JAFC是什么意思C在线翻译读音例句

/JAFCPublishedonWeb05/25/20102010AmericanChemicalSociet民间故事会在线阅读 y

em.2010,58,7383–73887383

DOI:10.1021/jf100385m

C-TypeStarchfromHigh-AmyloseRiceResistantStarch

GranulesModifiedbyAntise蜀中九日九日登高 nseRNAInhibitionofStarc霜叶红于二月花上一句 h

BranchingEnzyme

CUNXUWEI,†,‡BINXU,FENGLINGQIN,‡HUAGUANGYU,CHONGCHEN,

XIANGLENMENG,†LIJIAZHU,†YOUPINGWANG,‡MINGHONGGU,*,†

AND

QIAOQUANLIU*,†

†KeyLaboratoriesofCropGeneticsandPhysiologyoftheJiangsuProvinceandPlant

FunctionalGenomicsoftheMinistryofEducation,‡CollegeofBioscienceandBiotechnology,and

TestingCenter,YangzhouUniversity,Yangzhou225009,China

High-amylosestarchisasourceofresistantstarch(RS)whichhasagreatbenefitonhumanhealth.

Atransgenicriceline(TRS)enrichedamyloseandRShadbeendevelopedbyantisenseRNA

study,thenativestarchgranuleswereisolatedfrom

TRSgrainsaswellasthewildtype,andtheircrystallinetypewascarefullyinvestigatedbeforeand

-amyloseTRSrice,theC-typestarch,whichmightresultfromthecom-

binationofbothA-typeandB-typestarch,wasobservedandsubsequentlyconfirmedbymultiple

physicaltechniques,includingX-raypowderdiffraction,solid-statenuclearmagneticresonance,and

er,thechangeofstarchcrystallinestructurefromC-toB-type

atacouldaddtoourunder-

standingofnotonlythepolymorphstructureofcerealstarchbutalsowhyhigh-amylosestarchismore

resistanttodigestion.

KEYWORDS:Rice(OryzasativaL.);high-amyloseresistantstarchgranule;C-typestarch;X-ray

powderdiffraction;solid-statenuclearmagneticresonance

INTRODUCTION

Starch,themostimportantreservecomponentinhigherplants,

especiallyincerealendosperm,isusuallysynthesizedasasemi-

crystallinegranulecontainingdenselypackedpolysaccharides

w,thecrystal-

linityofnativestarcheshasbeenwellstudiedandtheirpoly-

morphismhasbeenrecognizedunderX-raypowderdiffraction

(XRD)(1-3).Typically,therearethreetypesofstarchcrystal-

linityreported,knownasA-,B-,andC-type(1-3).

TheA-typestarchmainlyexistsincerealendosperm,andtheir

crystallinestructureismostlyfavoredbytheamylopectinwith

shortlateralchainsandclosedbranchingpoints(1).Thecrystal-

linestructureofB-typestarch,contrarytothatofA-type,is

usuallyformedbytheamylopectinwithlongsidechainsand

distantbranchingpointsandcanbeobservedintubercropssuch

rearelimitedreportsfortheC-typestarch,

exceptinsmooth-seededpeasandbeans(4).Usually,theC-type

example,inpeascontainingC-typestarch,starchinthecenterof

thegranuleadoptsaB-typecrystallinestructurewhilethe

peripheryiscomposedoftheA-typecrystallinestructure(4).

Normally,thenormalcerealstarchesshowA-typecrystalline

structuresasmentionedabove(1),buttheB-typecrystalline

structureisalsoobserved,especiallyinhigh-amylosecerealstar-

mple,Yanoetal.(5)foundthatseveralricemutants

withhighamylosecontent(AC,35.4%vs29.4%ofwildtype)

elineGoami2

(previouslyknownasSuweon464),containing33%ofACand

abouttwiceasmuchaswildtypeIlpumbeyo,alsopresentsa

B-typestarchstructure(6).Inmaize,theB-typestarchisalsore-

portedinthehighamylosevarieties(2),butnotallthehigh-

ceandbarleymutants

withhighamyloseareidentifiedtocontainatypicalA-typeXRD

pattern(7,8).BesidesthetypicalA-andB-types,theC-typecereal

starchesarealsopresented,amand

Taoreportedthatthecrystaltypeofmaizestarchcouldbevaried

fromA-toB-viaC-typewhenACincreased,andthetransition

occurredatabout40%(2).

