Subcellular Biochemistry 79 Editor Claudia Stange Carotenoids in Nature Biosynthesis, Regulation and Function Subcellular Biochemistry Volume 79 Serieseditor J.RobinHarris UniversityofMainz,Mainz,Germany The book series SUBCELLULAR BIOCHEMISTRY is a renowned and well recognized forum for disseminating advances of emerging topics in Cell Biology and related subjects. All volumes are edited by established scientists and the individual chapters are written by experts on the relevant topic. The individual chaptersofeachvolumearefullycitableandindexedinMedline/Pubmedtoensure maximumvisibilityofthework. SeriesEditor J.RobinHarris,UniversityofMainz,Mainz,Germany InternationalAdvisoryEditorialBoard T.Balla,NationalInstitutesofHealth,NICHD,Bethesda,USA TapasK.Kundu,JNCASR,Bangalore,India A.Holzenburg,TexasA&MUniversity,CollegeStation,USA S.Rottem,TheHebrewUniversity,Jerusalem,Israel X.Wang,JiangnanUniversity,Wuxi,China Moreinformationaboutthisseriesathttp://www.springer.com/series/6515 Claudia Stange Carotenoids in Nature Biosynthesis, Regulation and Function 123 ClaudiaStange CentrodeBiologíaMolecularVegetal DepartamentodeBiología FacultaddeCiencias UniversidaddeChile Santiago,Chile ISSN0306-0225 SubcellularBiochemistry ISBN978-3-319-39124-3 ISBN978-3-319-39126-7 (eBook) DOI10.1007/978-3-319-39126-7 LibraryofCongressControlNumber:2016947054 ©SpringerInternationalPublishingSwitzerland2016 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof thematerialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthorsandtheeditorsaresafetoassumethattheadviceandinformationinthisbook arebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsor theeditorsgiveawarranty,expressorimplied,withrespecttothematerialcontainedhereinorforany errorsoromissionsthatmayhavebeenmade. Printedonacid-freepaper ThisSpringerimprintispublishedbySpringerNature TheregisteredcompanyisSpringerInternationalPublishingAGSwitzerland Preface Carotenoidsarecoloredpigmentswidespreaddistributedinnature.Theselipophilic molecules are synthesized in plants, algae, and some yeast and bacteria. In plants, carotenoids are synthesized in photosynthetic organs as well as in fruits, flowers, seeds, and reserve roots, providing attractive yellow, orange, and red colors. In plants and algae, carotenoids have important functional roles in photosynthesis, photomorphogenesis,andinphotoprotection.Theyalsogiverisetoapocarotenoids, such as the hormones abscisic acid and strigolactones, among other volatile terpenes.Additionally,theypossessantioxidantpropertiesactingasreactiveoxygen speciesscavengers.Inmammals,theyactasprovitaminAprecursorsandaspower- ful antioxidant molecules involved in the prevention of certain types of diseases. Carotenoid biosynthesis in plants is highly regulated, although all the processes involvedarenotcompletelyknown.Duringthepastdecades,hugeknowledgehas beenpublished,andalmostallcarotenogenicgeneshavebeenidentifiedandthose functions dissected as a result of molecular, genetic, and biochemical approaches utilizingdifferentplant,yeast,andalgaemodelsystems.Theinformationhasbeen used in genetic engineering for increasing abiotic stress tolerance, altering color and the nutritional value in plants, leading to the production of novel functional foods.Inthisbook,anextensiveandactualreviewofthemaintopicsofcarotenoid biosynthesis,regulation,andfunctioninhumanhealtharebroughttogether. The first chapters provide an introduction to carotenoid biosynthesis in yeast, bacteria, and plants and a profound exposition on the structures of carotenoid molecules. The second