CarrotsofManyColorsProvideBasicNutritionandBioavailableActingasaFuncti - PDF document

CarrotsofManyColorsProvideBasicNutritionandBioavailableActingasaFunctionalFoodSaraA.ArscottandSherryA.Tanumihardjo ABSTRACT:Hippocrates,aphilosopherwholivedfrom460to359BCisoftenquotedassaying,“Letyourfoodbethymedicineandyourmedicinebethyfood.”Havinglivedjustshyofacenturyatatimewhenlifeexpectancies Carrots(DaucuscarotaL.)aremorethanaversatileorangevegetable.Originalcarrotswerepurpleandyellow,initiallyde-scribedinthe10thcenturyinIranandnorthernArabia(Simon2000).ThesecarrotsspreadeastandwestfromthiscentertobeknownacrosstheMiddleEast,NorthAfrica,Europe,andChinabythemid-15thcentury.Yellowcarrotswerepreferredinnorthern MS20091032Submitted10/16/2009,Accepted11/23/2009.AuthorsarewithUniv.ofWisconsin-Madison,InterdepartmentalGraduatePrograminNutri- carrotshavebeenpublished(Rubatzkyandothers1999;Simon2000).Thisreviewdescribesthenutritionalvalueoforangeaswellasthatofothercarrots,nutritionalimprovementsresearchershavemadethroughbiofortiÞcation,thebioavailabilityofpig-mentsfromcarrotsandtheirimpactonvitaminAstatus,and,Þnally,theputativehealthbeneÞtsattributedtocarrots.Wholefood-basedapproachestoenhancehealthbyutilizingfunctional ComprehensiveReviewsinFoodScienceandFoodSafety OH OH OH HO alpha-carotenebeta-carotenebeta-cryptoxanthinzeaxanthinlycopene Figure1---ThebasicstructuresofcommoncarotenoidsfoundandquantiÞedinhumanserum.Threeoftheseareprecursorstoretinol(vitaminA).-and-Carotenearefoundinorangecarrotsandsomevarietiesofredandpurplecarrots;luteinpredominatesinyellowandyellowÐpurplecarrots;andlycopeneisinredandthenovelpurpleÐorangeÐredcarrot(Millsandothers2008).foodsupply(Table1),upfrom18%in1970(USDA-ERS2009).In2007,carrotssuppliedanestimated37%oftheavailablefreshvegetable-carotene,themajorprovitaminAcarotenoidintheU.S.diet(Table2).Thepopularityofcarrotshasalsobeeninßuencedbytheintro-ductionoftheconvenientprepackagedÒcut&peelÓorÒbabycar-rots,Ómakingcarrotsaleadingvegetablesnackitem(USDA/ERS1997).AnnualU.S.percapitaconsumptionofcarrotsin2005was11.6poundsperperson.Fresh-marketcarrotsaccountfornearlythree-fourthsofallcarrotsconsumedintheUnitedStates,andat$0.10perserving,aresecondonlytopotatoasthemostaf-fordablevegetable(USDAERS2004).U.S.carrotconsumption,whichriseswithageandincome,varieswithrace(USDAERS2007).In2005,non-HispanicWhites(10.7pounds)andAsians(10.3pounds)atethemostcarrotsathome,followedbyHispan-ics(6.8pounds)andBlacks(4.9pounds).WorldproductionWhilecarrotsarenotamajorstaplefoodinanypartoftheworldduetolowenergydensity,theyareconsideredapri-maryvegetableinmanycountries.China,Russia,andtheUnitedStatesarethetop3producersofcarrotsglobally,contributingalmost50%oftheworldcarrotcrop(Table3).Productionandavailabilityofcarrots,andnearlyallhorticulturalcommoditiesthatcontaincarotene,areincreasingworldwide(Simon1990).Table4illustrateshowcarrotproductioninallmajorregionshasoutpacedpopulationincreases,althoughthisdoesnotnecessar-ilyreßectpercapitaavailabilityorconsumptionbecauseitdoesnotreßectmeasurableuses,suchasfarminputs,exports,endingstocks,andindustry.Theincreaseofworldcarrotproductionhasalsooutpacedtotalworldvegetableproduction(Rubatzkyandothers1999).Carrotsareatemperateregioncropandproduc-tionintropicalregionsismorelimited;however,advancementsinsubtropicalcarrotshavecontributedtoincreasedproductioncapacityinSouthAmerica(Simon2000).Carrotyieldshavere-mainedsteadyinmuchofAfrica,butproductiontrendsshowincreases(Table4),andexpandedproductionintheseareasisdesirable.NutritionalValue Thestoragerootofthecarrotisthemostcommonlyconsumedportionoftheplant,althoughthetenderyoungfoliageisocca-sionallyusedasastir-friedherbandinsaladsinChinaandJapan(Rubatzkyandothers1999).Theroots,however,arethesolefo-cusofthisreview.CarrotsdonotsupplyasigniÞcantamountofcaloriestothehumandiet,butdosupplynutritionintheformofphytochemicals,suchascarotenoids,anthocyanins,andotherCOMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... Table1---VitaminAandcarotenoidcontributionsfrommajorfoodgroupstoU.S.foodsupply,percapitaperdayintheUnitedStates. VitaminACaroteneFoodgrougRAEg(%) Meat,poultry,andÞsh3456(320)00(0Dairyproducts1890(175)143(2Eggs691(64)00(0Fatsandoils886(82)136(2Fruits259(24)510(7Legumes,nuts,andsoy00(00)07(0Vegetables2894(268)5596(82Whitepotatoes00(00)00(0Darkgreen,deepyellow2398(222)4746(69Othervegetables508(47)850(12Grainproducts540(50)48(0Sugarsandsweeteners00(00)00(0Miscellaneous184(17)360(5 2004datafromUSDA/ERSFoodAvailability(percapita)DataSystem2009.Representsdisappearanceoffoodintothemarketingsystemandincludessubstantialquantitiesofnonediblefoodportionsandfoodlosttohumanusethroughwaste,trimming,cooking,andspoilageinthehomeandmarketingsystem.Datatypicallyoverestimateamountsoffoodandnutrientspeopleactuallyingest.gRAE(1retinolactivityequivalent)gall-trans-retinol,12gdietarytrans-carotene,or24gotherdietaryprovitaminAcarotenoid.phenoliccompounds.Thegreatestnutritionalinterestincarrotsstemsfromtheirphytochemicalcontent,butresearchhasalsofo-cusedoncarrotsasasourceofÞber.InTable5,thenutrientcom-positionofcarrotsiscomparedtoothercommonlyconsumedvegetablesintheU.S.dietthatcontributetototal-carotenein-takebasedontheNationalHealthandNutritionExaminationSurvey(NHANES)2003to2004,allagegroups.Nutrientcontentofcarrotscanvarywithcultivar(Nicolleandothers2004),season(Horvitzandothers2004),environmentalconditions(Rosenfeldandothers1998),andmaturity(PhanandHsu1973).MacronutrientsandmicronutrientsCarrotrootisapproximately88%water,1%protein,7%car-bohydrate,0.2%fat,and3%Þber(USDA2008).Thecarbohy-dratefractionisalmostexclusivelysimplesugars,predominantlysucrose,glucose,andfructose,withasmallamountofstarch(USDA2008).CarrotscontributesigniÞcantlytodietaryvitaminAintakethrough-and-caroteneandmodestlytoothernutri-ents.A100-gservingofrawcarrot(about0.75cupchoppedcar-rot)contributesthefollowingpercentagesoftheRecommendedDailyAllowanceofafemaleaged19to30y:120%vitaminA(asretinolactivityequivalents),4.5%vitaminE,3%calcium,4%magnesium,7%potassium,and11%Þber.CarrotscontaintheBvitaminsthiamin,riboßavin,andniacininappreciablequantitieswhencomparedwithothercommonlyconsumedvegetables(Table5).Nicolleandothers(2004)foundthatpotassiumwasthemostabundantmineralin20cultivarsoforange,yellow,white,andpurplecarrots,withameanof579mg/100gfreshweight(FW)andarangefrom443to758mg/100gFW.Somemineralswerelessaffectedbycultivar,suchascalcium,whilemineralssuchascopperandzincvariedmorebycultivar.Moreover,ascalciumlevelsincreased,ironcontentalsoincreased.Therewasnocorrelationbetweencolorandmin-eralortraceelementcontents,butdarkorangecarrots(highcarotene)hadthehighestmineralcontent.CarrotÞberContentofdietaryÞberanddigestiblecarbohydratecanvarybetweencultivarsandalsoduringprocessingandstorage(Svanbergandothers1997).FoodreferencetablesandreportsinTable2---Supplyof-carotenefromselectedvegetablecropstoU.S.population. -caroteneavailaPercaercaavaila-carotene Vegetalecrog/d% Bellpeppers28215Broccoli25854Brusselssprouts0141Cabbage3Carrots3988968037Celery2511Collardgreens023320Sweetcorn3Cucumbers2410Kale029239Iceberglettuce86245Romaineand6413225leaflettuceOnions93000Potatoes17062Pumpkin2170215Snapbeans0949Spinach0856126Sweetpotatoes2192542320Tomatoes78485 Freshvegetables,retailweight.FromUSDAEconomicResearchService2009,calculatedastheresidualofcommodityÕstotalannualavailablesupplyaftersubtractingmeasurableuses,suchasfarminputs,exports,endingstocks,andindustrialuses.FromUSDA-NCCCarotenoidDatabaseforU.S.Foods---1998.Referencesrawvegetableconcentrationsexceptwherenoted.Reßectspercapitafreshvegetableavailabilitydatawhichdoesnotaccountforsomevegetablessuchasfreshgreenpeas;variousAsianvegetables,suchasbokchoy,turnips,andrutabagas;freshbeets;parsnips;leeks;scallions(greenonions);rhubarb;domestically-producedgreenhousevegetables;andotherspecialtyanddehydratedvegetables.Allsquashvarieties.Acornwintersquashonly,raw.Squash-caroteneconcentrationsvaryconsiderablybasedonvariety.theliteraturevaryconsiderablywithregardtoÞbervalues,likelyduetodifferentmethodsofÞberanalysis(Marlett1992).ReportsoftotaldietaryÞbervaluesrangefrom2.42%(RaniandKawa-tra1994)to6.4%(DaSilvaandothers2007).Otherpublishedvaluesinclude2.5%(Marlett1992),2.8%(USDA2008),3.4%(Svanbergandothers1997),3.63%(Souciandothers2008),and4.4%(RamuluandUdayasekhararao2006).TheinsolubleÞbers,celluloseandhemicellulose,constitutethegreatestportion(50%to92%)ofthetotaldietaryÞberwithaverysmallamountoflignin(4%).ThesolubleÞbersconsistoffermentablehemicellu-loseandpectin(Marlett1992)andconstitute8%to50%oftotalÞber.CarrotÞberhasbecomeofinteresttofoodprocessersduetothelargequantitiesofcarrotwastecreatedinthecut&peelcarrotandcarrotjuiceindustries.Carrotpomaceisthewetcarrotshavingsproducedfromcarrotprocessingwhichissubsequentlydriedtoformapowder.TheÞbercompositionofcarrotpomacecandifferfromwholecarrotdependingontheprocessingmethod.Carrotpomacemadefromcarrotpeelshadareducedinsolubletosolu-bledietaryÞberratiowhenblanchedbeforedrying(Chantaroandothers2007),increasingitsperceivedqualityasafunctionalÞberforaddingtootherfoods.Thewaterretentionandswellingca-pacitiesofcarrotpomacewererelativelyhighcomparedtootheragriculturalbyproductssuchasapple,pear,andorangepomace(Chantaroandothers2007).TherelativelyhighsolubledietaryVol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafetyTable3---Carrotproductionbycountry,top20produc- RankCountryProduction(MT) 1China83955002RussianFederation17300003UnitedStatesofAmerica16017904Poland9350005Ukraine7065006UnitedKingdom6771447Italy6415588Japan6300009Germany55500010Netherlands43000011France41780012Turkey38000013Mexico37851714India35000015Belgium32000016Indonesia30867517Belarus30600018Australia30256019Canada30145020Morocco300000 Source:FAO-ERS2005.