Advances in Studies of Genetic Improvement of Sugarcane

AdvancesinStudiesofGeneticImprovementofSugarcane

MingfuWEN，JunxianYANG*，FangyinPAN，WenlongWU，YueguiCHEN，JinyanGUAN

GuangzhouSugarcaneIndustryＲesearchInstitute/GuangdongKeyLabofSugarcaneImprovement＆Biorefinery，Guangzhou510316，China

AbstractSugarcane（Saccharumspp．）isalargeperennialherbaceousplantthatiscultivatedintropicalandsubtropicalregionsofthe

world，anditisalsooneofthemostefficientcropsintheworldinconvertingenergyfromsunlightintochemicalenergy．Asanessentialsugar

sugarcaneisreceivinganincreasingconcernforitsvarietyimprovement．Traditionalbreedingandcultivationtechniquescropandenergycrop，

havecontributedalottoincreasingsugarcaneyieldandsucrosecontent．Inrecentyears，developmentandapplicationofbiotechnologyprovidemuchhelpforgeneticimprovementofsugarcane．Forconvenienceofbreedersfullyknowingadvancesinstudiesofsugarcanegeneticimprove-ment，thispaperelaboratedconventionalbreeding，genomics，GMtechnology，andmolecularmarkerassistedbreeding．KeywordsSugarcane，Geneticimprovement，Genomics，Transgenetechnology，Molecularmarkerassistedbreeding

1Introduction

Sugarcane（Saccharumspp．）isthelargestsugarcanecropinChi-naandtheworld．Sucroseaccountsformorethan75%ofthetotalsugaryieldoftheworld，andmorethan90%ofChina'ssugaryield．Sugarcaneisalsothefirstgenerationbioenergycrop，anditcanbeusedasrenewableenergyforproductionoffuelethanolandbiomassproducts．Fromthe1970s，manycountriesstartedformu-latingthecleanenergystrategyofdevelopingfuelethanoltore-placeoil．Amongthesecountries，Brazilwasmostsuccessfulinusingsugarcanetoproducefuelethanolaspowerenergyofauto-mobile．In1979，thefirstbioenergyautomobileusingfuelethanolasdriveappeared．By2003，therewere15．5millioncarsusingmixedfuelofethanolandoil，andtherewere2．2millioncarscompletelyusingethanolasthefuel［1］．In2010，theyieldofbioenergyusingsugarcaneasmainmaterialhadreplacedabout

bringingittobethecountry47．6%energyconsumptionofBrazil，［2］

withhighestbioenergyutilizationrate．Sugarcaneisthegeneral-lyrecognizedC4plantwithhighphotosyntheticrateanddrymatteraccumulationability［3－4］，andisthecropwithhighestperunitare

［5］

yielduptooneton．AccordingtostatisticsoftheFoodandAg-ricultureOrganization（FAO），in2011/2012sugarcanecrushingseason，thetotalsugarcaneyieldofBrazilreached7．34×108t，withtheperunitareayieldupto76．45t/hm2．InIndiaandChi-na，theplantingarea，totalyield，andaverageyieldofsugarcane

3．42×108t，and69．25t/hm2，1．73×106was4．94×106hm2，

hm2，1．15×108t，66．52t/hm2separately．

2Targetofsugarcanegeneticimprovement

Sugarcaneisanessentialsugarcropandenergycrop，anditsvari-etyimprovementisreceivingcloseattention．Sugarcanehasuniquegeneticmode．Itisallopolyploidcropgeneratedthrougha

2016Accepted：November10，2016Ｒeceived：September1，

SupportedbyScienceandTechnologyPlanningProjectofGuangdongProvince（2014A030304012，2014A020208012，2015A030302009）；ScienceandTech-nologyPlanningProjectofZhanjiangCity（2015A03014）．*Correspondingauthor．E-mail：siriyjx@163．com

seriesofhybridizationofpolyploidSaccharumofficinarumL．（2n=80，X=10）asfemaleparentandSaccharumspontaneumL．（2n=40～128，X=8）asmaleparent．Thenumberofchro-mosomesis100－150，about75%－85%fromSaccharumoffici-narumL．，and15%－25%fromSaccharumspontaneumL．Inthefirstandsecondtimeofhybridizationprocess，chromosomesweredeliveredin2n+nspecialmanner．Agronomiccharacterssuchassugarcontentandyieldofhybridgenerationwererapidlyrecoveredandstabilized．Thisprocessiscallednobilityprocessofsugarcane［7］．However，suchuniquegeneticmodeincreasesthedifficultyinsugarcanegeneticimprovement．Withrapiddevelop-mentofmodernbiotechnology，anddropofgenomesequencingcosts，thedevelopmentofsugarcanefunctionalgenomicsandstructuralgenomicsplaysagreatroleinpromotingfurtherdevelop-mentofsugarcanegeneticimprovement．Varietyisthekeyforde-velopmentofthesugarcaneindustry．Finevarietynotonlyincrea-reducesproductionsestheperunitareayieldandsucrosecontent，

