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Transgenic Crop Plants: Principles and Development PDF

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Transgenic Crop Plants Chittaranjan Kole Charles H. Michler l l Albert G. Abbott Timothy C. Hall l Editors Transgenic Crop Plants Volume 1: Principles and Development Editors Prof.ChittaranjanKole Prof.AlbertG.Abbott DepartmentofGenetics&Biochemistry DepartmentofGenetics&Biochemistry ClemsonUniversity ClemsonUniversity Clemson,SC29634,USA Clemson,SC29634,USA [email protected] [email protected] Prof.CharlesH.Michler Prof.TimothyC.Hall Director InstituteofDevelopmental&Molecular HardwoodTreeImprovementand Biology RegenerationCenteratPurdueUniversity DepartmentofBiology NSFI/UCRCCenterforTreeGenetics TexasA&MUniversity WestLafayette,IN47907,USA CollegeStation,TX77843,USA [email protected] [email protected] ISBN:978-3-642-04808-1 e-ISBN:978-3-642-04809-8 DOI10.1007/978-3-642-04809-8 SpringerHeidelbergDordrechtLondonNewYork LibraryofCongressControlNumber:2009939123 #Springer-VerlagBerlinHeidelberg2010 Thisworkissubjecttocopyright.Allrightsarereserved,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting, reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublication orpartsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9, 1965,initscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violations areliabletoprosecutionundertheGermanCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnot imply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotec- tivelawsandregulationsandthereforefreeforgeneraluse. Coverdesign:WMXDesignGmbH,Heidelberg,Germany Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface TransgenicPlants–knownalsoasBiotechPlants, GeneticallyEngineeredPlants, orGeneticallyModifiedPlants–haveemergedamazinglyfastasaboonforscience andsociety.Theyhavealreadyplayedandwillcontinuetoplayasignificantrolein agriculture,medicine,ecology,andenvironment.Theincreasingdemandsforfood, feed,fuel,fiber,furniture,perfumes,minerals,vitamins,antibiotics,narcotics,and many health-related drugs and chemicals necessitate development and cultivation oftransgenicplantswithaugmentedorsuppressedtrait(s).Fromasingletransgenic plant(FlavrSavrTM tomatowithalongershelf-life)introducedforcommercializa- tionin1994,wehavenow13transgeniccropscovering800millionhectaresin25 countries of six continents. Interestingly, the 13.3 million farmers growing trans- genic crops globally include 12.3 million (90%) small and resource-poor farmers from 12 developing countries. Increasing popularity of transgenic plants is well evidencedfromanannualincreaseofabout10%measuredinhectaresbutactually of15%in“traithectares.”Consideringtheurgentrequirementoftransgenicplants and wide acceptance by the farmers, research on transgenic plants is now being conducted on 57 crops in 63 countries. Transgenic plants have been developed in over100plantspeciesandtheyaregoingtocoverthefields,orchards,plantations, forests, and even the seas in the near future. These plantshave been tailored with incorporationofusefulaliengenesforseveraldesirabletraitsincludingmanywith “stackedtraits”andalsowithsilencingofgenescontrollingsomeundesirabletraits. Development,applications,andsocio-politicalimplicationsoftransgenicplants are immensely important fields now in education, research, and industries. Plant transgenicshasdeservedlybeenincludedinthecoursecurriculainmost,ifnotall, leading universities and academic institutes all over the world, and therefore reference books on transgenic plants with a classroom approach are essential for teaching,research,andextension.Therearesomeelegantreviewsonthetransgenic plantsorplantgroups(includinga10-volumeseries“CompendiumofTransgenic Crop Plants” edited by two of the present team of editors C. Kole and T.C. Hall published by Wiley-Blackwell in 2008) and on many individual tools and techni- quesofgenetictransformationinplants.Allthesereviewscouldsurelyservewell v vi Preface thepurposeforindividualcropplantsorparticularmethodologies.Sincetransgenic plant development and utilization is studied, taught, and practiced by students, teachers,andscientistsofoveradozendisciplinesunderbasicscience,agriculture, medicine, and humanities at public and private sectors, introductory reference books with lucid deliberations on the concepts, tools, and strategies to develop and utilize transgenic plants and their global impacts could be highly useful for a broadsectionofreaders. Deployment of transgenic crop plants are discussed, debated, regulated, and sponsored by people of diverse layers of the society including social activists, policymakers,andthestaffofregulatoryandfundingagencies.Theyalsorequire lucid deliberations on the deployment, regulations, and legal implications of practicingplanttransgenics.Moreimportantly,depictionofthepositiveandrealis- tic picture of the transgenic plants should and could facilitate mitigation of the negative propaganda against transgenic plants and thereby reinforce moral and financialsupportfromallindividualsandplatformsofthesociety.Globalpopula- tionisincreasingannuallyby70millionandisestimatedtogrowupto8billionby 