Þberfraction,highpecticsubstancesinthepeels,ortheblanchingmightaccountforthisproperty.NawirskaandUklanska(2008)foundÒDolankaÓcarrottohavethehighestpercentageofsolubleÞberwhencomparedwithapple,cabbage,strawberry,blackcur-rant,andchokeberrypomace.However,previouslytheyfoundcarrotpomacetohavethelowesttotaldietaryÞberaswellasthelowestrelativepectinandsolublehemicellulosecomparedwithapple,cherry,blackcurrant,andpearpomace(NawirskaandKwasniewska2003).Differencesmayberelatedtodiffer-entindustrialmethodsofpomaceproductionaswellascarrotvariety.The-carotenecontentofcarrotpomacewasreducedap-proximately19%byblanchingandbydrying,althoughlow-temperaturedryingreducedthe-carotenemore(57%)thanthehigher-temperaturedrying(46%)likelyduetoincreaseddrytime(Chantaroandothers2007).Totalphenoliccontentandtotalan-tioxidantactivityofcarrotpomacewerebothreducedbyblanch-ingandincreaseddryingtemperatures.TheseresultssuggestthatthenutritionalqualityofafunctionalÞberobtainedfromcarrotpomaceisrelatedtotheprocessingmethod.PigmentsandOtherCompounds Carotenoidsareresponsiblefortheyellow,orange,andredcolorsofcarrots,whileanthocyanins,aclassofpolyphenoliccompounds,areresponsibleforthecolorofpurplecarrots.AllofthesepigmentshavebeenstudiedfortheirhealthbeneÞts,in-cludingprotectionfromcertaincancersandcardiovasculardis-ease,andconsumerinterestinnaturalwholefoodsrichinthesecompounds,oftenreferredtoasÒfunctionalfoods,Óisgrowing(HaslerandBrown2009).Modernvegetablebreedershaveiniti-ateddevelopmentofsomecoloredcarrotbreedinglines(Simonandothers1997;Lazcanoandothers2001)toincreasethenu-tritionalqualityandvisualappealofthefoodsupply.Carotenoidsareagroupofphytochemicalsthatcompriseafamilyofover700compoundsinnature(Brittonandothers2004)andareresponsibleforthepigmentationinmanyfruitsandvegetables.ThosethatpredominateandareoftenquantiÞedinhumanserumincludelutein,zeaxanthin,lycopene,and-and-carotene(Figure1).Theyaresynthe-sizedexclusivelyinplantsandserveasthesourceforallanimalcarotenoids.Carotenoidsareaccessorylight-harvestingpigmentstochlorophyllinthechloroplastsofphotosynthetictissuesandinthechromoplastsofnonphotosynthetictissuessuchasfruits,ßowers,andtherootsofcarrots.Otherfunctionsofcarotenoidsinplantsincludephotoprotectionbyquenchingtheexcessenergyofexcitedchlorophyllorsingletoxygen(Gross1991),andalsoasanattractanttopollinatorssuchasinßowers.Thecarotenoidsincarrotrootsmaylikelyservenoneofthepreviouslymen-tionedpurposesandmaybetheresultofmutation(Gross1991;Simonandothers2008).Carrotrootcarotenoidsoccuraspure-pigmentcrystalsinchromoplasts.Eachcrystalissurroundedbyamembranetoformacarotenebody(Ben-Shaulandothers1968).Thecrystallinenatureofthecaroteneincarrotsnega-tivelyimpactsbioavailability(Zhouandothers1996).Forcom-parison,inpeppers,pumpkin,andfruit,carotenoidsarefoundincarotenoid-carryinglipiddropletsofglobulouschromoplasts.Ingreenleaves,thecarotenoidsarelocatedinthephotosystemsoftheinnerchloroplastmembranesandaredirectlyassociatedwithlipoproteinsandlipids.Carrotrootsarerichincarotenoids.Sixcarotenes(-zeacarotene,andlycopene)canberoutinelyseparatedandquantiÞedintypicalanddarkorangecarrots(Si-monandWolff1987).ThepredominantcarotenoidsaretheprovitaminAcarotenes,thatis,-and-carotene,accountingfor13%to40%and45%to80%ofthecarotenoidsinorangecarrots,respectively(SimonandWolff1987;Gross1991).EarlyTable4---Worldcarrotproduction1970Ð2003. NorthandCentralSouthYearWorldAfricaAmericaAmericaAsiaEuroeOceania 19707908214109928117763049117198010499383132846526773591141199013696558177363040034304193204899132619950816374943502332110542674982108017484327ProductionchangePopulationchange FAOProductionYearbookStatistics.Productionvalues1000Mt.ChinaandAsiaUSSRnotincludedin1983estimate;Nigerianotincludedbefore1985.Productionvaluesareaverageofthepreceding3y.FormerUSSRrepublicsincludedinappropriateEuropeanorAsiangrouping.Changeincarrotproductionfrom1970to2003,percent.Changeinregionpopulationfrom1970to2003,percent.Source:UN2009.COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... Table5---Nutrientcompositionofseveralcommonfruitsandvegetablesthatcontributetointakeoftotal-caroteneforcomparisontocarrot. g/100gfreshweightmg/100gfreshweight VitA VegetaleWater%100gCHOProteinFatFierAshRAEVitCVitBVitBNiacinVitECaPKNaMgFe Carrot88419.61.00.22.81.08355.90.070.060.980.66333532069120.3Sweetpotato778620.11.60.13.01.07092.40.080.060.560.26304733755250.6Cantaloupe90348.20.80.20.90.6169370.040.020.730.0591526716120.2Tomatoes94183.90.90.21.20.542130.040.020.590.5410242375110.3Spinach91233.62.90.42.21.7469280.080.190.722.03994955879792.7Lettuce,iceberg96143.00.90.11.20.4252.80.040.250.120.1818201411070.4Broccoli91285.23.00.33.30.9150930.060.120.641.45486632527250.9Lettuce,romaine95173.31.20.32.10.6436240.070.070.310.1333302478141.0Collards90305.72.40.43.60.9333350.050.130.742.26145101692090.2Watermelon91307.50.60.10.40.2288.10.030.020.180.007111121100.2 Source:USDANatl.NutrientDatabaseforStandardReference:http://www.ars.usda.gov/nutrientdata gRAE(1retinolactivityequivalent)gall-trans-retinol,12gdietaryall-trans-carotene,or24gotherdietaryprovitaminAcarotenoid.varietiesoforangecarrotsintheUnitedStateshadcarotenecon-tentsintherangeof70ppm,butbreedingefforts,throughphe-notypicrecurrentselectionofrootswithadarkerorangecolor,increasedthatvalueto90ppmbythe1970s(Simonandoth-ers2009).Modernspectrophotometricdetectionofhighlevelsofrootcaroteneshasallowedbreederstopushtheconcentrationabove130ppmintypicalorangeanddarkorangecarrots,thatis,high-carotenemass,canreach500ppm(SimonandothersTotalrootcarotenoidcontentcanvarysigniÞcantlybetweencultivars(SimonandWolff1987;Heinonen1990;Nicolleandothers2004;Grassmannandothers2007)andisthemajorsourceofvariationinreportedcarrotcarotenoidconcentrations.Ad-ditionally,thegrowingseason,soil,maturity,andgeneticfac-torsalsoinßuencecarotenoidcontentofcarrots(HartandScott1995;Rosenfeldandothers1997,1998).-Caroteneishetero-geneouslydistributedacrossthecarrotrootsection,andinallyellow,orange,red,andpurplecarrotclassesisgenerallyhighestintheouterroot,orphloem,andmuchlowerinthexylem(core)(Baranskaandothers2006).Carrotrootsofthehigh-carotenemassline,however,exhibithighconcentrationsofinthexylem(Baranskaandothers2005).SpeciÞcandsensi-tiveHPLCtechniqueshavegreatlyimprovedcarotenoidanalysisofplanttissues;however,differentsamplepreparationandex-tractiontechniquescontributevariabilitytoreportedvegetablecarotenoidconcentrations(Scottandothers1996;KimuraandRodriguez-Amaya1999).Table6listssomecurrentreportedcarotenoidconcentrationsoforangeanddarkorangecarrots,aswellasothercoloredcarrots.Inadditionto-and-carotene,sometypicalorangecarrotshavedetectablelevelsoflutein,althoughmodernbreedingeffortshaveattemptedtoeliminatetheyellowcore,whichwasperceivedasadefect(Simonandothers2009).LuteinhasnoprovitaminAactivity,butisfoundlocalizedinthemacularregionoftheeyeinhumansandmayhaveimportanceineyehealthandprotec-tionfromage-relatedmaculardegeneration(TanumihardjoandYang2005).Luteinisanimportantpigmentinyellowcarrotsandtheycancontainfrom1to5ppmlutein,butonlyverylowlev-elsofcarotenes(2to4ppm)comparedtoorangecarrots(95to311ppm)(Table6).Sunandothers(2009)reportedthatwhiteandyellowcarrotscontainedthelowesttotalcarotenoids(2and6ppmFW,respectively)comparedwithdarkorange(160ppmFW)orange(98ppm,FW)purpleÐyellow(92ppmFW)red(73ppmFW)andpurpleÐorange(40ppmFW)carrots.Nonethe-less,luteinand-carotenefromyellowcarrotsarebioavailablewhenfedtohumansinprocessedforms(MolldremandothersLycopeneisthecarotenoidprimarilyresponsibleforthecolorofredcarrots.WhilelycopenealsohasnoprovitaminAactivity,eventhoughitisahydrocarbon,itisthemostpotentantioxidantinvitroofallthecarotenoidsfoundinappreciableamountsinhumans(DiMascioandothers1989)andhasbeenimplicatedinprotectionfromcertaincancers(Giovannucci2002).Investi-gationsofthelycopenecontentofredcarrotsaremuchfewerthanforthecarotenoidcontentoftypicalorangecarrots,inspiteofthefactthatredcarrotsarecommonlyconsumedinIndia(Tan-umihardjoandothers2008).Reportedlycopenecontentofredcarrotsrangesfrom50to100ppm(Table6),whichissimilartoorgreaterthantypicaltomatoconcentrations.Theyear-roundav-erageofred,ripe,rawtomatoesis30ppmwitharangeof9to42ppm(USDACarotenoidDatabase1998).Redcarrotcultivarsalso-carotene,andlutein,theamountsvaryingbytypeofredcarrot.Agroupofpolyphenolicpigmentscalledanthocyaninsareresponsibleforthecolorofpurplecarrots.Currentcultivarsofpurplevarietiesincludesolidpurplecarrots,oftenreferredtoVol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafetyTable6---Carotenoidcontentofrawcarrotsofdifferent