Recently,wehavedevelopedseveralhigh-amylosetransgenic

ricelinesbyantisenseRNAinhibitionofthestarchbranching

enzymes(SBEs)(9,10).Thesetransgenicricegrainsarerichin

resistantstarch(RS)andhavebeenproventoshowasignificant

potentialtoimprovethelargebowelhealthinrats(11).Our

resultsfrommicrostructureandultrastructurestudiesrevealed

thatthesehigh-amylosestarchgranulesconsistofsemicompound

starch,muchdifferentfromthecompoundstarchinthestarch

granulesfromwildtyperice(9).

*,86-514-

87997217;E-mail,yzuwcx@().Phone,86-514-

87996648;E-mail,qqliu@();E-mail,gumh@

().

em.,Vol.58,No.12,2010Weietal.

Inthisstudy,thenativestarchgranuleswerefurtherisolated

fromthehigh-amylosegrainsaswellastheirwildtype,andtheir

crystallinetypewascarefullyinvestigatedbeforeandafteracid

stingly,inhigh-amyloserice,theC-typestarch

structurewaspresentedandconfirmedbymultiplephysical

techniques,includingXRD,13Ccross-polarizationmagic-angle

spinningnuclearmagneticresonance(13CCP/MASNMR),and

attenuatedtotalreflectance-Fouriertransforminfrared(ATR-

FTIR).Moreover,thechangeofstarchcrystallinityfromC-to

B-typeduringacidhydrolysiswasalsoobservedintheRS-rich

transgenicrice.

MATERIALSANDMETHODS

caricecultivarTe-qing(TQ)anditstrans-

genicline(TRS)

generatedfromTQaftertransgenicinhibitionoftwoSBEs(SBEIand

SBEIIb)throughantisenseRNAtechniqueandheldthehomozygous

transgene(9,10).TRS(inT8generation)andTQweresimultaneously

cultivatedintheexperimentfieldofYangzhouUniversity,Yangzhou,

China,in2009,andtheirmaturegrainswereusedtoisolatestarch

pea(PisumsativumL.)andpotato(Solanumtuberosum

L.)wereobtainedfromalocalnaturalfoodmarket.

starchgranuleswere

isolatedaspreviouslydescribed(9)exceptthatthesamplesandstarch

arentACwasdetermined

byusingacolorimetricmethodwithiodine-potassiumiodide(12).

d-modifiedstarchwas

preparedaccordingtothemethodofWangetal.(13)withaslightmodi-

msofisolatednativestarchweresuspendedin100mLof

tainerswereplacedinan

ovenat35Cforaperiodfrom0to20daysandgentlyshaken3timesby

he

certaintimeofhydrolysis,thesolventwascentrifuged(10min,3000g)and

thesupernatantwasusedformeasurementofthesolubilizedcarbohyd-

ratestoquantifythedegreeofhydrolysisbytheanthrone-H

2

SO

4

method(14).Theundissolvedresiduesweresubsequentlywashedthree

timesw敕勒歌北朝民歌古诗 ithddH

2

Oandtwotimeswithacetoneandthendriedat25

driedstarchesweregroundintopowdersandpassedthrougha100-mesh

overyyield(wt%)ofthestarchafteracid

hydrolysiswascalculatedbasedonthechangeofdriedstarchafterand

beforehydrolysis.

