreditedinstitutionsatthelevelofPhDorequiva- lent.LegalexpertswererequestedtohaveatleastaJDor equivalent.ByDecember2013,morethan300peoplewho metthestrictqualificationrequirementshadsignedthe statement.Thestatementwaswidelytakenupinthe mediaandreportedinnumerousoutletsandevidence providedthereincontinuestobecitedwidely.Inatime whenthereismajorpressureonthesciencecommunity fromcorporateandpoliticalinterests,itisofutmostim- portancethatscientistsworkingforthepublicinterest takeastandagainstattemptstoreduceandcompromise therigourofexaminationofnewapplicationsinfavorof rapidcommercializationofnewandemergingtechnolo- giesthatareexpectedtogenerateprofitandeconomic growth.Thedocumentcontinuestobeopenforsignature onthewebsiteoftheinitiatingscientificorganization ENSSER(EuropeanNetworkofScientistsforSocialand EnvironmentalResponsibility)atwww.ensser.org. Competinginterests Theauthorsdeclarethattheyhavenocompetinginterests. Authors ’ contributions Allauthorscontributedequallytothewritingofthedocument.Allauthors readandapprovedthefinalmanuscript. Authors ’ information EACisretired. Authordetails 1 EuropeanNetworkofScientistsforSocialandEnvironmentalResponsibility (ENSSERBoard/Secretariat),Marienstrasse19/20,10117Berlin,Germany. 2 InstituteofIntegrativeBiology,SwissFederalInstituteofTechnologyZurich, Universitätstrasse16,8092Zurich,Switzerland. 3 GenØkCentreforBiosafety, Forskningsparken,PB6418,9294Tromsø,Norway. 4 ComitédeRechercheet d ’ InformationIndépendantessurlegénieGénétiqueCRIIGEN,40rue Monceau,75008Paris,France. 5 UniversityofCaen,InstituteofBiologyIBFA andNetworkonRisks,QualityandSustainableEnvironmentMRSH, EsplanadedelaPaix,14032Caen,France. 6 EcoNexus,OxfordOX49BS,UK. 7 CentralEnvironmentalandFoodScienceResearchInstitute,POB393, H-1537Budapest,Hungary. 8 DepartmentofMedicalandMolecularGenetics, FacultyofLifeSciencesandMedicine,King ’ sCollegeLondon,Guy ’ sHospital, LondonSE19RT,UK. 9 TechnologyandPublicPolicy,Universityof Washington,Seattle,Washington98195,USA. 10 PlantAgriculture,University ofGuelph,K0K3K0Warkworth,Ontario,Canada. 11 ConsumersUnion,101 TrumanAvenue,Yonkers,NY10703,USA. 12 ClareHall,Universityof Cambridge,HerschelRoad,CambridgeCB39AL,UK. 13 CentreforIntegrated ResearchinBiosafety,SchoolofBiologicalSciences,UniversityofCanterbury, Christchurch,NewZealand. 14 NavdanyaHauzKhas,NewDelhi110016,India. 15 CentrefortheStudyofEnvironmentalChange,CSEC,LancasterUniversity, Lancaster,UK. 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WorldHealthOrganization/FoodandAgricultureOrganizationoftheUnited Nations2000[ftp://ftp.fao.org/codex/Publications/Booklets/Biotech/ Biotech_2009e.pdf] Submit your manuscript to a journal and bene“ t from: 7 Convenient online submission 7 Rigorous peer review 7 Immediate publication on acceptance 7 Open access: articles freely available online 7 High visibility within the “ eld 7 Retaining the copyright to your article Submit your next manuscript at 7 springeropen.com Hilbeck etal.EnvironmentalSciencesEurope (2015) 27:4 Page6of6 DISCUSSIONOpenAccess NoscientificconsensusonGMOsafety AngelikaHilbeck 1,2* ,RosaBinimelis 1,3 ,NicolasDefarge 1,4,5 ,RicardaSteinbrecher 1,6 ,AndrásSzékács 1,7 , FernWickson 1,3 ,MichaelAntoniou 8 ,PhilipLBereano 9 ,EthelAnnClark 10 ,MichaelHansen 11 ,EvaNovotny 12 , JackHeinemann 13 ,HartmutMeyer 1 ,VandanaShiva 14 andBrianWynne 15 Abstract Abroadcommunityofindependentscientificresearchersandscholarschallengesrecentclaimsofaconsensus overthesafetyofgeneticallymodifiedorganisms(GMOs).Inthefollowingjointstatement,theclaimedconsensus