costs，butalsoextendsthecrushingperiod，increasesequipment

andobtainshighereconomicbene-utilizationrateofsugarfactory，

fits．WithreferencetosugarcaneproductionanddevelopmentinChinaandforeigncountries，itisnecessarytoconstantlyupgradevarieties，topromotesustainableandhealthydevelopmentofsug-arcaneindustryandsatisfythemarketdemands．Accordingtore-quirementsofmodernsugarcaneindustry，thesugarcanebreedingtargetscanbeclassifiedintotwotypes：（i）improvementofvarie-tytraits，includingimprovingsugarcontent（sucrosecontent，brix，apparentpurity，andgravitypurity），increasingyield（tille-ringcapacity，effectivestalknumber，stalkdiameter，plantheight，growthrate，sproutingrate，andratoonperformance），in-creasingstressresistance（diseaseresistance，insectresistance，lodgingresistance，droughtresistance，coldresistanceandbarrenresistance），andselectingvarieties（sugarcanehairanddefolia-tion）［8－9］；（ii）cultivationofdifferentpurposesofvarieties，mainlyincludingsugartypesugarcane（fiberfraction≤14%），energytypesugarcane（fiberfraction≥30%），andsugarenergytypesugarcane（fiberfraction＜30%）［10］．

MingfuWENetal．AdvancesinStudiesofGeneticImprovementofSugarcane3Advancesinconventionalbreedingofsugarcane

3．1DevelopmentstagesofsugarcanebreedingThedevel-opmentofsugarcanebreedingcanbedividedinto5stages［9］．（i）Thestageofbreedingusingtropicalstrain．Thisstagestartedfrom1858whenBarbadosreportedthatsugarcanecanbearfruit．Atthisstage，maincharacteristicsofvarietiesincludedhighsugarcontent，lowfiberfraction，highpurity，lowadaptation，weakra-toon，andpoordiseaseresistance，suchasH109，B716，andQ813，etc［11］．（ii）Thestageofbreedingusingnobilityprocesstoselectnoblevarieties．In1885，Soltwedelmadeexperimentofhy-bridizationbetweensugarcaneandErianthusarundinaceus［12］；in13，MoquetteandWakkerobtainedhybridvarietyoftropicalstrainBlackCheribonandIndianstrainGansha［13］；in17，Ko-bushybridizedIndianstrainChunniandtropicalstrain［14］；in1911，WilbrinksuccessfullyhybridizedIndianstrainGanshaandtropicalstrain［15］；later，JeswietbackcrossedthetropicalstrainwiththehybridgenerationobtainedbyWilbrink，rapidlyrecov-eredandstabilizedsugarcontentandyieldofhybridgenera-tion［16］；in1921，JeswietsuccessfullybredPOJ2878finevarietythroughhybridization［17］．Thisstageopenedthecurtainofinter-specifichybridization，andrecoveredandstabilizedsugarcontentandyieldofhybridgeneration，andbredexcellentvarietieswith

highsugarcontent，

highfiberfraction，highadaptation，strongra-toon，andhighdiseaseresistance，suchasCo281andCo290［18］．（iii）Thestageofbreedingusingnoblevarieties．From1930to1950，itwasthestageofhybridizationusingnoblevarietiestose-lectexcellentnewvarieties．TypicalexamplesincludedCo419bredusingPOJ2878×Co290in1937，popularNCo310inthe1950sand1960sbredin1939，andH32－8560bredbyHawaiiＲesearchCenterin1945（accountingformorethan60%ofthelo-calplantingarea）［17］．Thisstagemainlyusednoblevarietiestohybridizeandbreedmoregermplasmresources．（iv）Thestageofbreedingusingnoblehybridvarieties．From1950to1965，noblehybridvarietieswereusedtoselectmodernsugarcanevarietieswithhighsugarcontent，highyield，highstressresistance，andexcellentratoon，andthesenewvarietiesweredistributedinallsugarcaneplantingareas［19］．Atthisstage，excellentgermplasmresourcesbredthroughnobilityprocesswereusedashybridpar-ent，tofurtherconsolidateexcellentgenesandselectbettermodernsugarcanevarieties．（v）Thestageofexpandingvarietygeneticconstitutionsource．Atthisstage，itmainlywasengagedinrapid-lyimprovingsugarcontent，yieldandotheragronomictraitsthroughconstanthybridizationorbackcrossingtoincreasefunda-mentalsubstances［20］．