-amylase

degradedstarchgranuleswerepreparedaccordingtothemethoddesc-

ribedbyLietal.(15)ednativestarches

(50mg)weresuspendedin5mLof0.1Mphosphatesodiumbuffer(pH6.9)

uslicheniformisR-amylase(Sigma-

Aldrich)wasadded,withafinalconcentrationof0.01%(w/v).Theamy-

lolysiswascarriedoutinanovenat37hedesired

timeofhydrolysis,undissolvedresidueswereisolatedbycentrifugation

(10min,3000g),andthesupernatantwasmeasuredforsolubilizedcarbo-

iduesweredriedandtherecoveryyieldafterenzyme

hydrolysiscalculatedasabove.

lysisofisolatednativeandacidmodified

starchgranuleswascarriedoutonanXRD(D8,Bruker,Germany)

accordingtothepublishedmethod(16).Thesampleswereexposedtothe

nningregionofthediffraction

angle(2)wasfrom3to40withastepsizeof0.02andacounttimeof

specimenswerestoredinadesiccator,whereasaturated

solutionofNaClmaintainedaconstanthumidityatmosphere(relative

humidity(RH)=75%)for1weekat25Cbeforemeasurements.

-FTIRmeasurementwascarried

outaccordingtothemethod(17)ctra

wereobtainedusingaVarian7000FTIRspectrometerwithaDTGSdete-

ctorequippedwithaATRsinglereflectancecellcontainingagermanium

crystal(45incidence-angle)(PIKETechnologies,USA).Foreachmea-

surement,64scanswitha4cm

-1

resolutionwerecoaddedbeforeFourier

ctrumofwaterrecordedinthesameconditionwas

awerecorrectedbyabaselinein

theregionfrom1200to800cm

-1

beforedeconvolutionwasappliedusing

umedlineshapewasLorentzianwithahalf-width

of26cm

-1

rbance

valuesat1047,1022,and995cm

-1

wereextractedfromthespectraafter

watersubtraction,baselinecorrection,anddeconvolution.

Solid-State13CCP/-resolutionsolid-

state13CCP/MASNMRexperimentswerecarriedoutatB

0

=9.4Tona

responding13C

swerepackedina7mm

ZrO

2

rotorandspunatthemagicangle(54.7)with6kHzofspinrate.

1H-13CCP/MASspectrawererecordedwithacontacttimeof1.2msand

micalshiftswerereferencedtotetramethyl-

silane(TMS)lly,8000-12000transientswereaccumu-

specimenswerestoredinadesiccator,

whereasaturatedsolutionofNaClmaintainedaconstanthumidityatmo-

sphere(RH=75%)atmospherefor1weekat25Cbeforemeasurements.

RESULTS

dtheeffectof

enzymeoralkalinetreatmentonstarchcrystallinestructure

duringstarchisolation,weisolatedthenativestarchfrommature

ricegrainswithoutenzymeandalkalinetreatmentinwhichthe

isolatednativestarcheswereidentifiedtohavenodamageon

granules(datanotshown).InTRSrice,the

max

(maximumabso-

rptionwavelength)andbluevalueofiodine-starchcomplexwere

significantlyhigherthanthoseofwildtypeTQ,andasexpected,

theACofTRSnativestarchwasnearlytwice(58.32%vs29.98%)

asmuchasthatoftheisolatednativestarchfromwildtyperice

grains.

HighResistancetoAcidandEnzymeHydrolysisofTRSNative

1showstherecoveryyieldofnativestarchafter

em.,Vol.58,No.12,20107385

herTRSor

TQnativestarch,theresidualstarchwasgraduallydecreasedwith

thepassageofhydrolysistime,butafter20daysofacidhydro-

lysis,therecoveryyieldofTRSstarch(33.5%)wassignificantly

higherthanthat(6.9%)ofnormalTQstarch(Figure1a).The

highrecoveryofresidualstarchinTRSricewasalsoobserved

afterR-amylasehydrolysis(Figure1b).Duringthetimecourseof

hydrolysis,TQstarchwasmorereadilyhydrolyzedthanTRS

72h,onlyabout52%oftheTRSnativestarchwas

hydrolyzed,whereasnearly89%oftheTQnormalstarchwas

esultssuggestedthatTRSstarchhadamuch

higherresistancetoeitheracidorenzymehydrolysisthanTQ

starch.

TheXRDPatternofTRSNativeStarchSimilartoThatofPea

C-TypeStarch.