Total( Carrotcolor-Carotene-CaroteneLuteinLyco Orange221282.6ndB27690.40.6C4069nmnmD13to3132to660.6to1.8nmE47.147.5ndndH10to2218to38nmnmI57to7045to524to5ndDarkorangeA311854.417B451130.70.9D75.81721.0nmF96to192215to311NRNRYellowA0.51.85.1ndB0.23.62.40.04CtrtrnmnmDnd3.31.4to2.3nmITrTr5to10ndA1.13.43.261B0.2220.250Indto435to40Tr-385to100I2to32to39to10ndle–orangeA412311ndB10281.10.2C87161nmnmI62to100658to10ndle–yellowB21530.4Dnd3.1to3.81.8to2.2nmAnd0.060.09ndB0.050.341.70.04CndndnmnmIndndtrnd Carotenoidconcentrationsofrawcarrot,ppm.References:ASurlesandothers(2004);BSunandothers(2009);CAlasalvarandothers(2001);DNicolleandothers(2004);EEL-Qudah(2009);FSimonandothers(1989);HRodriguez-Amayaandothers(2008);IGrassmannandothers(2007).notdetected;nmnotmeasured;NRnotreported.asÒblackcarrots,Óandcarrotswithpurplephloemandwhite,yellow,ororangexylem(core).Purplecarrotswithawhitecorecontainverylowlevelsofcarotene(4to6ppm),whereaspurpleÐorangecarrots(38to130ppm)cancontainasmuchormoretotalcaroteneastypicalorangecarrots(Grassmannandothers2007)(Table6).Thecontinuedresearchincarrotcul-tivardevelopmenthasproducedanovelpurpleÐorangeÐredcultivarthatcontainsapproximately40ppmcarotenesand62ppmlycopene(Millsandothers2008).Whilethesecolorfulcar-rotsarestillanoveltyformodernconsumers,theysharetheßavorattributesoftheirorangecounter-parts(Alasalvarandoth-ers2001)andaregenerallywell-likedbyconsumers(Surlesandothers2004).Colorfulcarrotswithavarietyofpigmentshavethepotentialtocontributetothediet,notonlytheprovitaminA O HO OH OR O HO HO O OH OH cyanidininapoyl-e-6"-feruloyl-e-6"-(4-coumaroyl)chloro Figure2---Themainphenoliccompoundsfoundincarrotsarechlorogenicacids.PolyphenolsthatarecommonlyquantiÞedinpurplecarrotsinclude5anthocyanins.Thebasicanthocyaninbackboneiscalledcyanidin.TheRgroupsarevariousglycosides.carotenes,butalsothebeneÞcialhealtheffectsoftheirrespectivePhenoliccompoundsCarrotscontainphenoliccompoundswithasinglearomaticringknownasphenolicacids.Themainphenoliccompoundsfoundincarrotsarechlorogenicacids,whicharehydroxycin-namicacidderivativesformedbytheesteriÞcationofcinnamicacids,suchascaffeic,ferulic,andp-coumaricacids,with(-)-quinicacid.Thesecompoundscontributetotheorganolepticpropertiesoffreshandprocessedcarrots(Rubatzkyandothers1999).Recently,consumptionofchlorogenicacidsincoffeehasbeenassociatedwithreductionsinseveralchronicdiseases(Hig-donandFrei2006).Studiesestimateapproximately30%intesti-nalabsorptionofchlorogenicacids(Farahandothers2008)andextensivemetabolismoftheremainderbycolonicmicroßora(Olthofandothers2003).Alasalvarandothers(2001)identiÞed11differentphenolicacidsinorange,purple,yellow,andwhitecarrots.Chlorogenicacid(Figure2)predominatedineverycolorcarrot,itsconcentra-tionbeing54.1,8.5,4.5,and4.4mg/100ginpurple,orange,white,andyellowcarrots,respectively.TotalconcentrationofalltheidentiÞedphenolicacidswasgreatestinthepurplecarrotsandfollowedthesamecolororderaschlorogenicacidconcen-tration.Grassmannandothers(2007)foundasimilarrankingbycoloroftotalphenolicsandincludedredcarrots,whichhadaslightlyhigherconcentrationthanyellow,orange,andwhite.To-talphenoliccontentincarrotsareoftenexpressedingallicacidequivalentsasmeasuredusingtheglobalFolinÐCiocalteumethodCOMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... (SingletonandRossi1965),whichalsoreactswithßavonoidssuchastheanthocyaninsinpurplecarrots.Therefore,purplecar-rotsarethemostconcentratedintotalphenolicsbythismethodwhencomparedwiththeothervarieties.AccordingtoCaetanoandLeal(2006),carrotswereclassiÞedasaÒlowphenoliccontentÓvegetable(100mgcatechinequiv-alents/100gFW),havingjust12.9mg,butwerejoinedinthatgroupingbytomato(30.8mg),greenbean(36.8mg),redonion(73.9mg),andspinach(82.1mg).Incomparison,collardgreens(348.3mg)andredcabbage(213.1mg)wereintheÒhighphe-noliccontentÓgroup.Chuandothers(2002)distinguishedfree,orsoluble,phenolicsfromboundphenolicsbysubmittingveg-etableextractresiduestoadditionalbasedigestion.Theyfoundthatbroccolihadthehighesttotalphenolics,followedbyspinach,onion,redpepper,carrot,cabbage,potato,lettuce,celery,andcucumber.Onaverage,thefree-phenoliccontentwas76%ofthetotalphenolics.Carrotshad37.6%asboundphenolics,secondonlytopotatowiththehighestbound-phenoliccontent.Boundphenolicsarefoundmostlyinesterformandareassociatedwithcell-wallcomponents(Kangandothers2008).Theylendcross-linkingabilitytoconferstructuralstabilityinthecell-wallma-trix.Theseboundphenolicsmaysurviveuppergastrointestinaldigestionandbereleasedinthecolonwheretheymayhavepositivehealtheffects.Chuandothers(2002)estimatethatcar-rotsandpotatoescouldreleaseapproximatelyone-halfoftheirphenolicsinthecolon,comparedwithone-fourthinmanyothervegetables.PhenoliccompoundsandantioxidantcapacityGajewskiandothers(2007)foundpurplecarrots,richinan-thocyanins,tohavegreaterinvitroantioxidantcapacitythanorangeandyellowcarrotsasmeasuredfrommethanolicextract,whichmaymoreefÞcientlyextracthydrophilic(phenolics),ratherthanhydrophobic(carotenoids)compounds.Invitrocapacity,asmeasuredbyoxygenradicalabsorbancecapacity(ORAC),showedlittledifferencebetweendifferentcoloredcar-rots,includingorange,yellow,purple,andwhite,whichmaybeduetoanextractionmethodoptimizedforhydrophobic,thereforecarotenoid,extraction(Nicolleandothers2004).Bothpheno-liccompoundsandcarotenoidsarestronginvitroPhenolicfractionsfromredwinearepotentfreeradicalscav-engers(Ghiselliandothers1998)and-caroteneisconsideredastrongquencherofsingletoxygen(Schaferandothers2002).Sunandothers(2009)evaluateddifferentcoloredcarrotsfortheirrelativeantioxidantcapacityindex(RACI)byassessingbothhydrophilicandhydrophobicextracts.Theyfoundthatchloro-genicacidwasamajorinvitroantioxidantindifferentcoloredcarrotsandthatthephenolics(phenolicacidsandßavonoids)madegreatercontributiontothetotalantioxidantcapacitythancarotenoids.Carotenoidswerethemajorantioxidantsinthehy-drophobicextracts.Grassmannandothers(2007)alsofoundthatantioxidantcapacityofthehydrophilicextractcorrelatedwithto-talphenolics,andthatofthehydrophobicextractcorrelatedwithcarotenoids.Thesestudiesdemonstratethatdifferentmethodsandextractionproceduresmeasuredifferentcompoundsandthatstudiesthatexplicitlymeasurebothhydrophilicandhydropho-bicextractsmayprovidegreaterinformation.Itisimportanttokeepinmindthatinvitrostudiesdonotnecessarilyreßectvivoactivityofmanyofthesecompounds.Researchisbegin-ningtouncoverthenumerousmetabolitesformedfrommanyingestedparentcompounds(Kayandothers2005)andemerg-ingevidenceispointingtowardsmechanismsofactionthatgobeyondthemodulationofoxidativestressandmayinvolvecellmodulationthroughintracellularsignalingcascades(Crozierandothers2009).PhenoliccompoundsasphytoalexinsPhenoliccompoundsmayhavearoleinplantresistancetofungalandbacterialagents.Theyaccumulateincarrotsinresponsetocold,injury,orethyleneexposure(Rubatzkyandothers1999).Totalphenolics(Alasalvarandothers2004),chloro-genicacid,isochlorogenicacid(HagerandHoward2006),trans-caffeoylquinicacid,andpara-hydroxybenzoicacidanditsesters(Babicandothers1993)increaseduringstorageiftheßeshiswounded,afterprocessingsuchasshredding,orun-derexposuretoethylene.Theincreasesinphenoliccontentoffresh-cutcarrotsduringstorageareassociatedwithincreasedphenylalanineammonialyase,awound-inducedenzyme.Com-poundsproducedbyplantsinresponsetopathogeninfectionorwoundingarereferredtoasphytoalexinsbecausetheirpro-ductionmaybeadefensemechanism.Thebittercoumarincompound6-methoxymellein,aphenolicoxidationproduct,preferentiallyaccumulatesinthetipsofwhole,unwoundedcar-rotsinresponsetoethylene(TalcottandHoward1999)andmaybeduetoahigherproportionofperidermtoxylem/phloematthetip,whererespiration,andthusphytoalexinproduction,wouldbeexpectedtobegreatest.Additionally,thetipistheleadinggrowthtissueandwouldbeexpectedtohavegreaterphytoalexinAnthocyaninsarewater-solublepigmentsthatcompriseagroupofover600compoundsthatprovidethered,purple,andbluecolorsofmanyfruits,vegetables,ßowers,andgrains.Theyareoftenusedinindustryasnaturalcolorantsinfoodsandbev-erages.Inplants,anthocyaninsfunctiontoprovidephotoprotec-tion,scavengefreeradicals,andattractanimalsforpollination(NeillandGould2003;Gould2004).Dietaryanthocyaninsmayplayaroleinhealthpromotionandprotectionfromcardiovas-culardisease(Reed2002;Mazza2007)andcancer(Hou2003;WangandStoner2008)byactingasdietaryantioxidants,reduc-inginßammationandlipidoxidation,causinginductionofanti-inßammatoryandvasoprotectiveeffects,phaseIIenzymes,andapoptosis.Forexample,anthocyanin-richextractsfromchoke-berryandbilberrywereshowntorelaxcoronaryarteryringsiso-latedfrompighearts(BellandGochenaur2006).Anthocyaninshavealsobeenimplicatedinimprovedbrainandmemoryfunc-tions(Shihandothers2009).Anthocyaninsarepolyphenoliccompoundsthatarecom-prisedofananthocyanidinbackbone,2-phenylbenzopyrylium,alsoreferredtoastheßavyliumcation.The6commonan-thocyanidinbackbonesarecyanidin,malvidin,delphinidin,peonidin,petunidin,andpelargonidin.Thesebackbonescanvaryinthenumberandpositionofhydroxylgroups,methoxylgroups,andtype,position,andnumberofattachedsugarmoleculeswhichmayalsobeacylatedbyvariousaromaticoraliphaticacids.Theprimaryanthocyaninsfoundinpurplecarrots(sometimesreferredtoasblackcarrots),arederiva-tivesofcyanidin(Figure2),butpelargondinandpeonidinglycosideshavealsobeenidentiÞed(Kammererandothers2003).Theanthocyaninsofpurplecarrotshavenodirecteffectonßavor(Simon2000;Surlesandothers2004).AnalysisofpurplecarrotshasidentiÞed5mainanthocyaninderiva-tives:cyanidin-3-(2-xylose-6-glucose-galactoside)(Cy3XGG),cyanidin-3-(2-xylose-galactoside)(Cy3XG),cyanidin-3-(2-sinapoyl-glucose-galactoside)(Cy3XSGG),cyanidin--feruloyl-glucose-galactoside)(Cy3XFGG),andcyanidin-3-(2-(4-coumaroyl)glucose-galactoside)(Cy3XCGG)(Glabgenandothers1992).Totalanthocyaninconcentrationintherootsofpurplecar-rotscanvarywidelybetweencultivarsandevenwithinacul-tivarbasedonthedegreeofrootcoloring(LazcanoandothersVol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafety2001;Kammererandothers2004).Reportsoftotalanthocyanincontentrangefrom0mg/100gFWinorangecarrotsto350mg/100gFWindarkpurplecarrots(Kammererandothers2004;Simonandothers2008;Sunandothers2009).Forcomparison,totalanthocyaninconcentrationisapproximately113mg/100ginblueberries,117mg/100gincherries,and48mg/100ginraspberries(USDA2003).ThedailyintakeofanthocyaninsintheU.S.dietisestimatedtobe12.5mg/day(Wuandothers2006).TheanthocyaninsinpurplecarrotscontributedsigniÞcantlytoinvitroantioxidantcapacity(Sunandothers2009).Dietaryanthocyaninsappeartohavelowbioavailability,re-coveriesinurinerangingfrom0.004%to0.1%ofintake(Manachandothers2005;McGhieandWalton2007).Studieswithwholerawandcookedpurplecarrotsandcarrotjuiceinhu-mansdemonstratedthatpurplecarrotanthocyaninsarebioavail-ableandareabsorbedintact,butwithlowefÞciency(Kurilichandothers2005;Charronandothers2009).Thesefeedingstud-iessuggestthatsaturationofabsorptionofthecyanidin-basedanthocyaninsfrompurplecarrotsoccursbetweenintakesof250and350mol(equivalentto150to250gcarrot).Thepercentagerecoveryofnonacylatedanthocyaninswasgreaterthanacylatedanthocyaninsinstudieswithbothwholecarrotsandcarrotjuice,suggestingthatacylation,andnottheplantmatrix,inßuencesbioavailability.OtherrelevantcompoundsManycompoundscontributetocarrotßavorandsomeofthesemaycontributetoeffectsonhumanphysiology.Thecharacteris-ticÒfreshcarrotÓßavorhasbeenattributedtothevolatilecom-poundsmono-andsesquiterpenes,andalsotosugars(Simonandothers1980).Terpenesgenerallyimpartaharshorbitterßavorandtheseßavorattributeswereseentoincreasedirectlywithterpenecontentindifferentcarrotgenotypes(Simonandoth-ers1980;Kreutzmannandothers2008).Terpinolene(Figure3)appearstobethemostabundantvolatileandthecontentofto-talvolatilesvariesgreatlybetweengenotypes(Simonandothers1980;Alasalvarandothers1999,2001;HabeggerandSchnitzler2005;Kreutzmannandothers2008).Whilepurplecarrotculti-varsinonestudy(Alasalvarandothers2001)arereportedtohaverelativelylowterpinolenecontent,theamountsinothercolorsvarywidely.Cookingcarrotscanreducevolatilecontentby70%to95%(SimonandLindsay1983;Alasalvarandothers1999).Agroupofcompoundscalledpolyacetylenesisresponsibleforthebitteroff-ßavorofcarrots(CzepaandHofmann2004).ThesecompoundsarewidelydistributedintheApiaceaefam-ilyofplantsthatincludescarrot,celeriac,parsnip,andparsley,andhavebeenidentiÞedasphytoalexins,toxiccompoundspro-ducedbyplantsinresponsetoattackbypathogensorotherstresses(ChristensenandBrandt2006).Theyarepotentskinsen-sitizers,irritants,andalsoneurotoxicathighconcentrations,andhavetraditionallybeenviewedastoxicants.Morerecently,poly-acetyleneshavebeenimplicatedasbioactivecompoundswithpotentialeffectsonhumanphysiologyanddisease(Hansenandothers2003;ChristensenandBrandt2006).FourpolyacetyleneshavebeenidentiÞedincarrotroot,themostabundantarefalcarinol,falcarindiol,andfalcarindiol3-acetate(Figure3)(ChristensenandBrandt2006).Freshweightconcentrationsofpolyacetylenesrangefrom20to100mg/kg(CzepaandHofmann2004;Zidornandothers2005;ChristensenandKreutzmann2007).Concentrationsofcarrotpolyacetylenesvarybycultivarandappeartobelocalizedtothecarrotrootphloem(CzepaandHofmann2004;Baranskaandothers2005).Higherpolyacetyleneconcentrationswerefoundincarrotswithhighercarotenoidlevels,andyellowcarrots,withlowerlevelsofcarotenoids,hadlowerlevelsofpolyacetylenesthanorangecarrots(Baranskaandothers2005).Boilingcarrotsfor12min H OH terpinolenefalcarinolfalcarindiolfalcarindiol 3-acetate Figure3---Chemicalstructuresofterpinolene,falcari-nol,falcarindiol,andfalcarindiol3-acetate.Thesecom-poundsareinpartresponsibleforthecharacteristiccarrotßavor.reducedthefalcarinolcontentby70%comparedwithrawcarrots(Hansenandothers2003).Invitrostudiessuggestthatcarrotpolyacetyleneshaveantiin-ßammatoryactivityinmacrophages(Metzgerandothers2008),biphasicstimulatoryandcytotoxiceffectsonprimarymammaryepithelialcells(Hansenandothers2003),andcytotoxicactiv-ityagainstanumberofcelllines(Zidornandothers2005).Ratsfed10%freeze-driedcarrotpowderwith35gfalcarinol/gfeedorfalcarindiolextracthadreducednumbersoflargerclasscolonicaberrantcryptfoci(Kob¾k-Larsenandothers2005).Totestbioavailabilityinhumans,astudywasconductedinmalesubjectswith300,600,and900mLcarrotjuicethatcontained16,33,and49molfalcarinol,respectively(Brandtandothers2004).All3treatmentsresultedinarapidincreaseinplasmafalcarinolwithin30minofthetest-mealandpeakconcentra-tionsat2h.Themaximumconcentrationsinplasmawerewithintherangethatinvitrodataindicatepotentialstimulatoryeffects(Hansenandothers2003);however,plasmaconcentrationswerebelowthelevelsreportedtoelicitinvitroeffectsoncytotoxicity(Hansenandothers2003;Zidornandothers2005),antioxidantorphase2enzymeinduction(Ohnumaandothers2009),andantiinßammatoryeffects(Metzgerandothers2008).Whilesomeresearchershavesuggestedthatepidemiologicdatalinkingre-ducedlungcancerincidencewithcarrotscouldberelatedtofalcarinol(Brandtandothers2004),moreresearchoninvivoeffectsofthiscompound,whichtraditionallywasconsideredatoxicant,iswarranted.Manyothercompoundscontributetotheuniqueßavorofcarrots;andconsumersgenerallyprefercarrotswithhighsugarandlowvolatileterpenoidcontents(Rubatzkyandothers1999).CarotenoidsandphenoliccompoundsarethepredominantCOMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... phytochemicalsinallvarietiesandcolorsofcarrots;however,otherminorcompoundsandtextureinßuencetasteandpossiblythehealthpropertiesofthispopularandtastyvegetable.Carrots,likeotherwholefoods,areapackagethatoffersmorethanthesumofitscomponentcompounds.Food-BasedApproachestoIncreaseCaroteneIntake TworelatedmethodstoincreaseprovitaminAcarotenoidin-takearetopromotedietarydiversiÞcationthroughtheinclusionofmorevegetablesandtoincreasetheamountofcarotenoidperservingofvegetable.IncreasingvegetableorprovitaminAcarotenoidintakenotonlyimprovesvitaminAstatus,butwillalsocontributetoimprovedhealthoutcomes.VitaminAdeÞciencyIndevelopingcountries,wherevegetablecarotenoidscon-tributegreaterthan80%oftheavailablevitaminA(FAO-WHO2001),vitaminAdeÞciency(VAD)isamajorpublichealthprob-lem.ThehighestprevalenceofVADisinpartsofAfricaandSoutheastAsia.VADaffects140millionpreschool-agedchil-drenand1.2to3millionchildrendieeachyearasaresult(UN-SCN2004).AccordingtotheWorldHealthOrganizationÕsGlobalDatabaseonVitaminADeÞciency(WHO2009),45and122countrieshaveapublichealthproblemwithregardtoclin-icalandsubclinicalvitaminAdeÞciency,respectively.ClinicalVADinvolvesoverteyesigns,whilesubclinicalVADismuchhardertodiagnose(Tanumihardjo2004).Theoretically,sufÞcientvitaminAexistsinthefoodsupplytomeetglobalneeds.How-ever,inequitabledistributionandpovertyaremajorhindrancestoadequatevitaminAaccesscontributingtoVAD.Indeed,povertycontributestoinadequatedietaryqualitywhichleadstomicronu-trientdeÞciency(Tanumihardjoandothers2007).Inthesepopu-lations,food-basedstrategiesthatpromotedietarydiversiÞcationarecriticalinasustainableefforttoeliminateVAD(UnderwoodAlthoughpartoftheU.S.populationmightbegettingtoomuchpreformedvitaminAthroughmultivitaminsandfortiÞedfoods(AllenandHaskell2002;PennistonandTanumihardjo2003),theDietaryGuidelinesAdvisoryCommittee(2005)identiÞedvitaminAasashortfallnutrient,alongwithvitaminsCandE,calcium,magnesium,potassium,andÞber.Shortfallnutrientsarethoselikelytobeconsumedininadequateamounts.AccordingtotheContinuingSurveyofFoodIntakebyIndividuals(CSFII),1994to1996,theprobabilityofadequateintakeforvitaminAinmenandwomenwas47%and48%,respectively.BasedonNHANES2001to2002,44%ofAmericanshaveaninadequateintakeofvitaminAfromfood(Moshfeghandothers2005).AlthoughestimatesofinadequacybasedonserumvitaminAconcentra-tionsfromNHANES1988to1994arelow(Ballewandothers2001;Gillespieandothers2004),serumconcentrationsarenotasensitivemeasureofstatusandmayunderestimateinadequacy,especiallyinlow-incomegroups(Spannaus-Martinandothers1997).TheDietaryGuidelinesrecommendincreasedfruitandvegetableconsumptionfortheshortfallnutrientsvitaminsAandCandmagnesium.BiofortiÞcationoffoodscanincreasethenutritivevalueofthefoodsupply(Tanumihardjoandothers2008).Leadersofnutrition-orientedplantbreedingprogramshavetakenthisap-proachthroughincreasingselectionofnutrient-richcultivars.PlantbreedingtoincreasetheprovitaminAcontentofselectedfruits,vegetable,andstaplecropsisastrategythatcouldim-pactincreasedvitaminAconsumption(Simonandothers2009).Geneticandbreedingeffortshaveincreasedtotalcarotenoid-carotene,andlycopeneforcarrot,tomato,pepper,potato,muskmelons,squash(includingpumpkin),sweetpotato,maize,cassava,andlettuce(Simonandothers2009).Effortstoincreasethecarotenecontentinorangecarrotswereinitiatedinthelate1970sandhavecreatedahighcarotenecarrotwithupwardsof500ppmtotal-and-carotene(SimonandothersBiofortiÞcationwithprovitaminAisasoundstrategyfromtheperspectiveofbothunder-andover-nutritionofvitaminA.Pre-formedvitaminA,whichisfoundinanimalproductsandsupple-ments,isefÞcientlyabsorbedandepidemiologicalstudiessuggestthatchronichighintakesofpreformedAareassociatedwithhipfractures(Melhusandothers1998;Promislowandothers2002).CarotenoidsprovideasaferformofvitaminAduetoregulatedbioconversioninthebody(Tanumihardjo2008).Additionally,thisstrategyisapplicabletothedevelopingworldbecauseoftherelianceonplant-basedfoodsforupto80%oftheirvitaminACarrotsareanaturaltargetforcarotenoidbiofortiÞcationduetotheirhighlevelsofconsumptionintheUnitedStates,affordabilitytoconsumers(USDAERS2007),widespreadcultivationworld-wide,andpotentialforincreasedcarotenebiosynthesis.Becausecarotenesincarrots(andothernongreentissues)arenotneces-saryforgrowth,theyhavegreatergeneticvariationforprovita-minAcontentthanforstoragecarbohydrateorproteincontents(Simon1990).Early20thcenturyorangecarrotsrangedfrom79to90ppm(Simonandothers2009).Successfulbreedingforin-creasedcarotenecontentthroughvisualselectionfordarkeror-angerootshasbroughtcurrenttypicalorangecarrotstogreaterthan130ppmcarotene.Thishasbeenaccomplishedwithoutadverselyaffectingculinaryquality(Simon1988;Simonandoth-ers1989).AdarkorangeÒhigh-carotene-massÓcarrothasbeendevelopedthatcontainsupto500ppmcaroteneandisthehigh-estnaturalwholefoodsourceof-carotene(Simonandothers2009).TheseÒhigh-carotene-massÓcarrotshavebeenshowntoincreaseserum-caroteneovertypicalorangecarrotsinhumans(Tanumihardjoandothers2009a),andmodestly,butsigniÞcantly,increaselivervitaminAstoresinMongoliangerbils(Porter-Dostiandothers2006).CarotenoidBioavailability Ofthehundredsofcarotenoidsinnature,onlyabout40arepresentinthetypicalhumandiet,andthosecommonlyquantiÞedinhumanbloodare-carotene,lycopene,lutein,-cryptoxanthin(Figure1)(Khachikandothers1997;RaoandRao2007).Serumconcentrationsofcarotenoidsvaryaccord-ingtotheseasonoftheyear(Olmedillaandothers1994),whichlikelyreßectsseasonalßuctuationsinthefoodsupply(Zieglerandothers1987).Carrotsareauniquesourcefor-caroteneandthiscarotenoidinserumuniquelyindicateshighcarrotconsumption(Campbellandothers1994;Yangandothers2008;Tanumihardjoandothers2009a).Becausecarrotsareavailableyear-round,areawidelyeatenvegetableinmanypartsoftheworld,andmostvarietiesareaconcentratedsourceof-and-carotene,ef-fortshavebeenmadetomeasuretheirabilitytosupplydietarycarotenesandvitaminA.Approximately150high-carotenecar-rots(anamountgrownin1m)mayprovidetheannualtotalvitaminArequirementsforanadult(Simon1990).EffectsofcarrotmatrixandprocessingCarotenoidsmustÞrstbereleasedfromthefoodmatrixbeforetheycanbesolubilizedandincorporatedintolipidmicellarstructuresforuptakeintointestinalmucosalcells.Bioavailabilityof-caroteneisthefractionofcarotenoidthatisabsorbedandavailableforutilizationinphysiologicfunctionsorVol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafetyforstorage.Studiessuggestthatthecarrotmatrixhasanegativeeffectoncarotenoidabsorption.Serumresponsetobothacute(Brownandothers1989;Buluxandothers1998)andchroniccarrotfeeding(Micozziandothers1992;Torronenandothers1996)showedreducedbioavailabilitycomparedtopuriÞedCarrotscontainseveralfoodmatrixcomponentsthatmayaffectcarotenoidbioavailability.EvidencefromaferretmodelsuggeststhatthecrystallineformofcarotenoidsincarrotrootchromoplastsanddietaryÞberbothnegativelyaffectbioavail-ability(Zhouandothers1996).Detailedinvitroresearchmod-elingthegastricenvironmentsuggeststhatmembrane-boundcarotenoidsinspinachchloroplastsaremoreresistanttosol-ubilizationintooilthanmembrane-boundandfreecrystallinecarotenoidsinthecarotenebodiesofcarrots(Richandoth-ers2003a).However,blanching,whichdisruptsmostoftheor-ganelles,increasessolubilizationofboth-caroteneandluteinfromhomogenizedspinachbymorethan60%,butwasshowntohavelittleeffectonthesolubilizationof-caroteneincarrotjuice(Richandothers2003a).Inrawcarrotjuice,cellwallsareremovedandthecellu-larstructurescontainingchromoplastsaredisrupted.Blanchingofcarrotjuicefurtherdisruptsthemembrane-boundcarotenebodies,butleavescarotenecrystalsintact(Richandothers2003a).ItwashypothesizedthatthecaroteneÐcaroteneinter-actionsinthecrystalsaremorestableagainstsolubilizationthanthelipoproteinÐcarotenoidinteractionsoftheblanchedspinach.Thisisinagreementwiththeobservationthatheatingofcar-rotjuicedidnotsigniÞcantlyincreasecarotenebioavailabilityinferrets(Zhouandothers1996).Inamodeloftheduodenalenvironment,solubilizationofcarotenoidsintomixedmicelleswaslowestforcar-rotjuicecarotenes(luteincarrotjuiceblanchedfrozenspinachblanchedfrozenspinachcarrotjuice),suggestingthat,intheduodenum,themoreapolarcarotenesremaininginthefoodmatrixarelesslikelytobeavailableforabsorptionthanlutein(Richandothers2003b).Thisinvitroresearchissupportedbyobservationsinhumansthatxanthophyllshaveahigherbioavail-abilitythancarotenesfrommixedvegetablediets(VanhetHofandothers1999).Assuggestedabove,foodprocessing,includingheatingandmechanicalandenzymatictreatments,resultsinsoften-inganddisruptionoftheplantcellwallandorganelles(Richandothers2003a).Pureeingvegetablesreducesparticlesizeandincreasessurfacearea,whichmayincreaseavailabilityofcarotenoidstotheintestinallumen(Erdmanandothers1993;Parker1996).Thisreleasefromthefoodmatrixbeforeabsorp-tioniscalledbioaccessibility(Tanumihardjo2002).Plasmare-sponseof-carotenefromasinglemealofcookedcarrots(65.1%7.4%)washigherthanfromraw,groundcarrots(41.4%7.4%)inileostomyvolunteersascalculatedfromacarotenoidmass-balancestudy(Livnyandothers2003).Mildheattreat-mentofcarrotsresultedinasmall,nonsigniÞcantincreasein-caroteneabsorptioninpreruminantcalves(Poorandothers1993).Inhumans,achronic4-wkfeedingofthermallypro-cessedandpureedcarrotandspinachincreasedplasmacarotene3timesoverthatoftherawvegetables(Rockandothers1998).Inhealthyadults,-and-carotenehadgreaterbioavailabilityfromasingletestmealinthefollowingorder:commercialcarrotpuree(thermallytreated)rawÐgratedcarrots(Edwardsandothers2002).AnotherrelevantcarrotmatrixcomponentiscarotenoidÐproteincomplexes.Denaturingofcarotenoid-proteincomplexesbyheatingmightimprovecarotenoidavailability(Dietzandothers1988).Two-and-carotenebindingproteinshavebeenidentiÞedincarrotrootchromoplasts(Milicuaandothers1991;Bryantandothers1992;Zhouandothers1994);however,theirassociationwithonlyasmallfractionofthetotalcarrotcarotenoidsmakesitunlikelythattheyhaveamajorinßuenceoncarotenoidbioavailability(Zhouandothers1994).CarrotssupplyvitaminAVitaminAisderivedfromthedietaspreformedretinolorprovitaminAcarotenoids.The3majordietarycarotenoidsthathaveprovitaminAactivityare-cryptoxanthin.Twoofthesecarotenoids,namely-and-carotene,arehighlyconcentratedinorangecarrotandsomevarietiesofpurpleandred.Carrotsupplies37%oftheU.S.availablesupplyof-carotene(Table1).Inadditiontothema-trixandprocessingfactorsindicatedpreviously,bioavailabilityofcarotenoidsisinßuencedbyanumberofotherfactorssuchasthecarotenoidspecies,theamountofcarotenoidinthediet,thepresenceofabsorptionenhancersorinhibitors,andthevita-minAstatusofthehost,amongothers(CastenmillerandWestCarotenoidsareabsorbedintointestinalepithelialcellsviasimplediffusionfrommixedmicellesandpossiblybyreceptor-mediatedtransport(YonekuraandNagao2007).Inhumans,provitaminAcarotenoidsarepartlyconvertedtovitaminAintheintestinalmucosaby-carotene15,15throughproductionofretinal,reductiontoretinol,andester-iÞcationtofattyacids.Theresultingretinylestersandothercarotenoidsarethenpackagedintochylomicraforsecretionintothelymphanddeliverytothebloodstream.Inadditiontonewlyabsorbedcarotenoidsfromameal,endogenouscarotenoidsarepresentinplasma,makingthemeasurementofabsolutebioavail-abilityofcarotenoidsandvitaminA-valueofingestedvegetablesdifÞcult(Parkerandothers1999).Nevertheless,plasmaresponse-caroteneisroutinelyusedtomeasurerelativebioavailabilityfromdifferentfoodsorsupplements.Manystudieshaveconclusivelyshownthat-caroteneisbioavailablefromcarrots(RaoandRao1970;Brownandoth-ers1989;Micozziandothers1992;Buluxandothers1998;Mullerandothers1999;Tanumihardjoandothers2009a)andthatprocessingviacookingorpureeingimprovesbioavailability(Rockandothers1998;Edwardsandothers2002;Livnyandoth-ers2003).Therelativebioavailabilityof-carotenefromcarrotshasbeenmeasuredbycomparingwholeplasma-carotenere-sponsefromcarrotstothatfrompuriÞed-carotenesupplemen-tation.Brownandothers(1989)reported21%bioavailabilityof-carotenefromintakeof29mgfromcookedcarrotsinhealthymen.Torronenandothers(1996)founda45%bioavailabilityof-caroteneinhealthywomenfromdailyintakesof12mgasei-therrawcarrotsorcarrotjuicefor6wk.Huangandothers(2000)provided12mgof-caroteneasstir-friedshreddedcarrotsandreporteda33%bioavailabilityinmen.Micozziandothers(1992)provided12mg-caroteneascookedcarrotstohealthymenfor6wkandreported18%bioavailability.Thevariationbetweenstudiesislikelyrelatedtostudydesign,subjectpopulation,and-carotenesupplementformulation(FaulksandSouthon2005).Thesestudiesdemonstratethatbioavailabilityof-carotenefromcarrotsislessthanfrom-carotenesupplements,butdonotpro-videabsoluteamountsnorindicationsofhowwellcarrotscansupplyvitaminA.Usinganextrinsicstableisotopereferencemethodandre-sponseinthetriacylglycerol-richlipoproteinfractionofplasma,Edwardsandothers(2002)wereabletomeasurenotonlytherelativeavailabilityof18.6mg-carotenefromdifferentcarrotpreparations,butalsotoestimatethevitaminAyield.Themassofabsorbed-and-carotenewasalmost2-foldgreaterfromacommercialcarrotpureethanfrommashedÐboiledcarrots.However,themassofassimilatedvitaminAwasonlymarginallyCOMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... greaterforthepureethanfortheboiledÐmashedcarrots.Theap--carotenetovitaminAconversionefÞciencies(44%and59%,inthepureeandboiled-mashedtreatments,respectively)weredifferentandsuggestedalevelofregulationofvitaminAproductionbytheintestine.The-carotenetovitaminAconver-sionefÞcienciesforbothcarrotpreparationswere23%to28%lowerthantheefÞcienciesproposedbytheInstituteofMedicineMicronutrientPanelof12-caroteneto1gretinoland-caroteneto1gretinol(InstituteofMedicine,FoodandNutritionBoard2001).Thehypothesesforthelowerconver-sionefÞciencyinclude:under-estimationinthemodelused,therelativelyhigh-caroteneamount,andtherelativelyrepletevi-taminAstatusofthesubjects.EvidencesuggeststhatabsorptionefÞciencydecreaseswithincreasingamountfed(Ol-son1987)andconversionefÞciencyincreasesinmorevitaminA-deÞcientindividuals(Tanumihardjo2008).AstudyutilizingintrinsicallylabeledvegetablesfoundcarotenetovitaminAconversionfactorsof15-carotenetogretinoland21to1gforanintakeof10.3mgfromsteamedÐpureedcarrotsandsteamedÐpureedspinach,re-spectively(Tangandothers2005).TheseresultsforcarrotareclosetotheInst.ofMedicineÕsvalue,butthespinachconversionfactorishigher,suggestingthat-caroteneismorebioavailablefromcarrotchromoplaststhanfromspinachchloroplasts.Inastudyoflactatingwomenfedastandardizedamountof6mg-caroteneintheformofasupplement,rawgratedcar-rot,orpureedpapayafor60d,theserumretinolconcentra-tionsimprovedinallofthetreatmentgroupsascomparedwiththeplacebo(Ncubeandothers2001).Aswithmoststudiesofcarotenoidbioavailability,therewasmuchindividualvariation.However,inallindividualswhowereseverelymalnourished,therelativedoseresponsetest,aqualitativemeasureofvitaminAliverreserves,improvedinallinterventiongroups.Greaterimprovementoccurredinthepapayagroupcomparedwiththecarrotgroupsuggestingthatcrystalline-carotenefromcarrotchromoplastsmaybelessbioavailablethan-carotenedissolvedinlipiddropletsofyellowÐorangefruitchromoplasts.HighercarotenebioavailabilityfromorangefruitwasalsosupportedbyastudyinIndonesianschoolchildren,althoughgreenleafyveg-etableswereincludedinthecarrot-fedgroup(DePeeandothersVitaminAisrequiredforvisualdarkadaptationindimlight.InsufÞcientvitaminAcanresultinnight-blindness.Astudyinnight-blind,pregnantNepaliwomenusedrecoveryfromim-paireddarkadaptationasafunctionalendpointfordailysup-plementationwithamaranthleaves,carrots,goatliver,vitaminA-fortiÞedrice,orretinylpalmitate(Haskellandothers2005).Cookedandpureedcarrotsperformedaswellasalloftheothertreatmentgroupsinimprovingdarkadaptationandself-reportedrecoveryfromnight-blindness.Thebodyofevidencefromthesecarrotstudiessuggeststhatcarrotsareabioavailablesourceof-carotene;thatprocess-ingthevegetablebygrating,heating,and/orpureeingimprovesthebioavailability;andthatthecarotenebioavailabilitymaybeintermediatetofruitsandleafygreenvegetables.Additionally,carrotsappeartobeavaluablesourceofvitaminAfordeÞcientpopulations,especiallyifprocessedandprovidedinadequateBioavailabilitystudieswithcarrotsofvariouscolorsTheÞrstcarrotswerepurpleandyellow,buthavemainlybeensupplantedintheWestbytheorangecultivarsoftoday.Color-fulspecialtycarrotshavebeenÒre-discoveredÓbymodernplantbreedersinterestedinimprovinganddiversifyingthenutritionofthefoodsupply.Inadditiontoeffortstoimprovetheßavor,texture,andhorticulturalqualityofcoloredcarrots,researchhasfocusedonthebioavailabilityandputativehealthbeneÞtsofthedifferentpigmentsthatcolorthesecarrots.Therelativebioavailabilityoflycopeneinredcarrotswasde-terminedin2similarcrossoverstudiesinhumansfedcarrotortomatopastemufÞnsat5mglycopene/dayfor11d(Horvitzandothers2004).TheÞrststudyincludedmufÞnsmadefromredcarrots,whitecarrots,andtomatopaste.ThesecondstudydeterminedifcarrotÞberaffectedlycopeneavailabilitybyfeed-ingtomatopastemufÞnswithorwithoutwhitecarrots.Lycopene-carotenewereavailablefromredcarrot,butlycopeneab-sorptionwasnegativelyaffectedbycarrotÞber.Combinedresultsfrombothstudiessuggestedthatlycopeneinredcarrotisabout44%asavailableasthatfromtomatopaste.Astudywithredcar-rotlycopeneinvitaminA-depletedMongoliangerbilsconÞrmedthatlycopeneisbioavailableandsuggestedapossibleinterac-tionbetween-caroteneandlycopenethatdecreaseslycopeneavailability(Millsandothers2007).Additionally,becausetheredcarrotalsocontained-carotene,andthuscouldsupplyvitaminA,thevitaminAvalueoftheredcarrotwascalculatedtobe-caroteneto1gretinolinthismodel.ThisismorefavorablethanthecurrentInstituteofMedicinevalueof12toginhumans(InstituteofMedicine,FoodandNutritionBoardInvitroresearchfoundredcarrotspreventedoxidationofcholesterolduringheatingcomparablytopurple,orange,anddarkorangecarrots(Sunandothers2009).InconsiderationoftheseÞndings,redcarrotscouldserveasanoptionfordeliveringimportantvitaminAnutritionandtheantioxidantlycopene.Luteinbioavailabilityfromyellowcarrotwasexaminedinhu-mansbyfeeding1.7mglutein/dayfromyellowcarrotsoraluteinsupplementdissolvedinoil,andwhitecarrotsasanegativecon-trol(Molldremandothers2004).Thesubjectswerefedcarrotsmoothies,mufÞns,andsoupforbreakfastandlunchfor7d.TheluteinfromyellowcarrotssigniÞcantlyincreasedserumconcen-trationsandwasfoundtobe65%asbioavailableastheluteinsupplement.Theyellowcarrottreatmentalsomaintainedserum-caroteneconcentrations,whereastheluteintreatmentdidnot.Bioavailabilityofcrystallinelutein,whichistheformfoundinmostsupplements,ishighlyvariablebetweenandwithinsub-jects(Tanumihardjoandothers2005).Whileyellowcarrotsarenotaconcentratedsourceofluteincomparedtoothervegeta-bles,especiallygreenleaves,theymayserveasanalternativebioavailablesourceoflutein.-carotene,dark-orangecarrotswerecomparedwithtyp-icalorangecarrotstodetermineifincreasedcarotenecontentwasbioavailableduringasustained-feedingcross-overstudy(Tanumi-hardjoandothers2009a).Orange,dark-orange,andwhitecarrotmufÞnsproviding2.6,7,and0mg-carotene/d,respectively,werefedfor11dtohealthyyoungadults.Comparedwithbase-line,thedark-orangeandorangecarrotmufÞnsincreasedserum-caroteneconcentration127%and85%,respectively.Thisin-creasewasdifferentbetweentreatmentsintheÞrst20-dperiodalone,thatis,thedark-orangecarrottreatmentsigniÞcantlyraised-and-caroteneserumconcentrationsabovethetypicalorangecarrottreatment.Dark-orangecarrotswerealsostudiedinMongoliangerbils(Porter-Dostiandothers2006),whichallowsthemeasurementoflivervitaminAstores,thegold-standardforevaluatingvitaminAstatus(Tanumihardjo2004).Whendietswereequalizedtocar-rotcontent,thedark-orangecarrottreatmentresultedindoubletheliver-carotenecontentcomparedwithtypicalorangecar-rots,butonly10%greaterlivervitaminAstores.ThevitaminAconversionfactorswereestimatedtobe9to11to1gretinolforthetypicalorangecarrotsandabout23gtogforthedark-orangecarrots(Simonandothers2008).ThisstudyutilizedgerbilswithadequatevitaminAstatus.BecauseimprovedbioconversionwithlowerlivervitaminAreserveshasVol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafetybeendemonstratedinanimalmodels(Tanumihardjo2008),fur-therresearchwithvitaminA-depletedanimalsorhumansubjectsiswarrantedandmayshowimprovedconversionoftheprovita-minAcarotenoidsinthehigh-carotenecarrots.Theclassicdef-initionofbioavailabilityincludesall-caroteneabsorbedfromthefood,yethumanstudiesarenotabletoassessthestoredintissues.Therefore,studiesthatmerelylookatserumresponseinhumansneedtobecautiouslyinterpreted.Ifdark-orangecarrotswerewidelyadopted,theycouldreadilyincreaseconsumptionof-caroteneandpotentiallyimpactvitaminAsta-tusofindividualsatriskfordeÞciency.PurplecarrotsarecoloredbyafamilyofblueÐredpigmentscalledanthocyanins.ThevitaminAvalueofpurpleÐorangecar-rots,andpurpleÐorange-redcarrotswerecomparedtoorangecarrotsinMongoliangerbils(Porter-Dostiandothers2006;Millsandothers2008).Liverstoresof-caroteneandvitaminAinthegerbilsdidnotdiffersuggestingthatthehigherphenolicandanthocyanincontentsofpurplecarrots(Alasalvarandothers2001;Nicolleandothers2004;Grassmannandothers2007;Sunandothers2009)donotinterferewiththebioavailabilityofcarotenefrompurplecarrots.Morerecently,astudyof5healthy,youngfemalesubjectscomparedtherelativebioavailabilityof10.3mg-carotenefromanacutefeedingofpurpleÐorangeandorangecarrotbreakfastsmoothies,withwhitecarrotsmoothiesasanegativecontrol(Arscottandothers2009).Analysisofarea-under-the-curveforplasma-carotenefor0to144hshowedthattheorangeandpurple-orangecarrotsincreasedplasmacarotene5.4-and4.5-old,respectively,comparedtothewhitecarrots,indicatingasmall,butsigniÞcant32%greaterresponsefromtheorangethanfromthepurpleÐorangecarrots.However,whenonlytheÞrst24hwerecompared,therewasnodifferenceinplasma-caroteneresponsebetweenpurpleandorangecar-rottreatments.Anthocyaninsarerapidlyabsorbed,appearinginplasmaby30minandachievingmaximumconcentrationby4h(Kurilichandothers2005;Charronandothers2009),soiftheyweretoexertaneffecton-caroteneabsorptioninthisstudy,itwouldbeexpectedinthistimeperiod.ThisstudyconÞrmedthat-carotenewasasbioavailablefrompurpleÐorangeasorangeHealthEffects Dietsrichinfruitsandvegetableshavebeenassociatedwithreducedriskofdegenerativediseasesincludingsomecancers(RiboliandNorat2003;Leeandothers2009;Zhangandoth-ers2009)andcardiovasculardisease(CVD)(Menteandothers2009).Carotenoidsareabundantinplant-basedfoodsandhavebeenimplicatedasthebeneÞcialsubstancesinthesedietsinthepreventionofdisease.TheproposedmechanismsincludeprovitaminAactivity,antioxidantfreeradicalscavengeractivitythatoffersprotectionagainstLDLoxidation(KrinskyandJohnson2005),increasedcell-to-cellcommunicationviagapjunctions(BertramandVine2005),andimmunomodulatoryeffects(ChewandPark2004).CardiovasculardiseaseTheepidemiologicliteraturerelatingcarotenoidstoCVDisconßictingandhasbeenreviewedbyVoutilainenandothers(2006).Clinicalinterventiontrialsthathavefocusedonsupple-mentationwithlargedosesof-carotenehavenotshownreduc-tionsinCVDand,infact,insomecaseshaveincreasedCVDriskincertainpopulations(Rapolaandothers1997).Explana-tionsproposedforthediscrepancybetweenobservationalandinterventionstudiesinclude:thepossibilitythat-caroteneisnottheactivecompound,butonlyanindicatorofintakeofsomeothernutrient,phytochemical,orbeneÞcialdietaryorlifestylefactor;therelativelyhighdosagesusedintheinterventionstud-iesprovidedamuchhigherÒexposureÓthannaturalfoodsources;andsubjectsintheinterventiontrialswerealreadyinhigherriskgroupsthatmighthavebeeninprogressedstatesofdisease.Population-basedstudiesalsoshowanassociationbetweendi-etshighinÞberanddecreasedriskforCVD(VanHornandothers2008).High-ÞberdietsareassociatedwithlowerLDLcholesterollevels,body-massindex,blood-pressure,andtriglyceridelevels.SolubleÞbersappeartobethemosteffectiveinloweringplasmaLDLcholesterollevels.Fibersmakeup3%to4%ofcarrotweight,ofwhichover50%issolubleÞber(Svanbergandothers1997).AscarrotsarearichsourceofbothcarotenoidsandÞber,andoneofthemajorvegetablesconsumedinthewesternworld,anumberofstudieshavelookedathowcarrotsmayexerteffectsinrelationtothediseaseprocessofCVD.Theresultsofalarge-scaleprospectivestudyreportedthatboth-and-caroteneintake,andcarrotconsumption,butnottoco-pherols,vitaminC,orothercarotenoidswereinverselyrelatedtoCVDmortalityinelderlymen(Buijsseandothers2008).TheywereunabletoÞndacorrelationbetweenCVDriskandmultipleantioxidantscoresrulingoutthatadietrichinmultipleantioxi-dantsisprotective.Becausecarrotswerethemajorsourceofboth-and-carotene,thebeneÞcialassociationmayberelatedtocarrotsthemselves,aresultofsomeothersubstancesincarrots,orahealthydietorlifestylethatishighincarrots.Interventiontrialswithcarrotsarefewandlimitedtoshort-termmeasurableendpoints.Theeffectsofcarrotsonlipidmetabolismareconßicting.Humansubjectsthatconsumed200gcarrotdailyfor3wkhad11%reductioninserumcholesterol,50%increaseinfecalbileacidsandfecalfats,andamodest,butsigniÞcant25%increaseinstoolweight(Robertsonandothers1979).Theeffectsoncholesterolandfecalbileacidsandfatpersistedfor3wkaftertreatmentended.However,noeffectsonserumcholes-terolwereobtainedwithhigherlevelsofcarrotsorcarrotÞberinhealthymen(Jenkinsandothers1979)orwomen(Wiskerandothers1994).Astudyinyoungfemalesfedbetween405and688gcarrot(15gÞber)for3wkfoundthatÞbersincarrotswerehighlyfermentable(soluble)andalsohadgoodstoolbulkingabil-itycomparabletotheinsolubleÞbersofcerealgrains(Wiskerandothers1994).However,differentprocessingofthecarrottreat-ments,thatis,raw,blanched,orcanned,alteredthedistributionofÞbertypes,butdidnothaveaneffectonthesephysiologi-calparameters,oronserumhigh-densitylipoproteincholesterollevelsorfecalbileacidsecretion.Differencesbetweenstudieswithregardtophysiologicalresponsetocarrotscouldbeduetodifferencesinproportionsofdigestiblecarbohydratesbetweencultivars(Svanbergandothers1997).OxidativemodiÞcationofLDLcholesterolhasbeenconsideredtoplayaroleinatherogenesisandcoronaryarterydisease(Diazandothers1997).DietaryantioxidantsareproposedtoprotectagainstLDLoxidation.Ina2-wkinterventioninhealthymenfed330mLtomatojuice,carrotjuice,or10gspinachpowder,all3treatmentsenhancedlipoproteincarotenoidconcentrations,butonlytomatojuicereducedplasmalipidperoxidationasmeasuredbyplasmamalonaldehydeandexvivoLDLoxidation(Bubandothers2000).Inasimilarstudy,neithertomatonorcarrotjuicehadaneffectonlipidperoxidation(Brivibaandothers2004).Researchinanimalshasexaminedtheeffectsofwholecar-rotandÞbersextractedfromcarrotpomaceonlipidmetabolismandantioxidantstatus.Cholesterol-fedratson15%carrotdiethadlowerlivercholesterolandtriglyceridesandalsoreducedserumcholesterolthroughapparentreductioninabsorptionandincreaseinthepercentageofcholesterolexcretedasbileacids(Nicolleandothers2003).Carrotalsoimprovedantioxidantsta-tusmarkersthroughreductionofurinaryexcretionofthiobar-bituricacidreactivesubstances(TBARS),reducedTBARSintheCOMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY—Vol.9,2010 Carrotsofmanycolors... heart,increaseinplasmavitaminE,andincreaseinplasmaferric-reducingabilityofplasma(FRAP).Inmice,carrotalsoloweredplasmaandlivercholesterolandtriglyceridesandinducedanin-creaseofneutralsterolfecalexcretion(Nicolleandothers2004).FRAPvalueswerealsoincreasedandtheplasmavitaminEtotriglycerideratiowasimprovedbycarrotfeeding.ThesolubleÞberfractionofcarrots,speciÞcallythepectinsandfermentablehemicelluloses,mightberesponsibleforthereductionincholes-terolcomparedtocellulose.Carrotpomaceisthebyproductofthecarrotjuicingindustry,aswellastheÒbabyÓcarrotorcut-and-peelindustry.Researchersareinterestedindescribingthecharacteristicsandpotentialhealth-promotingeffectsofthisbyproductasafunctionalÞberforaddingtofoods.TheinsolubleÞberfraction(IFF)ofcarrotpomacehadhigherwater-andoil-holdingcapacities,cation-exchangeca-pacity,glucose-adsorbingability,andamylaseinhibitionactivitythancellulose(Chauandothers2004).CarrotpomaceIFFalsoreducedserumcholesterolandtriglycerides,increasedserumHDL:LDL,andincreasedfecallipids,cholesterol,andbileacidsinhamsters(Hsuandothers2006).ResearchbythissamegroupsuggestedthatmicronizationofthecarrotpomaceIFFmayim-proveitscholesterol-loweringeffect(Chouandothers2008)andalsomeasuresofintestinalhealth(Chauandothers2007).DataonthecontributionofcarrotstoprotecthumansfromthepathogenesisofCVDareinconclusive.FeedinganimalscarrotsdemonstratesbeneÞcialeffectsonlipidmetabolismandantiox-idantaction,butinhumans,carrotshavenotyetclearlyshowndesiredeffectsontheseparameters.Theprotectiveeffectsofan-tioxidantsmayfunctionthroughmechanismsunrelatedtooxida-tionofLDL(Keaneyandothers1993;Diazandothers1997);thus,itmaybethatcarrotsexerttheireffectsonalternativebiomarkersofthepathogenesisofCVD.AssuggestedearlierforCVD,thecarotenoidassociationwithprotectionfromcancersmayactuallybeindicativeofsomeotherfactorssuchasahealthydietorlifestyle,oranotherbeneÞcialphytochemicalornutrient.Asystematicreviewoftheassocia-tionbetweenhightotalcarotenoidintakeorserumconcentrationandlungcancerriskindicatedasmall,butsigniÞcant,reductioninrisk,butsigniÞcantreductionswerenotfoundforindivid-ualcarotenoids(Gallichioandothers2008).Theevidencethatcarrotshaveanimpactontheprotectionfromcancermaybeprimarilythroughitsassociationwithahigh-vegetabledietarypattern.Nonetheless,apooledanalysisof13studiesfoundcar-rotintaketohaveasigniÞcantinverseassociationwithrenalcellcancer(Leeandothers2009).CarrotintakewasalsofoundtobeinverselyrelatedtobreastcancerincidenceamongChi-nesewomen(Zhangandothers2009).TheExpertReportoftheWorldCancerResearchFundandtheAmericanInstituteforCan-cerResearch(2007)judgedthatprobableevidencefortheroleofcarotenoid-containingfoodsincancerpreventionislimitedtothemouthandpharynx,larynx,andlung.