3．2BreedingofsugarcanevarietiesinbothChinaandfor-eigncountriesNewsugarcanevarietiesbredbyforeigncountriesmainlyincludeＲBvarietiesbredbyBrazilianFederalUniversityandIACvarietiesbredbyCampinasAgriculturalＲesearchInstitu-teofSaoPaulo［21］，suchasＲB99395andIAC86-2210；CPse-ries，Hseries，andHoCPseriesbredbyUSDAAＲSSugarcaneFieldStationCanalPoint，HawaiiＲesearchInstitute，andLouisi-anaSugarcaneＲesearchInstitute［22］；Co997，

Co1001，andCo52767

bredbyIndia［22］；Q174，Q205，andQ208bredbyAustralia［23］；

POJandEKvarietiesbredbyIndonesia．Thebreedingofsugar-caneinChinacanbedividedinto3stages［24－25］．（i）Breedingoflocalvarieties．Thisstagewasmarkedbyplantinglocalvarietiesassugarrefiningmaterials，typicalvarietiesincludedbamboocane，reedcane，andＲohancane．（ii）Introductionofforeignvarieties．Atthisstage，foreignvarietieswereintroducedandpop-ularized，andtypicalvarietiesincludedPOJ2878，POJ2725，NCo310，Co290，Co281，CP49-50，andCP34-120．（iii）Self-breedingvarieties．Atthisstage，varietiesbredbyChinastartedreplacingforeignvarieties．Inrecentyears，self-breedingvarietiesofvarietiesmainlyincludedYuetang，Guitang，Mintang，Yunzhe，Liucheng，andTaitang．Atpresent，newTaitang22，Yutang93-159，Yutang00-236，Guitang21，andLiucheng05-136takeup

thedominantpositionofsugarcaneplantinginChina．However，thegrowthrateforsugarcontentandyieldofnewsugarcanevarie-tiesbecomessmallerandsmaller．Thiscanbeprovedbyevalua-tionof4mainvarieties，Yuetang85-177，Yuetang99-66，Yuet-ang00-236，andYuetang03-393．Waclawovskyelal．［26］foundthatthegrowthrateofsugarcaneyieldintheworldremainedat

1%－1．5%inrecentyears，anditwilldeclineinfuture．ThismaybelargelybecausebreedingparentmainlycomesfromF4andF4ofPOJ2878［27］andgermplasmresourcesforimprovementareverylimited．Inthissituation，biotechnologymaybecomethekeyforsugarcanegeneticimprovement．

4Advancesinstudiesofsugarcanegeneticimprove-mentthroughbiotechnology

4．1SugarcanegenomicsSugarcanegenomicsisanindispen-sabletoolforfuturesugarcaneimprovement．However，complexgenomeofallopolyploidandinterspecifichybridizationofmodernstrainshinderstudiesofsugarcanegenomicsandapplicationofge-nomicsinsugarcanebreedingprocess．Bynow，inNCBIdata-base，thereare31555nucleotidesequences（including1216mＲ-NAsequences）availablefromdifferentsugarcanevarietiesand284818ESTsequences（http：//www．ncbi．nlm．nih．gov/）．TheseESTsequencescomefromcDNAlibraryandmanytran-scriptsofmorethan70varietiesofsugarcane，

andmaterialsmain-lyincludeseedlings，roots，stalks，leaves，flowers，andseedsof

sugarcane，

andcallustreatedbynonbiologicalstressandseedlingsinfectedbyendogenousnitrogen-fixingbacteria［28］．Atpresent，

bacterialartificialchromosome（BAC）libraryforsugarcaneisbuiltbyhybridstrainＲ570withnumberofchromosomeof2n=115．Thislibrarycontains283158clonesandcovers1．3timesofpolyploidygenomeofthisvariety（itispredictedthatthegenomeis10Gb）［29］．ItisreportedthatscientistsfromBrazilandothercountriesarebuildingfinephysicalmapforthisBAClibrary［30］．Meanwhile，scientistsarebuildingBAClibraryandthelibraryclonesequencingforBrazilianvarietySP80-3280［31］．Besides，in

cooperationwithShenzhenBGI，SugarcaneＲesearchCenterofChineseAcademyofAgriculturalScienceisbuildingBAClibraryandwholegenomesequencingforthinstalkwildsugarcanevariety

