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FOODANDNUTRITION RESEARCH

ADVISORYBOARDS

DavidRodríguez-Lázaro

Loong-TakLim

MichaelEskin

IsabelFerreira

CrispuloGallegos

Se-KwonKim

KeizoArihara

SERIESEDITORS

GEORGEF.STEWART(1948–1982)

EMILM.MRAK(1948–1987)

C.O.CHICHESTER(1959–1988)

BERNARDS.SCHWEIGERT(1984–1988)

JOHNE.KINSELLA(1989–1993)

STEVEL.TAYLOR(1995–2011)

JEYAKUMARHENRY(2011–2016)

FIDELTOLDRÁ(2016– )

ADVANCESIN FOODANDNUTRITION RESEARCH

Editedby FIDELTOLDRÁ

InstitutodeAgroquímicayTecnologíadeAlimentos(CSIC), Valencia,Spain

AcademicPressisanimprintofElsevier

50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates 525BStreet,Suite1650,SanDiego,CA92101,UnitedStates TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom 125LondonWall,London,EC2Y5AS,UnitedKingdom

Firstedition2019

Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans,electronic ormechanical,includingphotocopying,recording,oranyinformationstorageandretrievalsystem, withoutpermissioninwritingfromthepublisher.Detailsonhowtoseekpermission,further informationaboutthePublisher’spermissionspoliciesandourarrangementswithorganizationssuch astheCopyrightClearanceCenterandtheCopyrightLicensingAgency,canbefoundatourwebsite: www.elsevier.com/permissions.

Thisbookandtheindividualcontributionscontainedinitareprotectedundercopyrightbythe Publisher(otherthanasmaybenotedherein).

Notices

Knowledgeandbestpracticeinthisfieldareconstantlychanging.Asnewresearchandexperience broadenourunderstanding,changesinresearchmethods,professionalpractices,ormedical treatmentmaybecomenecessary.

Practitionersandresearchersmustalwaysrelyontheirownexperienceandknowledgeinevaluating andusinganyinformation,methods,compounds,orexperimentsdescribedherein.Inusingsuch informationormethodstheyshouldbemindfuloftheirownsafetyandthesafetyofothers,including partiesforwhomtheyhaveaprofessionalresponsibility.

Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors,assume anyliabilityforanyinjuryand/ordamagetopersonsorpropertyasamatterofproductsliability, negligenceorotherwise,orfromanyuseoroperationofanymethods,products,instructions,orideas containedinthematerialherein.

ISBN:978-0-12-817171-4

ISSN:1043-4526

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Prefacexiii

1.Geneticdeterminantsofbeverageconsumption:Implications fornutritionandhealth1

MarilynC.Cornelis

1. Introduction2

2. Determinantsofbeverageconsumption3

3. Geneticdeterminantsofbeverageconsumption4

4. Implicationsofgeneticknowledgeonbeverageconsumption28

5. Conclusions35 Acknowledgment36 References36 Furtherreading52

2.Currentfeedingstrategiestoimproveporkintramuscularfat contentanditsnutritionalquality53

C.M.Alfaia,P.A.Lopes,M.S.Madeira,J.M.Pestana,D.Coelho,F.Toldrá, andJ.A.M.Prates

1. Introduction54

2. Qualityofporkfatandfattyacids55

3. Feedingstrategiestoincreaseporkintramuscularfatcontent60

4. Feedingstrategiestoimproveporkfattyacidprofile70

5. Concludingremarksandchallenges84 Acknowledgments85 References86 Furtherreading94

3.Dairyfoodsandpositiveimpactontheconsumer’shealth95 SilvaniVerruck,CelsoFasuraBalthazar,RamonSilvaRocha,RamonSilva, ErickAlmeidaEsmerino,TatianaColomboPimentel,M ^ onicaQueirozFreitas, MarciaCristinaSilva,AdrianoGomesdaCruz, andElaneSchwindenPrudencio

1. Introduction96

2. Fermentedmilks98

3. Cheese112

4. Butter121

5. Icecream132

6. Dairydesserts141

7. Conclusions145 References145 Furtherreading164

4.Food-derivedbioactivepeptidesandtheirroleinameliorating hypertensionandassociatedcardiovasculardiseases165 AdvaitaGanguly,KumakshiSharma,andKaustavMajumder

