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BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm The Molecular Biology and Biochemistry of Fruit Ripening BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm The Molecular Biology and Biochemistry of Fruit Ripening Edited by GRAHAM B. SEYMOUR MERVIN POOLE JAMES J. GIOVANNONI GREGORY A. TUCKER A John Wiley & Sons, Inc., Publication BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm Thiseditionfirstpublished2013©2013byJohnWiley&Sons,Inc. Wiley-BlackwellisanimprintofJohnWiley&Sons,formedbythemergerofWiley’sglobalScientific,Technical andMedicalbusinesswithBlackwellPublishing. Editorialoffices: 2121StateAvenue,Ames,Iowa50014-8300,USA TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 9600GarsingtonRoad,Oxford,OX42DQ,UK Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyfor permissiontoreusethecopyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell. Authorizationtophotocopyitemsforinternalorpersonaluse,ortheinternalorpersonaluseofspecificclients,is grantedbyBlackwellPublishing,providedthatthebasefeeispaiddirectlytotheCopyrightClearanceCenter,222 RosewoodDrive,Danvers,MA01923.ForthoseorganizationsthathavebeengrantedaphotocopylicensebyCCC, aseparatesystemofpaymentshasbeenarranged.ThefeecodesforusersoftheTransactionalReportingServiceare ISBN-13:978-0-8138-2039-2/2013. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesand productnamesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheir respectiveowners.Thepublisherisnotassociatedwithanyproductorvendormentionedinthisbook.This publicationisdesignedtoprovideaccurateandauthoritativeinformationinregardtothesubjectmattercovered.Itis soldontheunderstandingthatthepublisherisnotengagedinrenderingprofessionalservices.Ifprofessionaladvice orotherexpertassistanceisrequired,theservicesofacompetentprofessionalshouldbesought. 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CoverdesignbyNicoleTeut Setin11/13ptTimesbyAptara®Inc.,NewDelhi,India Disclaimer Thepublisherandtheauthormakenorepresentationsorwarrantieswithrespecttotheaccuracyorcompletenessof thecontentsofthisworkandspecificallydisclaimallwarranties,includingwithoutlimitationwarrantiesoffitness foraparticularpurpose.Nowarrantymaybecreatedorextendedbysalesorpromotionalmaterials.Theadviceand strategiescontainedhereinmaynotbesuitableforeverysituation.Thisworkissoldwiththeunderstandingthatthe publisherisnotengagedinrenderinglegal,accounting,orotherprofessionalservices.Ifprofessionalassistanceis required,theservicesofacompetentprofessionalpersonshouldbesought.Neitherthepublishernortheauthorshall beliablefordamagesarisingherefrom.ThefactthatanorganizationorWebsiteisreferredtointhisworkasa citationand/orapotentialsourceoffurtherinformationdoesnotmeanthattheauthororthepublisherendorsesthe informationtheorganizationorWebsitemayprovideorrecommendationsitmaymake.Further,readersshouldbe awarethatInternetWebsiteslistedinthisworkmayhavechangedordisappearedbetweenwhenthisworkwas writtenandwhenitisread. 1 2013 BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm Contents ListofContributors ix Preface xi Chapter1 BiochemistryofFruitRipening 1 SoniaOsorioandAlisdairR.Fernie Introduction 1 CentralCarbonMetabolism 4 EthyleneinRipening 7 Polyamines 9 Volatiles 10 CellWallMetabolism 11 ConcludingRemarks 13 References 13 Chapter2 Fruit—AnAngiospermInnovation 21 SandraKnappandAmyLitt Introduction 21 FruitintheFossilRecord 30 FruitVariationandAngiospermPhylogeny 32 FruitDevelopment 33 FruitasaDriverofAngiospermDiversity 36 Acknowledgments 38 References 38 Chapter3 EthyleneandtheControlofFruitRipening 43 DonGrierson Introduction 43 EthyleneandClimactericandNonclimactericFruits 46 AMolecularExplanationforSystem-1andSystem-2Ethylene 48 EthyleneandRipeningGeneNetworksinFlowerandFruitDevelopment 53 EthylenePerceptionandSignaling 54 EthyleneResponseFactors 60 v BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm vi CONTENTS EthyleneandRipeningGeneExpression 60 Conclusions 67 Acknowledgments 68 References 68 Chapter4 CarotenoidBiosynthesisandChlorophyllDegradation 75 PeterM.Bramley Introduction 75 DistributionofCarotenoidsandChlorophyllsinFruit 75 