part covers the function and regulations of carotenoids in photosynthesisaswellasduringplant,fruit,storageroot,andalgadevelopment.We also included chapters that present an actual overview on plastids – accumulating carotenoids–ontheepigeneticmechanismsthatcontrolcarotenoidbiosynthesisand on the oncoming topic regarding apocarotenoids. To finish, some chapters argue about the metabolic engineering of carotenoids in plants and seeds that point the way of carotenoid biotechnological application. Additionally, important topics on theeffectofabsorptionmechanismsandcarotenoidsasantioxidantsandvitaminA precursorsforhumannutritionwerealsoincluded. v vi Preface Thechaptersincludedineachsectionwerepreparedandreviewedbyexpertsin thefield.Iwouldliketothankeachoftheauthorsforacceptingtocontributetothis book, for their dedicated effort providing carefully prepared manuscripts, and for theirpromptattentiontoanswertorequestedinformation. IwishtoexpressmygratitudetothestaffofSpringer,inparticulartoThijsvan Vlijmen(SeniorPublishingEditor)andtoSaraGermans-Huisman(SpringerSenior EditorialAssistant)fortheirkindassistanceandpatience. Itistheaimoftheeditorthatthisbookwillbeofbenefitandreferencesourceto anyoneresearchingtheareaoncarotenoidsynthesisandregulation. Santiago,Chile ClaudiaStange Contents PartI BiosynthesisofCarotenoids 1 CarotenoidDistributioninNature........................................ 3 JenniferAlcaíno,MarceloBaeza,andVíctorCifuentes 2 BiosynthesisofCarotenoidsinPlants:EnzymesandColor ........... 35 CarolinaRosas-SaavedraandClaudiaStange 3 StructuresandAnalysisofCarotenoidMolecules...................... 71 DeliaB.Rodriguez-Amaya PartII RegulationofCarotenoidsBiosynthesis 4 CarotenoidsandPhotosynthesis.......................................... 111 HidekiHashimoto,ChiasaUragami,andRichardJ.Cogdell 5 RegulationofCarotenoidBiosynthesisinPhotosyntheticOrgans.... 141 BriardoLlorente 6 RegulationofCarotenoidBiosynthesisDuringFruitDevelopment... 161 JoannaLado,LorenzoZacarías,andMaríaJesúsRodrigo 7 CarotenoidBiosynthesisinDaucuscarota............................... 199 KevinSimpson,ArielCerda,andClaudiaStange 8 CarotenoidsinMicroalgae................................................. 219 Vitalia Henríquez, Carolina Escobar, Janeth Galarza, andJavierGimpel 9 Apocarotenoids:ANewCarotenoid-DerivedPathway................. 239 JuanCamiloMorenoBeltranandClaudiaStange 10 PlastidsandCarotenoidAccumulation .................................. 273 LiLi,HuiYuan,YunliuZeng,andQiangXu vii viii Contents 11 EvidenceofEpigeneticMechanismsAffectingCarotenoids........... 295 Jacobo Arango, Jesús Beltrán, Jonathan Nuñez, andPaulChavarriaga PartIII CarotenoidsforHumanHealth 12 ManipulationofCarotenoidContentinPlantstoImprove HumanHealth .............................................................. 311 EnriquetaAlós,MariaJesúsRodrigo,andLorenzoZacarias 13 Modern Breeding and Biotechnological Approaches toEnhanceCarotenoidAccumulationinSeeds......................... 345 M.L.FedericoandM.A.Schmidt 14 CarotenoidsasaSourceofAntioxidantsintheDiet.................... 359 AnaAugusta OdorissiXavierandAntonioPérez-Gálvez 15 CarotenoidsinAdiposeTissueBiologyandObesity.................... 377 M.LuisaBonet,JoseA.Canas,JoanRibot,andAndreuPalou 16 AbsorptionofCarotenoidsandMechanismsInvolved inTheirHealth-RelatedProperties....................................... 415 BraulioCervantes-Paz,ClaudiaI.Victoria-Campos,and JosédeJesúsOrnelas-Paz Index............................................................................... 455 Part I Biosynthesis of Carotenoids