1. Introduction166

2. Pathophysiologyofhypertension167

3. Food-derivedbioactivepeptides175

4. Structuralfeaturesofbioactivepeptides183

5. Molecularmechanismsoffood-derivedantihypertensivepeptides190

6. Conclusion193 References193 Furtherreading207

5.Effectsofphytochemicalsagainstdiabetes209 MerveBacanli,SevtapAydinDilsiz,NurşenBaşaran,andA.AhmetBaşaran

1. Introduction209

2. Diabetes210

3. Diabetesandoxidativestress211

4. Phytochemicals213

5. Phytochemicalsanddiabetes216

6. Conclusion229 References230

6.DMHF(2,5-dimethyl-4-hydroxy-3(2H)-furanone),avolatilefood componentwithattractivesensoryproperties,brings physiologicalfunctionsthroughinhalation239 K.Arihara,I.Yokoyama,andM.Ohata

1. Introduction240

2. WhatistheMaillardreaction?241

3. VolatilecomponentsgeneratedbytheMaillardreaction242

4. DMHFandfoods244

5. Physiologicalfunctionsofodors245

6. DMHFonphysiologicalparametersthroughinhalation246

7. Conclusions254 References254

7.Challengesandopportunitiesregardingtheuseofalternative proteinsources:Aquacultureandinsects259

BelenGómez,PauloE.S.Munekata,ZhenzhouZhu,FranciscoJ.Barba, FidelToldrá,PredragPutnik,DanijelaBursacKovacevic,andJoseM.Lorenzo

1. Introduction260

2. Aquacultureproductsnutritionalcomposition265

3. Insectsnutritionalcomposition272

4. Extractionmethodstorecoverproteins278

5. Analysisoftheextracts279

6. Purificationandfractionationstages283

7. Developmentofnewproductsbasedoninsectproteinsandaquaculture products284

8. Challengesandfutureperspectivesofaquacultureproductsasprotein sources286

9. Challengesandfutureperspectivesofinsectsasproteinssource288 Acknowledgments289 References289

8.Mycotoxinsinfoodandfeed297

JelkaPleadin,JadrankaFrece,andKsenijaMarkov

1. Introduction298

2. Propertiesanddivision300

3. Factorsfacilitatingmycotoxinsynthesis302

4. Toxicogenicmolds305

5. Occurrenceinfoodandfeed308

6. Toxiceffects312

7. Majorgroupsandrepresentatives314

8. Analyticalmethods325

9. Preventativemeasures328

10. Reductionmethods331

11. Conclusion337 Acknowledgment338 References338 Furtherreading345

Contributors

C.M.Alfaia

CentrodeInvestigac ¸aoInterdisciplinaremSanidadeAnimal(CIISA),FaculdadedeMedicina Veterina ´ ria,UniversidadedeLisboa,AvenidadaUniversidadeTecnica,Po ´ loUniversita ´ rio doAltodaAjuda,Lisbon,Portugal

K.Arihara

SchoolofVeterinaryMedicine,KitasatoUniversity,Tokyo,Japan

MerveBacanli

FacultyofPharmacy,DepartmentofPharmaceuticalToxicology,HacettepeUniversity, Ankara,Turkey

CelsoFasuraBalthazar

UniversidadeFederalFluminense(UFF),FaculdadedeVeterina ´ ria,Nitero ´ i,Brazil

FranciscoJ.Barba

NutritionandFoodScienceArea,PreventiveMedicineandPublicHealth,FoodScience, ToxicologyandForensicMedicineDepartment,FacultyofPharmacy,Universitatde Vale ` ncia,Vale ` ncia,Spain

A.AhmetBaşaran

FacultyofPharmacy,DepartmentofPharmacognosy,HacettepeUniversity,Ankara,Turkey

NurşenBaşaran

FacultyofPharmacy,DepartmentofPharmaceuticalToxicology,HacettepeUniversity, Ankara,Turkey

DanijelaBursacKovacevic

FacultyofFoodTechnologyandBiotechnology,UniversityofZagreb,Zagreb,Croatia

D.Coelho

CentrodeInvestigac ¸ a ˜ oInterdisciplinaremSanidadeAnimal(CIISA),FaculdadedeMedicina Veterina ´ ria,UniversidadedeLisboa,AvenidadaUniversidadeTecnica,Po ´ loUniversita ´ rio doAltodaAjuda,Lisbon,Portugal

MarilynC.Cornelis

DepartmentofPreventiveMedicine,NorthwesternUniversityFeinbergSchoolof Medicine,Chicago,IL,UnitedStates

