Wild Crop Relatives: Genomic and Breeding Resources . Chittaranjan Kole Editor Wild Crop Relatives: Genomic and Breeding Resources Cereals Editor Prof.ChittaranjanKole DirectorofResearch InstituteofNutraceuticalResearch ClemsonUniversity 109JordanHall Clemson,SC29634 [email protected] ISBN978-3-642-14227-7 e-ISBN978-3-642-14228-4 DOI10.1007/978-3-642-14228-4 SpringerHeidelbergDordrechtLondonNewYork LibraryofCongressControlNumber:2011922649 # Springer-VerlagBerlinHeidelberg2011 Thisworkissubjecttocopyright.Allrightsarereserved,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting, reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublicationor partsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9,1965,in itscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violationsareliableto prosecutionundertheGermanCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnotimply, evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotectivelawsand regulationsandthereforefreeforgeneraluse. Coverdesign:deblik,Berlin Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Dedication Dr.NormanErnestBorlaug,1 the Father of Green Revolution, is well respected for his contri- butions to science and society. There was or is not and never will be a single person on this Earth whose single-handed ser- vice to science could save millions of people from death due to starvation over a period of over four decades like Dr. Borlaug’s.EventheNobelPeace Prize he received in 1970 does not do such a great and noble person as Dr. Borlaug justice. His life and contributions are well known and will remain in the pages of history of science. I wish here only to share some facets of this elegant and ideal personality I had been blessed to observeduringmypersonalinter- actionswithhim. Itwasearly2007whileIwas at the Clemson University as a visiting scientist one of my lab colleagues told me that “some- body wants to talk to you; he appears to be an old man”. I took the telephone receiver casually and said hello. The response from the other side was – “I am Norman Borlaug; am I talking to Chitta?” Even a million words would be insufficient to define and depict the exact feelingsandthrillsIexperiencedatthatmoment! 1ThephotoofDr.BorlaugwaskindlyprovidedbyJulieBorlaug(NormanBorlaugInstituteforInternationalAgriculture,Texas A&MAgriculture)thegranddaughterofDr.Borlaug. v vi Dedication IhadseenDr.Borlaugonlyonce,waybackin1983,whenhecametoNewDelhi, IndiatodelivertheCoromandalLectureorganizedbyProf.M.S.Swaminathanonthe occasion of the 15th International Genetic Congress. However, my real interaction withhimbeganin2004whenIhadbeenformulatinga7-volumebookseriesentitled GenomeMappingandMolecularBreedinginPlants.Initially,Iwasneitherconfident of my ability as a series/book editor nor of the quality of the contents of the book volumes. I sent an email to Dr. Borlaug attaching the table of contents and the tentativeoutlineofthechaptersalongwithmanuscriptsofonlyafewsamplechapters, including one authored by me and others, to learn about his views as a source of inspiration(orcaution!)Iwasalmostsurethatapersonofhisstaturewouldhaveno time and purpose to get back to a small science worker like me. To my utter (and pleasant) surprise I received an email from him that read: “May all Ph.D.’s, future scientists,andstudentsthataredevotedtoagriculturegetaninspirationasitrefersto your work or future work from the pages of this important book. My wholehearted wishesforasuccessonyourimportantjob”.Igotashotinmyarm(andinmindfor sure)!Restisapleasantexperience–thesevenvolumeswerepublishedbySpringerin 2006and2007,andwerewelcomeandlikedbystudents,scientistsandtheirsocieties, libraries, and industries. As a token of my humble regards and gratitude, I sent Dr. Borlaug the Volume I on Cereals and Millets that was published in 2006. And here started my discovery of the simplest person on Earth who solved the most complexandcriticalproblemofpeopleonit–hungeranddeath. Just one month after receiving the volume, Dr. Borlaug called me one day and said,“Chitta,youknowIcannotreadalotnow-a-days,butIhavegonethroughonly on the chapters on wheat, maize and rice. Please excuse me. Other chapters of this andothervolumesoftheserieswillbeequallyexcellent,Ibelieve”.Hewashighly excitedtoknowthatmanyotherNobelLaureatesincludingProfs.ArthurKornberg, Werner Arber, Phillip Sharp, G€unter Blobel, and Lee Hartwell also expressed generous comments regarding the utility and impact