TheXRDpatternsofnativestarchesofTQ,TRS,

pea,RDpatterns

werecarefullycomparedwithknowndiffractionpatternsofA-,

B-,andC-typecrystallinity(1,2).Thenormalnativestarchfrom

TQricegrainshowedstrongreflectionat2about15and23

andanunresolveddoubletat17,182,whichwasverycloseto

thetypicalA-typeXRDpatterninmostordinarycerealstar-

ches(1,2).Thepotatostarchpresentedthestrongestdiffraction

peakataround172andafewsmallpeaksataround2values

of24,22,and15.Anadditionalpeakalsoappearedatabout5

2.These陈元方候袁公 spectraweretypicalcharacteristicsofB-typestarch

fromtubercrops(1,2).Thepeastarchhadbeenreportedtobea

typicalC-typecrystallinityrevealedbyXRD(1,2).Onthepea

starchXRDspectra,onlyonepeakappearedat232,whichwas

indicativeoftheA-typepattern,whilethepeakataround52

wasthecharacteristicofB-typepattern(1,2).Whencompared

withtheabovethreetypecrystallinity,theXRDpatternofTRS

-

ever,thepresenceofsomeadditionalA-typepeaksindicatedthat

Sstarchwas

oteworthythatthesca-

tteringintensitiesfor15and232diffractionpeaksdecreased,

whereasasharpreflectionpeakatanglesof202wasobserved

kof202wasatypicalamylose-lipid

complexdiffractionpeak(1,2),whichwasinagreementwiththe

resultofhighACinTRSstarch.

elopmentof

samplingdeviceslikeATR-FTIRcombinedwithproceduresfor

spectrumdeconvolutionprovidedopportunitiesforthestudyof

starchexternalregionstructure(17).Theoriginalanddeconvo-

lutedATR-FTIRspectraintheregion1200-900cm

-1

offour

dsat1045and

1022cm

-1

hadbeenlinkedwithorder/crystallineandamorphous

regionsinstarch,respectively(17).Theratioofabsorbance1045/

1022cm

-1

wasusedtoquantifythedegreeoforderinstarch

ityratiosof1045/1022and1022/995cm

-1

might

thereforebeusefulasaconvenientindexofFTIRdataincom-

parisonswithothermeasuresofstarchconformation(18).The

relativeintensitiesofFTIRbandsat1045,1022,and995cm

-1

wererecordedfromthebaselinetopeakheight,andtheratiosfor

1045/1022and1022/995werecalculatedasshowninTable1.

Onthebasisofboththespectraandcalculateddata,theATR-

FTIRcharacteristicsofTRSstarchwasmuchclosetothoseof

peaorpotatostarch,especiallyontheIRratioof1045/1022or

1022/995cm

-1

.InTRSstarchspectra,thebandat1022cm

-1

was

lesspronouncedthaninTQandpea,whichwassimilartothatin

potato(Figure3).TheseresultsalsoimpliedthatTRSstarchwas

aC-typestarch,amixtureofA-typeandB-typestarches,which

wasinqualitativeagreementwiththedatafromXRDanalysis.

id-state13C

CP/MASNMRpatternsfornativeTQandTRSstarchesare

ntialsimilaritieswereobservedinthe

onancesat61.8ppm

wasassignedtoC-6,andthelargesignalaround68-78ppmwas

ctraofnativestarches.

ooftheAbsorbances1045/1022and1022/995cm-1

for

NativeStarches

IRratio1045/1022(cm

-1

)IRratio1022/995(cm

-1

)

TQ0.691.67

TRS0.890.60

pea0.800.78

potato1.090.75

em.,Vol.58,No.12,2010Weietal.

collectivelyassociatedwithC2,C3,onanceat

81.8ppmwasassociatedwithC4site,andtheresonanceataround

theabove

peaks,theweakpeakappearedat94.3ppmcouldarisefromthe

ssignmentsoftheresonances

werebasedontheliteraturereports(19,20).

Tworemarkabledifferenceswereobservedbetweenthe13C

CP/,

theC1resonancesofTQstarchoccurredastriplets,whichwasa

typicalA-typecharacteristic(19,20).TheC1resonancesofTRS

starchalsooccurredasinconspicuoustriplets,especiallyweak

peakat101.4ppm,whichshowedthatTRSstarchwasaC-type

ond

peakat102.9ppmappearedonlyasashoulderonthedownfield

C-1resonanceinTQstarch,however,thatinTRSappearedasa

strongpeak,whichshowedthatthecontentofamylose-lipid

complexwashigherinTRSthanthatinTQ.