AdrianoGomesdaCruz

InstitutoFederaldeEducac ¸ao,Ci^ enciaeTecnologiadoRiodeJaneiro(IFRJ), DepartamentodeAlimentos,RiodeJaneiro,Brazil

SevtapAydinDilsiz

FacultyofPharmacy,DepartmentofPharmaceuticalToxicology,HacettepeUniversity, Ankara,Turkey

ErickAlmeidaEsmerino

UniversidadeFederalFluminense(UFF),FaculdadedeVeterina ´ ria,Nitero ´ i,Brazil

JadrankaFrece

FacultyofFoodTechnologyandBiotechnology,UniversityofZagreb,Zagreb,Croatia

M ^ onicaQueirozFreitas

UniversidadeFederalFluminense(UFF),FaculdadedeVeterina ´ ria,Nitero ´ i,Brazil

AdvaitaGanguly

ComprehensiveTissueCentre,UAHTransplantServices,AlbertaHealthServices, Edmonton,AB,Canada

BelenGo ´ mez

CentroTecnolo ´ gicodelaCarnedeGalicia,Ourense,Spain

P.A.Lopes

JoseM.Lorenzo

CentroTecnolo ´ gicodelaCarnedeGalicia,Ourense,Spain

M.S.Madeira

KaustavMajumder

DepartmentofFoodScienceandTechnology,UniversityofNebraska-Lincoln,Lincoln, NE,UnitedStates

KsenijaMarkov

FacultyofFoodTechnologyandBiotechnology,UniversityofZagreb,Zagreb,Croatia

PauloE.S.Munekata

CentroTecnolo ´ gicodelaCarnedeGalicia,Ourense,Spain;DepartmentofFood Engineering,CollegeofAnimalScienceandFoodEngineering,UniversityofSaoPaulo, SaoPaulo,Brazil

M.Ohata

CollegeofBioresourceSciences,NihonUniversity,Tokyo,Japan

J.M.Pestana

TatianaColomboPimentel

InstitutoFederaldoParana ´ (IFPR),CampusParanavaı´,Paranavaı´,Brazil JelkaPleadin

CroatianVeterinaryInstitute,LaboratoryforAnalyticalChemistry,Zagreb,Croatia

J.A.M.Prates

ElaneSchwindenPrudencio

UniversidadeFederaldeSantaCatarina(UFSC),DepartamentodeCi^ enciaeTecnologiade Alimentos,Floriano ´ polis,Brazil

PredragPutnik

FacultyofFoodTechnologyandBiotechnology,UniversityofZagreb,Zagreb,Croatia

KumakshiSharma

Health,SafetyandEnvironmentBranch,NationalResearchCouncilCanada,Edmonton, AB,Canada

MarciaCristinaSilva

InstitutoFederaldeEducac ¸ao,Ci^ enciaeTecnologiadoRiodeJaneiro(IFRJ), DepartamentodeAlimentos,RiodeJaneiro,Brazil

RamonSilva

UniversidadeFederalFluminense(UFF),FaculdadedeVeterina ´ ria,Nitero ´ i;InstitutoFederal deEducac ¸ao,Ci^ enciaeTecnologiadoRiodeJaneiro(IFRJ),DepartamentodeAlimentos, RiodeJaneiro,Brazil

RamonSilvaRocha

UniversidadeFederalFluminense(UFF),FaculdadedeVeterina ´ ria,Nitero ´ i;InstitutoFederal deEducac ¸ao,Ci^ enciaeTecnologiadoRiodeJaneiro(IFRJ),DepartamentodeAlimentos, RiodeJaneiro,Brazil

FidelToldra ´

InstitutodeAgroquı´micayTecnologı´adeAlimentos(CSIC),Valencia,Spain SilvaniVerruck

UniversidadeFederaldeSantaCatarina(UFSC),DepartamentodeCi^ enciaeTecnologiade Alimentos,Floriano ´ polis,Brazil

I.Yokoyama

SchoolofVeterinaryMedicine,KitasatoUniversity,Tokyo,Japan

ZhenzhouZhu

CollegeofFoodScienceandEngineering,WuhanPolytechnicUniversity,Wuhan,China

Preface

Thisvolumeof AdvancesinFoodandNutritionResearch compileseight chaptersreportingavarietyofinterestingtopicslikethegeneticdeterminantsofbeverageconsumptionanditshealthimplications,thecurrent feedingstrategiestoimproveporkintramuscularfatcontentanditsnutritionalquality,theimpactofdairyfoodsonconsumers’health,thebenefits offood-derivedbioactivepeptidesforhypertensionandassociatedcardiovasculardiseases,theeffectsofphytochemicalsagainstdiabetes,thevolatile foodcomponentDMHFthatimprovespalatabilityandphysiological functions,theuseofalternativeproteinsourcesfromaquacultureand insects,andtheoccurrenceandpreventionofmycotoxinsinfoods.