68AsianAgriculturalＲesearch2016

GXS87－16（2n=）［32］．ThebuildingofsugarcaneBAClibraryanddetailedphysicalmapinformationareofutmostimportancetounderstandingstructureofsugarcanegenome．Inthewholegenomesequencing，sorghumisacropwithcloseaffiliationwithsugar-cane，anditsgenomesequenceisofmuchhelpforstudiesofsug-arcanegenome，andcompletionofsorghumgenomesequencingprovidesimportantcomparativegenomicstoolforsugarcanege-nomicstudies［33－34］．Wangetal．［35］carriedouthybridizationinBAClibraryusing1961singlecopysorghumoligonucleotideprobesandsugarcanecommercialvarietyＲ570，obtained20sug-arcaneBACs，witheachBACcorrespondingtosorghumchromo-somearm．About95．2%sequencesofcodingareaofsugarcaneBACsmatchthesorghumsequences．Ifusingsorghumgenomeastemplatetosequencethecontig，itcancover78．2%of20BACs．About53．1%sugarcaneBACsmatchsorghumsequences．Inare-asthatcanbelinked，209genesofsugarcanehavebeenannota-ted，202sorghumgeneshavebeenannotated，including17genesuniquetosugarcane，andallhavebeenverifiedbysugarcaneex-pressedsequencetags（ESTs），in12genesuniquetosorghum，onlyonehasbeenverifiedbysorghumESTs．In17genesuniquetosugarcane，12genesdonothavematchingproteininGenBanknon-redundantproteindatabase，andtheymayothertypesofpro-teinparticipatingincodingsugarcanespecialprocess．Ｒelativeto

thesugarcane，

linealhomologicalareaofsorghumexpands，whichisrealizedmainlythroughincreaseinreversetranscriptiontranspo-son．Therefore，sugarcaneandsorghumgenomesarecollinearinmostgeneareas．SorghumgenomecanbeusedasDNAassemblytemplateofallopolyploidsugarcanegenomes．

4．2SugarcaneTransgenetechnologyTheinvitrocultureandregenerationsystemtechnologyhavebeenestablishedandgraduallyimprovedsince40yearsago［36］，whichisveryimportantfordevelopmentofsugarcanegenetictransformationsystem．Nev-ertheless，aneuploidypolyploidy，hugegenomeandcomplexge-neticbackgroundofsugarcanepresenttheproblemoflowtransfor-mationefficiencyoftransgenetechnology．Sincegenegunmethod（particlebombardment）featuresnotsubjecttohost，widetargetreceptortype，highcontrollability，simpleandrapidoperation，itisamethodmainlyusedintheearlyperiodofthesugarcanetrans-genetechnology［37－38］．Withconstantdevelopmentandoptimiza-tionofgenetictransformationmethodswithfeaturesoflowcosts，highsuccessrate，andsinglecopyofallogenicgenes，andhighgeneticstability，ithasbeenwidelyappliedinsugarcanetransgene

technology［39］

．Intheprocessofsugarcanegenetictransformation，mainrestrictivefactorsincludelowtransformationefficiency，ac-tivetransgene，mutationinbodycellclone，anddifficultback-crossing［40］．Itisthusrequiredtofurtheroptimizethetransforma-tionmethod，bettercontrolthetransgeneexpression，andrealizestableexpression．Thesugarcanetransgenestudiesfocusonin-creasingsucroseaccumulation，stalkyield，improvingdiseasere-sistanceandstressresistance［40－45］．Ｒesearchesindicatethatgenesparticipatinginmetabolismofcellwallhavedifferencesin

expression［40］．ThroughcDNA-SCoTanalysisofratoonstuntingdiseaseinducedsugarcanedifferenceexpressiongene，theyfound

manygenesparticipatingininteractionofratoonstuntingdis-ease［46］．ThroughsubtractivelibrarytechnologyandcDNAchiptechnologyanalysis，SSADHrelatedtosugarcanewaterstressre-sponsewasscreened［47］．Throughexcessiveexpressionordown-wardmodulationofviruscoatprotein，mＲNAexertsresistanceagainstSCMV［48］，SCYLV［49］，andFDV［50］．ThroughadjustingexpressionofSc-EＲSgene，itisabletostrenthenphotosynthesis

ofsugarcaneleavesandimprovedroughtresistanceofsugar-cane［51］．Thesekeygenesarefavorableforcultivationofnewsug-arcanevarietieswithhighyield，highsugarcontent，andhighad-aptation．Atpresent，non-commercialsugarcanetransgenestrains

havemadebreakthroughadvancesandsomestrainsareundergoing

thefieldexperiment［21，

40］

．However，limitationofregulationsoftransgeniccropswillretardthereleaseofcommercialtransgenicvarieties．Therefore，thebreedingandapplicationofanewtrans-genicvarietytakesaconsiderablelongtime．