ChlorophyllDegradationandRecycling 78 CarotenoidsandCarotenoidMetabolites 82 FuturePerspectives 100 Acknowledgments 102 Bibliography 102 Chapter5 PhenylpropanoidMetabolismandBiosynthesisofAnthocyanins 117 LauraJaakola Introduction 117 CinnamicAcids 119 Monolignols,Lignans,andLignin 120 Coumarins 120 Stilbenoids 122 Flavonoids 122 EngineeringElevatedLevelsofFlavonoidsandOtherPhenylpropanoids 128 Conclusion 129 References 129 Chapter6 BiosynthesisofVolatileCompounds 135 AntonioGranellandJose´ LuisRambla Introduction 135 MetabolicPathways 136 IdentificationofQuantitativeTraitLociforVolatiles 152 MetabolicEngineeringoftheFruitVolatilePathways 153 FuturePerspectives 154 References 155 Chapter7 CellWallArchitectureandMetabolisminRipeningFruitandthe ComplexRelationshipwithSoftening 163 ElielRuiz-MayandJocelynK.C.Rose Introduction 163 BuildingBlocksofFruitCellWalls 164 BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm CONTENTS vii TheArchitectureofFruitCellWalls 168 CellWallDynamicsinRipeningFruit 171 TheCuticularCellWallandFruitSoftening 177 Summary 179 Acknowledgments 180 References 180 Chapter8 RegulatoryNetworksControllingRipening 189 BetsyAmpopho,NatalieChapman,GrahamB.Seymour, andJamesJ.Giovannoni HormonalControl 189 GeneticNetworks 191 EpigeneticRegulation 200 References 201 Index 207 BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm List of Contributors BetsyAmpopho BoyceThompsonInstituteforPlantScienceResearch CornellUniversity Ithaca,NewYork,NY,USA. PeterMBramley SchoolofBiologicalSciences RoyalHolloway UniversityofLondon Egham,Surrey,UnitedKingdom. NatalieChapman PlantandCropScienceDivision UniversityofNottingham SuttonBonington,Loughborough,Leics,UnitedKingdom. AlisdairR.Fernie DepartmentofMolecularPhysiology Max-Planck-InstituteforMolecularPlantPhysiology Potsdam-Golm,Germany. JamesJGiovannoni DepartmentofAgriculture–AgriculturalResearchService BoyceThompsonInstituteforPlantScienceResearch CornellUniversity Ithaca,NewYork,NY,USA. AntonioGranell InstitutodeBiolog´ıaMolecularyCelulardePlantas ConsejoSuperiordeInvestigacionesCient´ıficas UniversidadPolite´cnicadeValencia Valencia,Spain. DonGrierson LaboratoryofMolecularPhysiologyandBiotechnology ZhejiangUniversity Zhejiang,China. DivisionofPlantandCropSciences SchoolofBiosciences UniversityofNottingham SuttonBoningtonCampus Loughborough,Leicestershire,UnitedKingdom. ix BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm x LISTOFCONTRIBUTORS LauraJaakola DepartmentofBiology UniversityofOulu Oulu,Finland. SandraKnapp DepartmentofBotany TheNaturalHistoryMuseum London,UnitedKingdom. AmyLitt TheNewYorkBotanicalGarden Bronx,NewYork,NY,USA. SoniaOsorio Max-Planck-InstituteforMolecularPlantPhysiology Potsdam-Golm,Germany. MervinPoole PlantScienceDivision UniversityofNottingham SuttonBoningtonCampus Loughborough,Leics,UnitedKingdom. JoseLuisRambla InstitutodeBiolog´ıaMolecularyCelulardePlantas ConsejoSuperiordeInvestigacionesCient´ıficas UniversidadPolite´cnicadeValencia Valencia,Spain. JocelynK.C.Rose DepartmentofPlantBiology CornellUniversity Ithaca,NewYork,NY,USA. ElielRuiz-May DepartmentofPlantBiology CornellUniversity Ithaca,NewYork,NY,USA. GrahamB.Seymour PlantandCropScienceDivision UniversityofNottingham SuttonBonington Loughborough,Leics,UnitedKingdom. GregoryA.Tucker SchoolofBiosciences UniversityofNottingham SuttonBoningtonCampus Loughborough,Leics,UnitedKingdom. BLBS125-fm BLBS125-Seymour Printer:YettoCome March7,2013 0:32 244mm×172mm Preface Evolutionhasfashionedmultiplemeansofprotectingseedanddispersingthemuponmaturation. Noneisasfascinatingnorasconsequentialtohumankindastheripeanddelectablefleshyfruit. Ripefruitscompriseasignificantandexpandingproportionofhumanandanimaldiets,which themedicalcommunitycontendsshouldonlybeincreased.Inadditiontobeingvisualdelights withseductivetastesandaromas,ripefruitsdeliveradiversearrayofantioxidantsandnutrients to those who consume them, in addition to healthy doses of carbohydrates and fiber. The chemistry of fruits comprises attributes that producers, processors, and distributors alike seek tounderstand,optimize,anddelivertoincreasinglyhealth-consciousconsumersexpectinghigh quality and diversity of choices. Plant