Chapter1 bringsnewinsightsonthegeneticdeterminantsofcommon beverageconsumptionandhowresearchinthisfieldiscontributinginsight towhatandhowmuchweconsumeandwhythisgeneticknowledge mattersfromaresearchandpublichealthperspective,andalsohowthe genome-wideassociationstudieshighlightanimportantbehavior-reward component(asopposedtotaste)tobeverageconsumptionwhichmayserve asapotentialbarriertodietaryinterventions. Chapter2 addressescurrent feedingstrategiestoameliorateporksensoryattributesandnutritional qualitybyincreasingintramuscularfatdepositionandimprovingfattyacid composition,respectively.Italsodiscussesfeedingsourcesofn-3polyunsaturatedfattyacidstopigs,mainlyfrommarineorigin(richin eicosapentaenoicanddocosahexaenoicacids),thatincreasetheircontent inpork,thusimprovingthehealthvalueofitsfattyacidprofile,andhow theinclusionofmicroalgaeandseaweedsinfeedrepresentsapromising approachforthemaintenanceanddevelopmentofthelivestocksector.

Chapter3 bringsnewinsightsonthepositiveimpactsoffermentedmilk, cheese,butter,icecream,anddairydessertscomponentsontheconsumer’s health.Aspecialfocusisgiventothemainmechanismsresponsibleand essentialforabetterunderstandingofnutritionalandfunctionalvaluesof thecomponentsofmilkanddairyproducts. Chapter4 presentspotent bioactivepeptides,especiallywithantiinflammatory,antioxidant,and angiotensin-convertingenzyme-Iinhibitoryactivity,fromdifferentfood sourcesasapromisingendeavortowardnutraceutical-baseddietarymanagementandpreventionofhypertension.Italsodiscusseshowtheunderstandingofthepathophysiologyofhypertensionandtheactionmechanismsof

thebioactivepeptideswouldhelpinthedesignandcharacterizationofmore potentpeptides. Chapter5 focusesontherelationshipbetweendiabetes mellitusandpreventiverolesofvariousphytochemicals.Someofthemsuch asflavonoids,lignans,andprophenylphenolscanplayapreventiveroleon diabetesviatheirantioxidantproperties. Chapter6 dealswithavolatile foodcomponentDMHF(2,5-dimethyl-4-hydroxy-3(2H)-furanone),with attractivesensoryproperties.Thissubstanceisanaromacompound,generatedinvariousfoodsbytheMaillardreactionduringcookingand processing,thatexhibitsastrawberry-likeflavorwhendilutedanda caramel-likearomawhenconcentrated.Itbringsphysiologicalfunctions andbioactivitythroughinhalation. Chapter7 addressesthepotentialof insectsandproductsderivedfromaquacultureassustainablealternative proteinsources.Italsodiscussesmajorchallengesliketheneedtoadapt technologiesandmethodsfortheproductionandwell-characterizationof thenewingredients,theevaluationofsuchnewproteinsinthedietand itssafetyofuse,includingpotentialallergies,andtheacceptanceby consumers.Finally, Chapter8 bringsthelatestdevelopmentsinmycotoxins infoods.Itdescribestheharmfuleffectsofmycotoxinsobservedinhumans andanimalslikecarcinogenicity,teratogenicity,immunetoxicity, neurotoxicity,hepatotoxicity,nephrotoxicity,reproductiveanddevelopmentaltoxicity,indigestion,andsoforth.Thechapterincludesthedescriptionofpreventativemeasurescapableofreducingthecontaminationtothe minimumaswellasmethodsformycotoxinreductionorelimination.

Insummary,thisvolumepresentsthecombinedeffortsof38professionalsdevelopingtheirresearchin8countries(Canada,USA,Brazil, China,Turkey,Japan,Croatia,Portugal,andSpain)withavarietyof backgroundsandexpertise.TheEditorwishestothanktheproductionstaff andallthecontributorsforsharingtheirexperienceandformakingthisbook possible.