4．3MolecularmarkerassistedbreedingAtpresent，thenumberofsugarcanechromosomesforgeneticmappinganalysisismorethan100，butgenomesequenceformolecularmarkerisverylimited．Mostgeneticmapsarebasedondominantmarkers，whichareusedassingledosemarkers．Posteritysegregationratioexistsasper1：1（markedas\"1\"），doesnotexist（markedas\"0\"）forstatisticalanalysis．Forallopolyploidsugarcane，suchstatisticalmethodwillonlyprovideanapproximatevaluewhenestimatingtherecombinationrateandlinkage；besides，someevidencesindicatethatsingledosemarkeronlydetectsabout70%polymorphismlo-ci［52］．13sugarcanemappinggroupswereusedtobuild18molec-ulargeneticlinkagemap，and1500－2000markerswereused［53］，includingＲFLP［］，AFLP［55］，TＲAP［］，EST-SSＲ［56］and

DAＲT［57］，indicatingallbuiltsugarcanegeneticmapsareincom-plete．Toobtainhighdensitygeneticmapscoveringthewholesug-arcanegenome，itneedsdevelopingmoreSNPmarkers．However，throughQTLpositioningofsugarcanerelatedtraits，ithasobtainedQTLlocirelatedtodiseaseresistance，stressresistance，yield，andsugarcontent［58－62］．Becauseofcomplexityofsugarcanege-nome，fortargettraits，mostgenomescannotbescanned，andsuchdefectlimitstheapplicationofthemolecularmarkerassistedselection．Atthesametime，thesugarcanegeneticmodeindicatesthatthegeneticlinkageunbalancewidelyexists［63］．Whentraitre-latedmolecularmarkersareusedtodetermineQTLofsugarcanethroughlinkageanalysisandcorrelationanalysis，lowdensitymarkerandroughgeneticstatisticalmethodsarestillrelativelydif-ficult．Insugarcanebreeding，itisachallengingtasktousemo-lecularmarkerassistedselection．Manyimportanttraitsarejointly

determinedbymanytraitloci，

andeachtraitonlycontributesalit-tletotheoverallphenotype［］

．ThesugarcaneQTLpositioningismainlybasedonsingledosemarkeranalysisorcompositeintervalmapping［63］．Toobtaineffectiveresults，itneedsdevelopingnewresearchmodelsandstatisticalmethods，andalsoconsidersinflu-enceofinteractionbetweenQTLandenvironmentandgenecorre-lation．Therefore，therearestillmanychallengestobesolvedin

MingfuWENetal．AdvancesinStudiesofGeneticImprovementofSugarcanesugarcanemolecularmarkerassistedbreeding．

5Conclusions

Sugarcaneisanessentialsugarcropandenergycrop，anditsvari-etyimprovementisreceivingcloseattention．Traditionalbreeding

andcultivationtechniqueshavecontributedalottoincreasingsug-arcaneyieldandsucrosecontent．Withrapiddevelopmentofmod-ernbiotechnology，relyingonitsimportanceinagricultureandin-dustries，sugarcaneattractsmanyscientiststomakejointeffortsinmolecularbiology，bioinformatics，andgenetics．Besides，withapplicationofnewgenerationoflowcostDNAsequencingtechnol-ogy，theallopolyploidsugarcanegenomesequencingwhichwascostlyinthepastbecomespossible．Infuture，biotechnologyge-neticimprovementtechniquewillacceleratetheprogressoftradi-tionalsugarcanebreedingandcultivatemoreexcellentsugarcanevarieties．

Ｒeferences

1］BOＲＲEＲOMAV，PEＲEIＲAJTV，MIＲANDAEE．Anenvironmental

managementmethodforsugarcanealcoholproductioninBrazil［J］．Bio-massandBioenergy，2003，25（3）：287－299．2］HOFSETZK，SILVAMA．Braziliansugarcanebagasse：Energyandnon-energyconsumption［J］．BiomassandBioenergy，2012，46：5－573．3］ＲAEAL，JACKSONMA，NGUYENCH，etal．Functionalspecialization

ofvacuolesinsugarcaneleafandstem［J］．TropicalPlantBiology，2009，2（1）：13－22．4］AＲNOULTS，BＲANCOUＲT-HULMELM．Areviewonmiscanthusbio-massproductionandcompositionforbioenergyuse：Genotypicandenvi-ronmentalvariabilityandimplicationsforbreeding［J］．BioenergyＲe-search，2015，8（2）：1－25．5］LOOMISＲS，WILLIAMSWA．Maximumcropproductivity：anestimate