of the book series on science and the academic society. While we were discussing many other textbooks and review book series that I was editing at that time, again in my night hours for the benefit of students, scientists, and industries, he became emotional and said to me, “Chitta, forget about your original contributions to basic and applied sciences, you deservedNobelPrizeforPeacelikemeforprovidingacademicfoodstomillionsof starving students and scientists over the world particularly in the developing countries.IwillrecommendyournamefortheWorldFoodPrize,butitwillnotdo enoughjusticetothesacrificeyouaredoingforscienceandsocietyinyoursleepless nightsoversomanyyears.TakesomerestChittaandgivetimetoPhullara,Sourav andDevleena”(hewassoparticulartoaskaboutmywifeandourkidsduringmostof our conversations). I felt honored but really very ashamed as I am aware of my almostinsignificantcontributionincomparisontohismonumentalcontributionand thousands of scientists over the world are doing at least hundred-times better jobs thanmeasscientistorauthor/editorofbooks!So,Iwasunabletoutteranywordsfor acoupleofminutesbutrealizedlaterthathemustbeentooaffectionatetomeandhis hugeaffectionisthebestawardforasmallscienceworkerasme! In another occasion he wanted some documents from me. I told him that I will send them as attachments in emails. Immediately he shouted and told me: “Youknow,Julie(hisgranddaughter)isnotathomenowandIcannotcheckemail myself.Juliedoesthisforme.IcantypemyselfintypewriterbutIamnotgoodin computer.Youknowwhat,Ihaveaxeroxmachineanditreceivesfaxalso.Sendme Dedication vii thedocumentsbyfax”.Herewastheever-presentchildinhim.Julieemailedmelater tosendthedocumentsasattachmenttoherasthe‘xeroxmachine’ofDr.Borlaugran outofink! AnotheroccasioniswhenIwastalkingwithhiminalowvoice,andheimmedi- atelychidedme:“YouknowthatIcannothearwellnow-a-days;Idon’tknowwhere Juliehaskeptthehearingapparatus,can’tyouspeaklouder?”Herewasthefatherly figurewhowaseagertoheareachofmywords! I still shed tears when I remember during one of our telephone conversations he asked:“YouknowIhaveneverseenyou,canyoucometoDallasinthenearfutureby chance?”IrememberweweregoingthroughafinancialpaucityatthattimeandIcould notmakeavisittoDallas(Texas)toseehim,thoughitwouldhavebeenagreathonor. In late 2007, whenever I tried to talk to Dr. Borlaug, he used to beckon Julie to bring the telephone to him, and in course of time Julie used to keep alive all the communicationsbetweenuswhenheslowlysuccumbedtohishealthproblems. The remaining volumes of the Genome Mapping and Molecular Breeding in Plantsserieswerepublishedin2007,andIsenthimallthesevenvolumes.Iwished tolearnabouthisviews.Duringthisperiodhecouldnotspeakandwritewell.Julie preparedaletterbasedonhiswordstoherthatread:“DearChitta,Ihavereviewed the seven volumes of the series on Genome Mapping and Molecular Breeding in Plants, which you have authored. You have brought together genetic linkage maps based on molecular markers for the most important crop species that will be a valuable guide and tool to further molecular crop improvements. Congratulations forajobwelldone”. During one of our conversations in mid-2007, he asked me what other book projects I was planning for Ph.D. students and scientists (who had always been his all-timebelovedfolks).Itoldhimthatthewealthofwildspeciesalreadyutilizedand to be utilized for genetic analysis and improvement of domesticated crop species have not been deliberated in any book project. He was very excited and told me to take up the book project as soon as possible. But during that period I had a huge commitment to editing a number of book volumes and could not start the series he wassointerestedabout. HissuddendemiseinSeptember2009keptmesomoroseforanumberofmonths thatIcould notevencommunicatemypersonallosstoJulie.Butinthemeantime,I formulated a 10-volume series on Wild Crop Relatives: Genomic and Breeding ResourcesforSpringer.AndwhomelsetodedicatethisseriestootherthanDr.Borlaug! IwrotetoJulieforherformalpermissionandsheimmediatelywroteme:“Chitta, ThankyouforcontactingmeandyesIthinkmygrandfatherwouldbehonoredwith thededicationoftheseries.Irememberhimtalkingofyouandthisundertakingquite often.Congratulations onallthatyouhave accomplished!”Thishelpedme alotas I could at least feel consoled that I could do a job he wanted me to do and I will alwaysremaingratefultoJulieforthishelpandalsofortakingcareofDr.Borlaug, notonlyashisgranddaughterbutalsoastherepresentativeofmillionsofpoorpeople