2025. This huge population, particularly its large section from the developing countries, will suffer because of hunger, malnutrition, and chemical pollution unlessweproducemoreandmoretransgenicplantsparticularlywithstackedtraits. Compulsiontomeet therequirementsofthisgrowingpopulationonearth andthe proven innocuous nature of transgenic plants tested and testified for the last 13yearscouldsubstantiatetheimperativenecessityofembracingtransgenics. Traditional and molecular breeding practiced over the last century provided enormous number of improved varieties in economic crops and trees including wheat andrice varietiesthatfosteredthe“green revolution.”However,thesecrop improvement tools depend solely on the desirable genes available naturally, crea- table by mutation in a particular economic species or their shuffling for desired recombinations.Transgenicbreedingopenedanovelavenuetoincorporateuseful aliengenesnotonlyfromothercross-incompatiblespeciesandgeneraoftheplant kingdom,butalsofrommembersoftheprokaryotesincludingbacteria,fungi,and viruses, and even from higher animals including mice and humans. An array of plantgenetic engineering achievements starting from developinginsect resistance transformingwiththecrygenesincottonfromthebacteriaBacillusthuringiensisto thepresent-daymolecularpharmingexpressingtheinterferon-agenefromhuman intobaccoevidenceforthispan-specificgenetransfer. Humanandanimal safetyisanothergeneralconcernrelatedtotransgenicfood or feed. However, there is no reliable scientific documentation of these health hazardsevenafter13yearsofcultivationoftransgenicplantsandconsumptionof aboutonetrillionmealscontainingtransgenicingredients.Utilizationoftransgenic plantshasreducedthepesticideapplicationsby359,000tonsthatwouldotherwise affect human and animal health besides causing air, water, and soil pollution and alsomitigatedthechanceofconsumptionofdeadmicrobesandinsectsalongwith foodsorfeeds. Geneflowfromtransgeniccropspeciestotheircross-compatiblewildrelatives isagenuineconcernandthereforerequiredtestingofatransgeniccropplantbefore Preface vii deployment followed by a comprehensive survey of the area for the presence of interfertilewildandweedyplantsbeforeintroductionofatransgeniccropisbeing seriouslyconducted. Additionofnovelgenotypeswithtransgenesinthegermplasmsisincreasingthe biologicaldiversityratherthandepletingit.Useofthegeneticallyengineeredplants hasalsoeliminatedgreenhousegasemissionof10millionmetrictonsthroughfuel savings.Infact,1.8billionlitersofdieselhasbeensavedbecauseofreducedtillage andplowingowingonlytoherbicide-resistanttransgeniccrops.Manytransgenics arenowbeingusedforsoilreclamation.Aboveall,cultivationoftransgeniccrops hasreturned$44billionofnetincometothefarmers.Perhapsthesearethereasons that 25 Nobel Laureates and 3,000-plus eminent scientists appreciated the merits and safety and also endorsed transgenic crops as a powerful and safe way to improve agriculture and environment besides the safety of genetically modified foods.Manyinternationalandnationalorganizationshavealsoendorsedhealthand environmental safety of transgenic plants; these include Royal Society (UK), National Academy of Sciences (USA), World Health Organization, Food and AgricultureOrganization(UN),EuropeanCommission,FrenchAcademyofMedi- cine,andAmericanMedicalAssociation,tonameafew. Production,contributions,andsocio-politicalimplicationsofbiotechplantsare naturally important disciplines now in education, research, and industries and therefore introductory reference books are required for students, scientists, indus- tries, and also for social activists and policy makers. The two book volumes on “TransgenicCropPlants”willhopefullyfillthisgap.Thesetwobookvolumeshave severaluniquefeaturesthatdeservemention.Theoutlinesofthechaptersforthese twobooksareformulatedtoaddresstherequirementsofabroadsectionofreaders. Students and scholars of all levels will obtain a lot of valuable reading material required for their courses and researches. Scientists will get information on con- cepts, strategies,and clues usefulfor their researches. Seed companiesand indus- tries will get information on potential resources of plant materials, and expertise, andalsofortheirownR&Dactivities.Inbrief,thecontentsofthisserieshavebeen designedtofulfillthedemandsofstudents,teachers,scientistsandindustrypeople, for small to large libraries. Students, faculties, or scientists involved in various subjectswillbebenefitedfromthisseries:biotechnology,bioinformatics,molecu- lar biology, molecular genetics, plant breeding, biochemistry, ecology, environ- mental science, bioengineering, chemical engineering, genetic engineering, biomedical engineering, pharmaceutical science,agronomy, horticulture, forestry, entomology,pathology,nematology,andvirology,justtonameafew. Ithasbeenourprivilegetoeditthe23chaptersofthesetwobooks,contributed by 71 scientists from 14 countries, and the list of authors includes one of the pioneers of plant transgenics Prof. Timothy C. Hall (one of the editors also); some senior scientists who have themselves edited books on plant transgenics; and many scientists who have written elegant reviews