［J］．CropScience，1963，3（1）：67－72．6］FAO．FAOSTATdata［EB/OL］．http：//faostat3．fao．org/home/index．

html，2011．7］BUTTEＲFIELDMK，D'HONTA，BEＲDINGN．Thesugarcanegenome：

asynthesisofcurrentunderstanding，andlessonsforbreedingandbio-technology［J］．ProcSAfrSugTechnolAss，2001，75：1－5．8］LIQW，DENGHH．Majorproblemsandstrategyinsugarcanebreeding

andvarietyextensioninChina［J］．SugarcaneandCanesugar，2011（4）：70－76．（inChinese）．9］MINGＲ，MOOＲEPH，WUK，etal．Sugarcaneimprovementthrough

breedingandbiotechnology［J］．PlantBreedingＲeviews，2006，27：15．10］LOUＲEIＲOME，BAＲBOSAMHP，LOPESFJF，etal．Sugarcane

breedingandselectionformoreefficientbiomassconversionincellulosicethanol［M］．Ｒoutestocellulosicethanol，SpringerNewYork，2011：199－239．11］KENNEDYAJ，ＲAOPS，OＲIOLP．Handbook，October2000［J］．

Cirad，2011．12］STEVENSONGC．Geneticsandbreedingofsugarcane［M］．London：

Longmans，Green，1965．13］JESWIETJ．Thedevelopmentofselectionandbreedingofthesugarcane

inJava［C］．ProcCongrInternSocSugarCaneTechn，1930，3：44－57．14］SMAＲTTJ，SIMMONDSNW．Evolutionofcropplants．ed．2［J］．

LongmanScientific＆Technical，1995．15］ETHIＲAJANAS．Sugarcanehybridizationtechniques［Z］．In：anony-mous（eds．），CopersucarInternationalSugarcaneBreedingWorkshop．

Copersucar，Brazil，1987：129－148．16］SIMMONDSNW．Evolutionofcropplants［M］．（London）：Longman

69

GroupLtd．，1976．［17］JESWIETJ．Thedevelopmentofselectionandbreedingofthesugarcane

inJava［C］．ProcCongrInternSocSugarCaneTechn．，1930：44－57．［18］ＲOACHBT．Originandimprovementofthegeneticbaseofsugarcane

［J］．ProcAustSocSugarCaneTechnol，19，11：34－47．［19］TEWTL．Worldsugarcanevarietycensus-year2000［J］．SugarCane

International，March/April2003：12－18．［20］WALKEＲDIT．Manipulatingthegeneticbaseofsugarcane［C］//Coper-sucarInternationalSugarcaneBreedingWorkshop．Copersucar，Piraci-caba，Brazil．1987：321－334．［21］CHEAVEGATTI-GIANOTTOA，ABＲEUHMCD，AＲＲUDAP，etal．

Sugarcane（Saccharum×officinarum）：areferencestudyfortheregu-lationofgeneticallymodifiedcultivarsinBrazil［J］．Tropicalplantbiol-ogy，2011．［22］DUNCKELMANPH，BＲEAUXＲD．Breedingsugarcanevarietiesfor

Louisianawithnewgermplasm［J］．ProcIntSocSugarCaneTechnol，1972，14：233－239．［23］HEINZDJ．WildSaccharumspeciesforbreedinginHawaii［J］．Proc

IntSocSugarCaneTechnol，1967，12：1037－1043．［24］LIQW，CHENZY，LIANGH．Modernsugarcaneimprovementtechnolo-gy［M］．Guangzhou：SouthChinaUniversityofTechnologyPress，

2000：11－13．［25］CHENＲK，XULP，LINYQ，etal．Modernsugarcanegeneticsand

breeding［

M］．Beijing：ChinaAgriculturePress，2011：2－12．［26］WACLASOVSKYAJ，SATOPM，LEMBKECG，etal．Sugarcanefor

bioenergyproduction：anassessmentofyieldandregulationofsucrosecontent［J］．PlantBiotechnolJ，2010，8：263－276．［27］JACKSONPA．Breedingforimprovedsugarcontentinsugarcane［J］．

FieldCropsＲesearch，2005，92（2）：277－290．［28］MANNEＲSJM，CASUＲE．Transcriptomeanalysisandfunctionalge-nomicsofsugarcane［J］．TropicalPlantBiology，2011，4（1）：9－21．［29］TOMKINSJP，YUY，MILLEＲ-SMITHH，etal．Abacterialartificial

chromosomelibraryforsugarcane［J］．TheoreticalandAppliedGenet-ics，1999，99（3－4）：419－424．［30］SETTAND，METCALFECJ，CＲUZGMQ，etal．Buildingthesugar-canegenomeforbiotechnologyandidentifyingevolutionarytrends［J］

．BmcGenomics，2014，15（4）：1－18．［31］DAL-BIANCOM，CAＲNEIＲOMS，HOTTACT，etal．Sugarcaneim-provement：howfarcanwego［J］．CurrentOpinioninBiotechnology，

2012，23（2）：265－270．［32］LIYＲ，YANGLT．Ｒesearchanddevelopmentprioritiesforsugarindus-tryofChina：Ｒecentresearchhighlights［J］．SugarTech，2015，17