from around the world and hundreds of plant and agricultural scientists who try to followhisphilosophyandworshiphimasafatherfigure. Itisanothersadexperienceofgrowingolderinlifethatwewalkaloneandmiss the affectionate shadows, inspirations, encouragements, and blessings from the fatherlyfiguresinourprofessional andpersonal lives. How Iwish Icould treatmy nextgenerationsinthesamewayaspersonalitieslikeMotherTeresaandDr.Norman Borlaugandmanyothergreatpeoplefromaroundtheworldtreatedme! viii Dedication Duringmostofourconversationsheusedtoemphasizeontheimmediateimpact ofresearchonthesocietyanditspeople.Acoupleoftimesheeventoldmethatmy works on molecular genetics and biotechnology, particularly of 1980s and 1990s, havehighfundamentalimportance,butIshouldalsodosomeworksthatwillbenefit peopleimmediately.Thisadviceelicitedachangeinmythoughtsandworkplansand sincethenIhavebeendevotedlyendeavoringtodevelopcropvarietiesenrichedwith phytomedicines and nutraceuticals. Borlaug influenced both my personal and pro- fessionallife,particularlymyapproachtoscience,andIdedicatethisseriestohimin remembranceofhisgreatcontributiontoscienceandsocietyandforallhispersonal affection,loveandblessingsforme. IemailedtheabovedraftofthededicationpagetoJulieforherviewsandIwishto completemyhumblededicationwithgreatsatisfactionwiththewordsofJuliewho served as the living ladder for me to reach and stay closer to such as great human beingasDr.Borlaugandexpressmydeepregardsandgratitudetoher.Julie’semail read:“Chitta,Thankyouforsendingmethedraftdedicationpage.Ireallyenjoyed readingitandIthinkyoucapturedmygrandfather’sspiritwonderfully....Sothank youverymuchforyourbeautifulwords.Iknowhewouldbeandishonored.” Clemson,USA ChittaranjanKole Preface Wild crop relatives have been playing enormously important roles both in the depictionofplantgenomesandthegeneticimprovementoftheircultivatedcounter- parts.Theyhavecontributedimmenselytoresolvingseveralfundamentalquestions, particularlythoserelatedtotheorigin,evolution,phylogeneticrelationship,cytolog- ical status and inheritance of genes of an array of crop plants; provided several desirable donor genes for the genetic improvement of their domesticated counter- parts;andfacilitatedtheinnovationofmanynovelconceptsandtechnologieswhile working on them directly or while using their resources. More recently, they have even been used for the verification of their potential threats of gene flow from genetically modified plants and invasive habits. Above all, some of them are con- tributing enormously as model plant species to the elucidation and amelioration of thegenomesofcropplantspecies. Asamatteroffact,asastudent,ateacher,andahumblescienceworkerIwas,still am and surely will remain fascinated by the wild allies of crop plants for their invaluable wealth for genetics, genomics and breeding in crop plants and as such shareadeepconcernfortheirconservationandcomprehensivecharacterizationfor futureutilization.Itisbynowawellestablishedfactthatwildcroprelativesdeserve seriousattentionfordomestication,especiallyfortheutilizationoftheirphytomedi- cinesandnutraceuticals,bioenergyproduction,soilreclamation,andthephytoreme- diationofecologyandenvironment.Whilethesevastlypositiveimpactsofwildcrop relativesonthedevelopmentanddeploymentofnewvarietiesforvariouspurposesin the major crop plants of the world agriculture, along with a few negative potential concerns,areenvisagedtheneedforreferencebookswithcomprehensivedelibera- tions on the wild relatives of all the major field and plantation crops and fruit and foresttreesisindeedimperative.Thiswasthedrivingforcebehindtheinceptionand publicationofthisseries. UnliketheprevioussixbookprojectsIhaveeditedaloneorwithco-editors,this timeitwasverydifficulttoformulateuniformoutlinesforthechaptersofthisbook series for several obvious reasons. Firstly, the status of the crop relatives is highly diverse. Some of them are completely wild, some are sporadically cultivated and someareattheinitialstageofdomesticationforspecificbreedingobjectivesrecently deemedessential.Secondly,thestatusoftheirconservationvarieswidely:somehave been conserved, characterized and utilized; some have been eroded completely exceptfortheirpresenceintheircenter(s)oforigin;someareat-riskorendangered duetogeneticerosion,andsomeofthemhaveyettobeexplored.Thethirdconstraint is the variation in their relative worth, e.g. as academic model, breeding resource, and/orpotentialas“newcrops.” ix