ChangeofCrystalTypeofTRSStarchduringAcidHydrolysis.

TheXRDpatternsofacid-modifiedTRSstarchesandtheir

ikingdifference

wasobservedforthepeakataround2valueof23amongthe

XRDspectraofTRSstarchafterdifferenttimeofacidhydrolysis.

NativeC-typestarchfromTRSgrainshowedonlyonebroad

peakat232.Thepeakbecamebroadfrom2to6daysofhydro-

lysisandthensplitintotwopeaksat22and24,whichwerethe

typicalB-typecharacteristics(1,2).Thedisappearanceofthecha-

racteristicA-typediffractionpeakandthedevelopmentoftypical

B-typediffractionpeakshowedthatthecrystaltypeofnative

TRSstarchmightchangefromtypicalC-typetoB-typeduring

acidhydrolysis.

DISCUSSION

ThecrystallinityofnativestarchcanbeclassifiedtoA-,B-,and

C-types(1,2).TheC-typestarchisusuallyacombinationofA-

andB-types,especiallyinmaize,withabout40%ofamylose(2).

Uptonow,inmostofthereportedhigh-amylosericemutants,

endospermstarchesarecharacterizedasaB-typepatternrevealed

byXRDanalysis(5,6).Inthisstudy,thestarchfromourdeve-

lopedhigh-amylosericeTRSwasdemonstratedastheC-typenot

onlybyXRDanalysisbutalsoconfirmedbythe13CCP/MAS

NMRandATR-FTIRtechniques.

13Csolid-stateNMRhasbeenemployedinexaminingthe

pectra,mostofthe

resonancescannotbedistinguishedorhavenotbeenassigned

amongtheA-,B-,andC-typestarches,buttheC-1carbon

atomshavechemicalshiftscharacteristicforeachtypestarch.

FortheA-typestarch,whichhasthreenonidenticalsugar

residues,theC-1peakregionisaclusterofthreepeaksat

∼102,101,and100ppm,B-typestarch,

whichhastwononidenticalsugarresidues,theC-1peaksignalis

aclusteroftwopeaksat∼101and100ppm,respectively.

BecauseC-typestarchhasthecharacteristicsofbothA-and

B-typecrystallinestructure,C-1spectraoftheC-typestarch

resonancesinthespectraofC-typestarchmainlydependonthe

relativeproportionsofA-orB-typecrystallinityinthesam-

ple(20).Ingeneral,theC-typestarchshowstripletsC-1spectra

iftheA-typecrystallinestructureispredominantinthesample,

andtwo-peakC-1spectraiftheB-typecrystallinestructureis

predominant(19,20).Inpresentstudy,theTRSstarchshowed

inconspicuoustripletsC-1spectra(Figure5),whichimpliedthat

TRSstarchexistedaC-typecrystallinitywithdominantA-type

crystallinestructure.

Thecrystallinepropertyofstarchcanbechangedbyacid

treatment,whichisveryhelpfultounderstandthefinestructure

ofstarchgranules(13).Inthepresentstudy,duringorafteracid

treatmentofTRSnativestarch,theXRDcharacteristicatA-type

diffractionpeakdisappearedwhilethetypicalB-typediffraction

henomenaindicatethatthecrystaltypeof

TRSstarchchangedfromtypicalC-typetoB-typeafteracid

ultwasquitedifferentfromotherreports

(13,16,21).Forexample,theacid-modifiedcornstarchesexhibit

thesamecrystallinetypeasthatofitsnativestarch(21),whilethe

crystaltypeofpeaandChines何其芳简介 eyamstarcheschangesfromC-type

toA-typeafteracidhydrolysis(13,16).Thisstructurechangeof

ourTRSstarchduringacidtreatmentmightbeduetothe

degradationofA-typecrystallinestarchfirstorfasterthanthat

ofB-typestarch.