（1）：9－12．［33］BEDELLJA，BUDIMANMA，NUNBEＲGA，etal．Sorghumgenome

sequencingbymethylationfiltration［J］．PLoSBiology，2005，3（1）：13．［34］AITKENKS，MCNEILMD，BEＲKMANPJ，etal．Comparativemap-pinginthePoaceaefamilyrevealstranslocationsinthecomplexpoly-ploidgenomeofsugarcane．［J］．BmcPlantBiology，2014，14（1）：190

－190．

［35］WANGJ，ＲOEB，MACMILS，etal．Microcollinearitybetweenau-topolyploidsugarcaneanddiploidsorghumgenomes［

J］．BMCGenom-ics，2010，11（1）：261．［36］LAKSHMANANP，GEIJSKESJ，AITKENKS，etal．Sugarcanebio-technology：thechallengesandopportunities［

J］．InVitroCellDevBi-ol-Plant，2005，41：345－363．

［37］LIYＲ．Theapplicationofbiotechnologyinsugarcane［M］．Beijing：

ChinaAgriculturePress，2009．（inChinese）．［38］WENGLX，DENGHH，XUJL，etal．Transgenicsugarcaneplantsex-pressinghighlevelsofmodifiedcry1Acprovideeffectivecontrolagainst

stemborersinfieldtrials［J］．TransgenicＲesearch，2011，20（4）：759

［［［［［［［［［［［［［［［［70AsianAgriculturalＲesearch2016

－772．

［39］MANICKAVASAGAMM，GANAPATHIA，ANBAZHAGANVＲ，etal．

Agrobacterium-mediatedgenetictransformationanddevelopmentofher-bicide-resistantsugarcane（Saccharumspecieshybrids）usingaxillarybuds［J］．PlantCellＲeports，2004，23（3）：134－143．［40］HPTTACT，LEMBKECG，DOMINGUESDS，etal．Thebiotechnology

roadmapforsugarcaneimprovement［J］．TropicalPlantBiology，2010，3（2）：75－87．［41］DEBIBAKASS，ＲOCHEＲS，GAＲSMEUＲO，etal．Prospectingsugar-caneresistancetosugarcaneyellowleafvirusbygenome-wideassocia-tion［J］．Theoretical＆AppliedGenetics，2014，127（8）：1719－1732．［42］ZHANGM，ZHUOX，WANGJ，etal．Phosphomannoseisomeraseaf-fectsthekeyenzymesofglycolysisandsucrosemetabolismintransgenic

sugarcaneoverexpressingthemanAgene．［J］．MolecularBreeding，2015，35（3）：1－10．［43］PＲIJIPJ，HEMAPＲABHAG．Sugarcanespecificdroughtresponsive

candidategenesbelongingtoABAdependentpathwayidentifiedfrom

．SugarbasicspeciesclonesofSaccharumsp．andErianthussp．［J］

Tech，2014，17（2）：130－137．［44］MCINTYＲECL，GOODEML，COＲDEIＲOG，etal．Characterisation

ofallelesofthesucrosephosphatesynthasegenefamilyinsugarcaneandtheirassociationwithsugar-relatedtraits［J］．MolecularBreeding，

2015，35（3）：1－14．［45］CHENZL，GUIYY，QINCX，etal．Isolationandexpressionanalysis

ofsucrosesynthasegene（ScSuSy4）fromsugarcane［J］．SugarTech，2015，18（2）：134–140．［46］CASUＲE，JAＲMEYJM，BONNETTGD，etal．Identificationoftran-scriptsassociatedwithcellwallmetabolismanddevelopmentinthestem

ofsugarcanebyaffymetrixgenechipsugarcanegenomearrayexpressionprofiling［J］．FunctIntegrGenomics，2007，7（2）：153－167．［47］CHENMH，ZHANGBQ，SONGXP，etal．cDNA-SCoTanalysisofdif-ferentiallyexpressedgenesinsugarcaneinducedbyLeifsoniaxylisub-sp，xyli［J］．ActaAgronomicaSinica，2013，39（6）：1119－1126．

（inChinese）．［48］ZHANGJS，GUOCF，WANGBM，etal．Cloningandexpressionanaly-sisofawaterstress-inducedALDHgenefromsugarcane［J］．Scientia

AgriculturaSinica，2009，42（8）：2676－2685．（inChinese）．［49］GILBEＲTＲA，GLYNNNC，COMSTOCKC，etal．Agronomicperform-anceandgeneticcharacterizationofsugarcanetransformedforresistance

tosugarcaneyellowleafvirus［J］．FieldCropsＲesearch，2009，111（1－2）：39－46．

［50］ZHUYJ，MCCAFFEＲTYH，OSTEＲMANG，etal．Genetictransfor-mationwithuntranslatablecoatproteingeneofsugarcaneyellowleafvi-rusreducesvirustitersinsugarcane［J］．TransgenicＲesearch，2011，