isshowntobeanartificialconstructthathasbeenfalselyperpetuatedthroughdiversefora.Irrespectiveof contradictoryevidenceintherefereedliterature,asdocumentedbelow,theclaimthatthereisnowaconsensuson thesafetyofGMOscontinuestobewidelyandoftenuncriticallyaired.Fordecades,thesafetyofGMOshasbeena hotlycontroversialtopicthathasbeenmuchdebatedaroundtheworld.Publishedresultsarecontradictory,inpart duetotherangeofdifferentresearchmethodsemployed,aninadequacyofavailableprocedures,anddifferences intheanalysisandinterpretationofdata.Suchalackofconsensusonsafetyisalsoevidencedbytheagreementof policymakersfromover160countries-intheUN ’ sCartagenaBiosafetyProtocolandtheGuidelinesofthe Codex Alimentarius -toauthorizecarefulcase-by-caseassessmentofeachGMObynationalauthoritiestodetermine whethertheparticularconstructsatisfiesthenationalcriteriafor ‘ safe .RigorousassessmentofGMOsafetyhasbeen hamperedbythelackoffundingindependentofproprietaryinterests.Researchforthepublicgoodhasbeen furtherconstrainedbypropertyrightsissues,andbydenialofaccesstoresearchmaterialforresearchersunwilling tosigncontractualagreementswiththedevelopers,whichconferunacceptablecontroloverpublicationtothe proprietaryinterests. Thejointstatementdevelopedandsignedbyover300independentresearchers,andreproducedandpublished below,doesnotassertthatGMOsareunsafeorsafe.Rather,thestatementconcludesthatthescarcityand contradictorynatureofthescientificevidencepublishedtodatepreventsconclusiveclaimsofsafety,oroflackof safety,ofGMOs.ClaimsofconsensusonthesafetyofGMOsarenotsupportedbyanobjectiveanalysisofthe refereedliterature. Background Overrecentyears,anumberofscientificresearcharticles havebeenpublishedthatreportdisturbingresultsfrom geneticallymodifiedorganism(GMO)feedingexperi- mentswithdifferentmammals(e.g.rats[1],pigs[2]).In additiontotheusualfierceresponses,thesehaveeliciteda concertedeffortbygeneticallymodified(GM)seeddevel- opersandsomescientists,commentators,andjournalists toconstructclaimsthatthereisa ‘ scientificconsensus ’ onGMOsafety[3-5]andthatthedebateonthistopic is ‘ over ’ [6]. Theseclaimsledabroaderindependentcommunityof scientistsandresearcherstocometogetherastheyfelt accountofthecurrentstateofdissentinthisfield,based onpublishedevidenceinthescientificliterature,forboth theinterestedpublicandthewidersciencecommunity. Thestatementthatwasdevelopedwasthenopenedupfor endorsementfromscientistsaroundtheworldwithrele- vantexpertiseandcapacitiestoconcludeonthecurrent stateofconsensus/dissentanddebateregardingthepub- lishedevidenceonthesafetyofGMOs. Thisstatementclearlydemonstratesthattheclaimed consensusonGMOsafetydoesnotexistoutsideofthe abovedepictedinternalcircleofstakeholders.Thehealth, environment,andagricultureauthoritiesofmostnations recognizepubliclythatnoblanketstatementaboutthe *Correspondence: angelika.hilbeck@ensser.org 1 EuropeanNetworkofScientistsforSocialandEnvironmentalResponsibility (ENSSERBoard/Secretariat),Marienstrasse19/20,10117Berlin,Germany 2 InstituteofIntegrativeBiology,SwissFederalInstituteofTechnologyZurich, Universitätstrasse16,8092Zurich,Switzerland Fulllistofauthorinformationisavailableattheendofthearticle ©2015Hilbecketal.;licenseeSpringer.ThisisanOpenAccessarticledistributedunderthetermsoftheCreativeCommons AttributionLicense(http://creativecommons.org/licenses/by/4.0),whichpermitsunrestricteduse,distribution,andreproduction inanymedium,providedtheoriginalworkisproperlycredited. Hilbeck etal.EnvironmentalSciencesEurope (2015) 27:4 DOI10.1186/s12302-014-0034-1