on invitation for quality books and leading journals. We believe that these two books will hopefully serve the purposes of the broad audience: those who are studying, teaching, practicing, supporting, funding, and also those who are debating for or against plant viii Preface transgenics.Thefirstvolumededicatedto“PrinciplesandDevelopment”elucidates the basic concepts, tools, strategies, and methodologies of genetic engineering, while the second volume “Applications and Safety” enumerates the utilization of transgenic crop plants for various purposes of agriculture, industry, ecology and environment, and also genomics research. The second volume also deliberates comprehensivelyonthelegalandregulatoryaspects;addressesthemajorconcerns; andfinallyjustifiesthecompulsionofpracticingplanttransgenics. Little more detail on the contents of the volume “Transgenic Crop Plants: Principles and Development” will hopefully substantiate its usefulness. This vol- umefocusesonthemethodsforconstructinggenevectors,introducingthesegene vectors into plant tissue, targeting gene insertion to specific tissues, methods for detectingtransgene expression, generationoftransgenicplants,andtypesoftraits and bioproducts that are targeted for these technologies. The first chapter of this volumepresentsglimpsesontheseaspectsandalsoonthoserelatedtodeployment oftransgenicplants. Oneimportantfactorthatdeterminessuccessfultransgeneinsertionisthedeci- sionofexplanttypeforuseintransformation.AcomprehensivereviewinChap.2is providedfrompreviousresearchwithbothherbaceousandwoodyplants.Theuse of morphogenic calli for cereals is discussed along with somewhat standardized protocols for each individual woody species. Gene transfer methods have been discussed including use of Agrobacterium, biolistics, electroporation, liposomes, microinjections,andbioactivebeadsinChap.3. Onceatransgeneisinserted,markershavebeenusedtoeitherscorethesuccess ofthetransgeneeventorscreenforthesuccessfulevents.Althoughmuchworkhas beendoneusingnptandgus,recentworkhaslookedatmarkergeneremovalinthe finaltransgeneproductinordertobelieenvironmentalconcerns.Furthermolecular characterizationwithSouthernblotanalysisandPCRconfirmdefinitivetransgene integrationandcopynumber.Chapter4hasbeendevotedtothesecriticalsteps. Stableandregulatedtransgeneexpression,asdescribedinChap.5,isnecessary for the transgenic plant to express the trait of interest for further research or commercial applications. The use of constitutive and tissue-enhanced promoters alongwithattentiontoattachmentregionswithintheDNA,introns,RNAintegrity factors,andtranscriptionfactorswilldeterminetransgeneexpressionandthelevels thereof. Besides using transgenes for introduction of nucleic acid for novel trait production,transgeneshavealsobeenusedtosilencenativegenesforapplications suchasresistancetonematodes,insects,andviruses.Asinterestingaretheapplica- tionstoreduceproductionofcompoundsinsomeplants,suchascaffeineincoffee and sulfur metabolites in onions, that are disagreeable to portions of the human populations. All of these silencing events rely on RNAi technology to degrade native RNA for those traits of interest. Chapter 6 provides a commentary on the employmentofRNAitechnologyandtheimplicationsandoutcomeofexpression andgenesilencinghavebeenexplicatedinChap.7. Researchershavealsobeensuccessfulwithorganelletransfer,whichhasappli- cationstomolecularpharming,ashavebeenenumeratedinChap.8.Thistechnique canbeemployedtoovercomesometransgeneexpressiondifficulties.Cellculture Preface ix biosynthesis and metabolic engineering are the focus of the last two chapters (Chaps.9 and 10) in this volume and these chapters offer an intriguing look at research into production of high-value energy and medicinal products, secondary metabolites, and plants with attractive esthetic qualities. Because we still have a rudimentary understanding of many biochemical pathways, we are continuing to gain new knowledge and insight into pathway function, but commercial plant systemsarestilllackinginmostdesirabletraitswheneconomicviability,environ- mentalsafety,andsustainabilityaretakenintoaccount. Wethankthe31scientistsfrom9countriesfortheirelegantandlucidcontribu- tionstothisvolumeandalsofortheirsustainedsupportthroughrevising,updating, and fine-tuning their chapters. We also acknowledge the recent statistics we have accessedfromthewebsitesofMonsantoCompanyon“ConversationsaboutPlant Biotechnology” and “International Service for the Acquisition of Agri-Biotech Applications on ISAAA Brief 39-2008: Executive Summary” and used them in thisprefaceandelsewhereinthevolume. We enjoyed a lot our Clemson-Purdue-Texas A&M triangular interaction, constant consultations, and dialogs while editing this book and also working with the editorial staff of Springer, particularly Dr. Sabine Schwarz, who had been supportivesincetheinceptiontillthepublicationofthisbook. Welookforwardtosuggestionsfromallcornersforthefutureimprovementof thecontentandapproachofthisbookvolume. ChittaranjanKole,Clemson,SC CharlesH.Michler,WestLafayette,IN AlbertG.Abbott,Clemson,SC TimothyC.Hall,CollegeStation,TX

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