20（3）：503－512．［51］MCQUALTEＲＲB，DALEJL，HAＲDINGＲM，etal．ProductionandevaluationoftransgenicsugarcanecontainingaFijidiseasevirus

（FDV）genomesegmentS9-derivedsyntheticresistancegene［J］．Aus-tralianJournalofAgriculturalＲesearch，2004，55（2）：139－145．

［52］WEIYW．Studiesongenedifferentialexpressionsregulatedwithethep-honandcloningofethylenereceptorgeneSc-EＲSinsugarcane（Sac-charumofficinarumL．）［D］．Nanning：GuangxiUniversity，2007．

（inChinese）．［53］ALWALAS，KIMBENGCA．MoleculargeneticlinkagemappinginSac-resourcesandachievements［M］．CＲCPress，Sci-charum：strategies，

encePublishers，2010．［］WUＲ，MACX，WUSS，etal．Linkagemappingofsex-specificdiffer-ences［J］．GeneticalＲesearch，2002，79（1）：85－96．［55］ANDＲUS，PANYB，THONGTHAWEES，etal．Geneticanalysisof

thesugarcane（Saccharumspp．）cultivar‘LCP85-384’．I．Linkage

mappingusingAFLP，SSＲ，andTＲAPmarkers［J］．Theoreticaland

2011，123（1）：77－93．AppliedGenetics，

［56］LIUXL，MAOJ，LUX，etal．Constructionofmoleculargeneticlink-J］．ActaAgro-agemapofsugarcanebasedonSSＲandAFLPmarkers［

nomicaSinica，2010，36（1）：177－183．（inChinese）．［57］OLIVEIＲAKM，PINTOLＲ，MAＲCONITG，etal．Functionalintegrat-edgeneticlinkagemapbasedonEST-markersforasugarcane（Saccha-rumspp．）commercialcross［J］．MolecularBreeding，2007，20（3）：

1－208．［58］WANGJ，ＲOEB，MACMILS，etal．Microcollinearitybetweenau-topolyploidsugarcaneanddiploidsorghumgenomes［J］．BMCGenom-ics，2010，11：261．［59］MAＲGAＲIDOGＲA，PASTINAMM，SOUZAAP，etal．Multi-trait

multi-environmentquantitativetraitlocimappingforasugarcanecom-mercialcrossprovidesinsightsontheinheritanceofimportanttraits［J］．MolecularBreeding，2015，35（8）：1－15．［60］BANEＲJEEN，SIＲAＲEEA，YADAVS，etal．Marker-traitassociation

studyforsucroseandyieldcontributingtraitsinsugarcane（Saccharumspp．hybrid）［J］．Euphytica，2015，205（1）：185－201．［61］SANTOSFＲC，PINTOLＲ，CAＲLINI-GAＲCIALA，etal．Marker-trait

associationandepistasisforbrownrustresistanceinsugarcane［J］．Eu-phytica，2014，203（3）：533－7．［62］GOUYM，ＲOUSSELLEY，CHANEAT，etal．Genomewideassocia-tionmappingofagro-morphologicalanddiseaseresistancetraitsinsug-arcane［J］．Euphytica，2015，202（2）：269－284．［63］PASTINAMM，PINTOLＲ，OLINEIＲAKM，etal．Molecularmapping

ofcomplextraits［C］．HenryＲJ，KoleC．Genetics，GenomicsandBreedingofSugarcane．CＲCPress，SciencePublishers，2010．［］WEIX，JACKSONPA，HEＲMANNS，etal．Simultaneouslyaccount-ingforpopulationstructure，genotypebyenvironmentinteraction，and

spatialvariationinmarker–traitassociationsinsugarcane［J］．Ge-nome，2010，53（11）：973－981．［17］DAPQ，ZHANGKL，YEYH，etal．Ｒestrictingfactorsandcounter-measuresofreforestationprojectinLhasa［J］．anScience＆Technology，2010（5）：13－16．（inChinese）．［18］XINGZ．Aquantitativeresearchonthebenefitofecologicalenvironment

oflandscapeplantin［M］．Beijing：SocialSciencesAcademicPress（China），2014．（inChinese）．

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［15］LIUZN，PANG，ZHANGHF，etal．Investigationandapplicationre-J］．JournalofYunnanAgriculturalsearchonlandscapeplantsinLhasa［

University，2016，31（4）：670－680．（inChinese）．［16］LIUZN，PANG，ZHANGHF，etal．Ｒesearchonlandscapeplantsin

Shannanof［J］．JournalofShandongAgriculturalUniversity（Nat-uralScienceEdition），2016，47（7）：82－．（inChinese）．