scientists have endeavored to unravel the mysteries of fleshyfruitbiologyandtheunderlyingmolecularandbiochemicalprocessesthatcontributeto fruitripeningandtheresultingdesirableattributesoffruitsandfruitproducts. Thisbookoffersausefuloverviewoffruitontologyandevolutionemphasizingtheexponen- tialgrowthinadvancesanddiscoveriesinripening-relatedchemistryandassociatedregulatory processesaccumulatedinthelastdecade.Thereaderwillappreciatethebroadanddeepimpact of comprehensive genomics and metabolomics in addition to the computational tools neces- sary to decipher the resulting data on the progress of the field. As a consequence of these all-encompassingapproaches,fruitbiologyhasadvancedfromtheinvestigationofsinglegenes and enzymatic reactions to the development of nuanced molecular regulatory models over- seeingcomplexbiochemicalpathwaysleadingtonumerousmetabolicoutputs.Lookingatthe physiological and molecular symphony of events impacting textural changes of the ripening fruit,thearrayofnovelphenolicmetabolites,orthenetworkofgenesandsignalingprocesses regulatingethylenehormoneresponse,itbecomesstrikinglyclearthatrecenttechnicaladvances have moved ripening biology forward at an astounding rate. This book captures the advances ofthefieldandcouchestheminanevolutionarycontextandafundamentalknowledgeoffruit biology, making it an excellent primer for those interested in the field and a comprehensive referenceforthosefamiliarwithit.TheMolecularBiologyandBiochemistryofFruitRipening is essential reading for any student of plant science and those especially interested in fruit biologyanditsrelationshiptohumandietandnutrition. xi BLBS125-c01 BLBS125-Seymour Printer:YettoCome March6,2013 18:9 244mm×172mm 1 Biochemistry of Fruit Ripening SoniaOsorioandAlisdairR.Fernie Introduction Thischapterisintendedtoprovideanoverviewofthekeymetabolicandregulatorypathways involved in fruit ripening, and the reader is referred to more detailed discussions of specific topicsinsubsequentchapters. The quality of fruit is determined by a wide range of desirable characteristics such as nutritional value, flavor, processing qualities, and shelf life. Fruit is an important source of supplementarydiet,providingminerals,vitamins,fibers,andantioxidants.Inparticular,theyare generallyrichsourcesofpotassium,folate,vitaminsC,E,andKaswellasotherphytonutrients such as carotenoids (beta-carotene being a provitamin A) and polyphenols such as flavonols (Saltmarshetal.,2003).Asimilar,butperhapsmoredisparate,groupofnutrientsisassociated with vegetables. Thus nutritionists tend to include fruits and vegetables together as a single “food group,” and it is in this manner that their potential nutritional benefits are normally investigated and reported. Over the past few decades, the increased consumption of fruits and vegetables has been linked to a reduction in a range of chronic diseases (Buttriss, 2012). This has led the WHO to issue a recommendation for the consumption of at least 400 g of fruits and vegetables per day. This in turn has prompted many countries to issue their own recommendationsregardingtheconsumptionoffruitsandvegetables.InBritainthishasgiven risetothefive-a-dayrecommendation.AportionintheUnitedKingdomisdeemedtobearound 80g;sofive-a-daycorrespondstoabout400gperday.Othercountrieshaveoptedfordifferent recommendations(Buttriss,2012),butallrecognizetheneedforincreasedconsumption. The rationale for the five-a-day and other recommendations to increase fruit and vegetable consumption comes from the potential link between high intake of fruits and vegetables and low incidence of a range of diseases. There have been many studies carried out over the last few decades. The early studies tended to have a predominance of case-control approaches whilerecentlymorecohortstudies,whichareconsideredtobemorerobust,havebeencarried out. This has given rise to many critical and systematic reviews, examining this cumulative TheMolecularBiologyandBiochemistryofFruitRipening,FirstEdition. EditedbyGrahamB.Seymour,MervinPoole,JamesJ.GiovannoniandGregoryA.Tucker. ©2013JohnWiley&Sons,Inc.Published2013byJohnWiley&Sons,Inc. 1
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