Resistantstarchreferstotheportionofstarchand/orstarch

productsthataredifficulttodigestwhentheypassthroughthe

gastrointestinaltract(22).TheproportionofRSwillbeincreased

whenthedietstarchcarriesmoregranularstructurenaturally

resistanttodigestion(23).InourTRSgrains,thereisnotonly

highlevelofamylosebutalsoofRS(9,10).Asexpected,theTRS

starchshowedahigherresistancetoeitheracidorenzyme

hydrolysisthanthatofitswildtypestarch(Figure1).Thismight

Figure4.13CCP/MASNMRspectraofnativest意境有仙气的诗句 arches.

em.,Vol.58,No.12,20107387

beattributedtonotonlythehighamylosebutalsothespecial

viousexperiments

showedthatthestarchgranulesfromtheregularriceTQwere

organizedascompoundstarchesanddissociatedtoseparate

individualstarchesduringstarchisolation,whilestarchgranules

fromTRSwereorganizedassemicompoundstarcheswithathick

continuousbandencirclingtheentirecircumferenceofthegra-

nules(9).ThesizesofTRSsemicompoundstarchesarelarger

thanthatofTQindividualstarches,soTRSstarcheshadalower

rateofacidhydrolysisthanTQstarches,presumablyduetotheir

,highamylosestarch

wasreportedtobelesssusceptibletoacidhydrolysisthannormal

uggestedthatthehighlycompact

amorphousregionsinhighamylosestarchgranules,resulting

fromextensiveinterchainassociationsofamylosepolymers,pre-

ventedpenetrationofacidintothegranules(24).TRSstarchhad

highconcentrationofamyloseinboththehilumandencircling

band(9),whichmightpartlyexplainwhyitwashighlyresistantto

acidhydrolysis.

Itisreportedthattheamountofnativestarchhydrolysis

byamylaseisinverselyrelatedtotheamylosecontent(15).

Moreover,thedoublehelicesinstarchgranulesalways

prefertoformacrystallinestructurethatresiststoenzyme

hydrolysis(25-27).Bothcrystallineregionsanddouble

helicesthemselvescanincreasetheresistancetoamylase

obablyexplains,atleastinpart,whyhigh

amylosestarchresistsamylasedigestionmorethannative

orwaxystarcheseventhoughtheywerelesscrystalline

(25-27).Otherwise,theA-,B-andC-typesofstarchesshow

lly,

theB-orC-typestarchshowsmoreresisancetoenzyme

hydrolysisthanthatofA-type(25,28).Thismightbewhy

theTRSC-typestarchinthepresentstudyhadahighresistant

abilitytoR-amylasedigestion.

Inconclusion,thehigh-amyloseTRSstarchwasinvestigated

byusingXRD,13CCP/MASNMR,andATR-FTIRtechniques

andsubsequentlyconfirmedtobeC-typecrystallinestructure,

whichresultedfromthecombinationofbothA-typeandB-type

acidhydrolysis,thecrystaltypeofTRSstarch

atacouldaddtoour

understandingofnotonlythepolymorphstructureofricestarch,

especiallyofhighamylosestarch,butalsowhyhighamylose

starchmoreresistanttodigestion.

ABBREVIATIONSUSED

AC,amylosecontent;ATR-FTIR,attenuatedtotalreflec-

tance-Fouriertransforminfrared;13CCP/MASNMR,13C

cross-polarizationmagic-anglespinningnuclearmagneticreso-

nance;RS,resistantstarch;SBE,starchbranchingenzyme;TQ,

Te-qing(wildtypericecultivar);TRS,transgenicRSriceline;

XRD,X-raypowderdiffraction.

ACKNOWLEDGMENT

-ChengShifromKansas

StateUniversityforhelpfuldiscussions,andtothereviewersfor

valuablecommentsandcorrections.

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ReceivedforreviewJanuary29,dmanuscriptreceived

March22,edMay17,udywasfinancially

supportedbygrantsfromtheNationalNaturalScienceFoundationof

China(30828021,30300215),theMinistryofScienceandTechnology

(2006AA10A102,2009ZX08011-003B),theGovernmentofJiangsu

Province(BK2009186,06KJA21018),andtheChinaPostdoctoral

ScienceFoundation(2).