-Carotene-andlycopene-containingfoodshaveaprobableassociationforpro-tectionfromesophagealandprostatecancers,respectively.Dataoncarrotscontributedtotheevidence-basefortheassociationofnonstarchyvegetablesandoralandlungcancersandlimited,butconsistent,evidencefromcase-controlstudiessupportedaroleofcarrotsinprotectionfromcancerofthecervix.Fewmechanisticstudiesofcarrotsandthecancerprocesshavebeenperformed.LymphocyteDNAstrandbreakagewasreducedinhealthymenconsuming330mLtomatojuice(40mglycopene)orcarrotjuice(22.3mg-caroteneand15.7mg-carotene)for2wk(Pool-Zobelandothers1997).Oxidativebasedamagewasre-ducedonlywiththecarrotjuice.Healthymenconsumingtomatojuiceorcarrotjuicefor2wkexperiencedatime-delayedmodu-lationofimmunefunctions(Watzlandothers2003).Researchbythesamegroupdemonstratedthatcarrotortomatojuiceconsumptiondidnotdecreaselipidperoxidationinplasmaorfeces(Brivibaandothers2004)orincreasecytotoxicorantipro-liferativeactivityoffecalwateroncolonadenocarcinomacells(Schnabeleandothers2008).Bothtomatoandcarrotjuiceonlyledtominorchangesinluminalbiomarkersrelevanttocoloncar-cinogenesis.Inhamsters,theinsolubleÞberfractionisolatedfromcarrotpomacereducedfecalammoniaoutputanddetrimentalcolonicbacterialenzymes,mucinase,andurease,potentiallyreducingtheexposureoftheintestinallumentoharmfulcompounds(Chauandothers2005).Thebitteroff-ßavorofcarrotsisattributedtoagroupofcom-poundscalledpolyacetylenes(CzepaandHofmann2004),whichhavebeenidentiÞedashavingantiinßammatory,antifungal,an-tibacterial,andplatelet-reducingactivity(ChristensenandBrandt2006).Polyacetyleneshaveshowninvitrobiphasiceffects,be-ingstimulatoryatlowconcentrationsandinhibitoryortoxicathighconcentrations,suggestingconcentration-dependenteffects(Hansenandothers2003).Theprimarypolyacetylenesidenti-Þedincarrots,falcarinolandfalcarindiol,haveshowninvitroantiinßammatoryactivity(Metzgerandothers2008)andanin-hibitoryeffectonthedevelopmentofpreneoplasticlesionsintheratcolon(Kob¾k-Larsenandothers2005).Falcarinolappearstobebioavailableinhumans(Brandtandothers2004),appearinginseruminconcentrationssimilartotherangeshowntohavevitrophysiologiceffect(Hansenandothers2003).Informationonthechemoprotectiveactivityofcarrotcol-orsotherthanorangeissparse.InMongoliangerbilsfedpurpleÐorange,orange,orangeÐred,orpurpleÐorangeÐredcar-rotsfor4wk,totalliverantioxidantcapacitywashigherinallthecoloredcarrotgroupsthanboththewhitecarrot-fedandvita-minA-supplementedgroups(Millsandothers2008).Blackcar-rotanthocyanin-richextractdisplayed80%invitroinhibitionofcolorectaladenocarcinoma(HT-29)andpromyelocyticleukemia(HL-60)cells(Netzelandothers2007).Invitroresearchdemon-stratedallanthocyanin-richextractsfrom7differentfruitsandvegetablessuppressedHT29cellgrowth,buttovariousdegrees,intheorder:purplecornchokeberryandbilberrycarrotandgraperadishandelderberry(Jingandothers2008).Anthocyaninchemicalstructureaffectedchemoprotection,sug-gestingfoodswithdifferentanthocyaninproÞleshavedifferinganti-canceractivities.Cautionininterpretinginvitrostudiesofanthocyaninsiswarrantedduetothehighconcentrationsem-ployedinvitroandthelowinvivobioavailabilityofthesecom-ThechemoprotectivemechanismsassociatedwithcarrotsareasinconclusiveasthoseforCVD.EvidencepointstobeneÞcialeffectsofcarrotsaspartofahealthylifestyle,whichincludesadietrichinavarietyoffruits,vegetables,andotherwholefoods.Excessivecarrotconsumption(1kg/djuicedorraw),whichhasbeenreportedinrarecases,maybeassociatedwithleucopenia(Shoenfeldandothers1982)andamenorrhea(Kemmannandothers1983;Mikkelsenandothers2009).SatietyandglucosemetabolismDietaryÞberincreasessatietyanddecreasesenergyintake(Howarthandothers2001).Fruitsandvegetablesareafoodgroupthatisusuallylowinfatcontentandenergydensity,andhighinwateranddietaryÞber.Thus,theymaycontributesigniÞcantlytohungercontrolandweightmanagement.Relativelyfewstud-ieshavespeciÞcallytestedtheeffectsoffruitsandvegetablesonsatiety,foodintake,andbodyweight(Rollsandothers2004;Ello-Martinandothers2007;Tanumihardjoandothers2009b).SomeeffortshavebeenmadetospeciÞcallyassesstheeffectsofcarrotsandtheircomponentsontheseparameters.Vol.9,2010COMPREHENSIVEREVIEWSINFOODSCIENCEANDFOODSAFETY ComprehensiveReviewsinFoodScienceandFoodSafety Figure4---Inanefforttogainpublicawareness,carrotseedsfrombiofortiÞedvarietiesaredistributedtolocalcommunityandyouthgardensthroughoutreachactivities.ThisresultsindirectconsumptionoftheseÒfunctionalÓcarrotsbyconsumers.Gustaffsonandothers(1994)foundthatinisocaloricmeals,thelargerthecarrotportion(100,200,and300gcarrotcontaining2.9,5.8,and8.7gÞber,respectively),thelowertheglucoseandinsulin/C-peptideresponsesandthehigherthesatietyscores.Theminimumservingsizecausinganeffectwas200g.However,effectsofprocessingandcookingwith2differentcarrotharvestshadmixedresults(Gustaffsonandothers1995).The1styear(4.4gÞberincarrots),therawcarrottreatmentenhancedsatiety,andreducedglucoseandC-peptideresponsesmorethanthecooked.The2ndyear(6.6gÞberincarrots)onlytheglucoseresponsewasaffectedbyprocessing.Thus,energydensityratherthancookingmethodmayhaveaffectedsatiety.Alternatively,theexponentiallyhigherviscosityelicitedbythehigherÞberdose(Svanbergandothers1995)mayhaveunexplainedmetabolicAnotherstudyexaminedwhetherthephysicalstructureorÞbercontentofcarrotsexertsaneffectonsatietybyfeeding200gwholecookedcarrots(structureandÞber),cookedpureedcar-rots(nostructure,someÞber),andcarrotnutrients(nostruc-ture,noÞber)tohealthyadultwomen(Mooreheadandothers2006).Theymeasuredsatietyaswellassubsequentenergyin-takestherestofthedayandfoundthatthewholecarrotsandblendedcarrotshadhighersatietyscoresthanthecarrotnutri-ents.Subsequentenergyintakesincreasedinthefollowingorder:wholecarrotsblendedcarrotscarrotnutrients.Thus,bothÞbercontentandstructureplayedabeneÞcialrole.Thisresearchissupportedbyarecentfruitandvegetableweight-lossinter-ventiontargetedtoobeseindividuals,whichfedcarrotsatleast3timesperweek,showingsmall,butsigniÞcant,weightlossat3moandbaselineweightmaintenanceat18mo(Tanumi-hardjoandothers2009b).Additionally,carrotsmayhaveplayedaroleinmaintainingglycemiccontrolintype2diabeticsubjectsfedcarrotcakefor24d(BrynesandFrost2007).Morestudyiswarrantedtocomparecarrotswithothervegetables,effectsofprocessing,andaddedÞberisolatesonsatietyandweight FunctionalfoodsprovidebeneÞtbeyondbasicnutrition.Bio-fortiÞedcarrotsnotonlyprovidevitaminAbutmaycontributetooptimalhealth.Moreresearchisneededtobetterunderstandthecontributionofcarrotstohealthandthemechanismsinvolved.CarrotfeedingstudiesinspeciÞcpopulations,suchasolderadultsorhypercholesterolemicindividuals,couldprovideinformationuniquetotheseoxidativelychallengedgroups.Combinedwithmeasurementofalternativemarkersofoxidativestress,suchastheproductionofF-isoprostanes,orsurrogatemarkersofcar-diovascularhealth,suchasplateletaggregationorendothelialfunction,newinsightsmaybeelucidatedconcerningtheeffectsofvariousphytochemicalsincarrots.Macronutrient,Þber,andphytochemicalanalysesofcarrotsusedinthesestudiesarecrit-icalbecausegeneticvariabilitymayprecludegeneralizationofeffectstoallcarrots.Additionally,moreinformationisneededre-gardingthemetabolicproductsandinvivomechanismsofactionofcarrotcompounds,suchasanthocyanins,phenolicacids,andpolyacetylenes.Inanefforttoenhanceconsumerawarenessoverthepastdecade,carrotseedsfrombiofortiÞedvarietieshavebeendistributedeveryspringtocommunityandyouthgardeninginitia-tivesleadingtodirectconsumptionofspecialtycarrotsofvariouscolors(Figure4).MostepidemiologicresearchpointstoadiethighinfruitandvegetablesincombatingdiseaseandcolorfulcarrotsmaycertainlycontributetothisbeneÞt.Acknowledgments ThisresearchisinpartialfulÞllmentoftherequirementsforSAAtoobtainthePhDdegreeinNutritionalSciences.Theau-thorsthankPhilippSimon,ProfessorofHorticulture,forshar-ingimportantresourcesandreadingthereviewaspartofthedissertationcommittee.ThisreviewwassponsoredinpartbyStandardProcessInc.,HatchWisconsinAgriculturalExperimentStationWIS04975,andtheUW-MadisonGraduateSchool.Re-searchconductedinDr.TanumihardjoÕslaboratorythatispartofthisreviewwassponsoredbyUSDA-IFAFSgrantnumber2000-4258,AmericanCancerSocietyIRG-580114402,Hatch-WisconsinAgriculturalExperimentStationWIS04533,andtheNatl.ResearchInitiativeoftheUSDACooperativeStateRe-search,EducationandExtensionService,grantnr2003-35200-References 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