FIDEL TOLDRA ´ Editor

Geneticdeterminantsofbeverage consumption:Implicationsfor nutritionandhealth

MarilynC.Cornelis*

DepartmentofPreventiveMedicine,NorthwesternUniversityFeinbergSchoolofMedicine,Chicago,IL, UnitedStates

*Correspondingauthor:e-mailaddress:marilyn.cornelis@northwestern.edu

Contents

1. Introduction2

2. Determinantsofbeverageconsumption3

3. Geneticdeterminantsofbeverageconsumption4

3.1 Earlyapproaches:Linkageandcandidategeneanalysis5

3.2 Recentapproaches:Genome-wideanalysis7

4. Implicationsofgeneticknowledgeonbeverageconsumption28

4.1 Researchtools28

4.2 Publichealth34

5. Conclusions35 Acknowledgment36 References36 Furtherreading52

Abstract

Beveragesmakeimportantcontributionstonutritionalintakeandtheirroleinhealth hasreceivedmuchattention.Thisreviewfocusesonthegeneticdeterminantsofcommonbeverageconsumptionandhowresearchinthisfieldiscontributinginsightto whatandhowmuchweconsumeandwhythisgeneticknowledgemattersfroma researchandpublichealthperspective.Theearliesteffortsingene-beveragebehavior mappinginvolvedgeneticlinkageandcandidategeneanalysisbuttheseapproaches havebeenlargelyreplacedbygenome-wideassociationstudies(GWAS).GWAShave identifiedbiologicallyplausiblelociunderlyingalcoholandcoffeedrinkingbehavior. NoGWAShasidentifiedvariantsspecificallyassociatedwithconsumptionoftea,juice, soda,wine,beer,milkoranyothercommonbeverage.Thusfar,GWAShighlightan importantbehavior-rewardcomponent(asopposedtotaste)tobeverageconsumption whichmayserveasapotentialbarriertodietaryinterventions.Lociidentifiedhavebeen usedinMendelianrandomizationandgene beverageinteractionanalysisofdisease

butresultshavebeenmixed.ThisresearchisnecessaryasitinformstheclinicalrelevanceofSNP-beverageassociationsandthusgenotype-basedpersonalizednutrition, whichisgaininginterestinthecommercialandpublichealthsectors.

1.Introduction

Waterisanessentialnutrientforlife( Jequier&Constant,2009).Beveragescontributeapproximately80%tototalwaterintakewiththeremainderprovidedbysolidfoods(EFSA,2010; Electrolytes&Water,2005).After water,coffee,tea,beer,milk,100%juice,sugarsweetenedbeverages(SSB) andwineareamongthemostwidelyconsumedbeveragesintheworld (Euromonitor,2018; Neves,Trombin,Lopes,Kalaki,&Milan,2012; Singhetal.,2015).Unlikeplainwater,beveragesarealsoimportantsources ofenergy,othervitaminsandmineralsaswellas1000sofnon-nutrients, manyofwhicharebioactive.Coffeeandtea,forexample,arenaturally energy-freebutimportantsourcesofcaffeineandpolyphenols.Beerand winebothcontainalcoholbutalsopresentwithuniqueconstituentsincludinghopsandresveratrol,respectively.JuiceandSSBarebothenergydense buttheformerisalsoanaturalsourceofvitamins.

Withtheirwideconsumptionandcontributionstonutrientintakethere isgreatinterestintherolebeveragesplayinhealth.Epidemiologicalstudies supportabeneficialroleofmoderatecoffeeintakeinreducingriskofseveral chronicdiseasesbutheavyintakeislikelyharmfulonpregnancyoutcomes (Pooleetal.,2017).Teamightalsoreduceriskoftype2diabetes(T2D), metabolicsyndrome(MetS)(Marventanoetal.,2016; Yang,Mao,Xu, Ma,&Zeng,2014),osteoporosis(Sunetal.,2017)andcardiovasculardiseases(CVD)(Pangetal.,2016).Winedrinkingmayhaveadose-dependent associationwithhealth:lowdosesmightprotectagainstbreastcancerand CVDwhilehighdosesoffernobenefitorincreasedrisk(Chenetal., 2016; deGaetano,DiCastelnuovo,Rotondo,Iacoviello,&Donati, 2002).Healthbenefitsorrisksspecifictobeerandmilkareunclear (Gijsbersetal.,2016; Guoetal.,2017; Kaplan,Palmer,&Denke,2000; Larsson,Crippa,Orsini,Wolk,&Michaelsson,2015; Lee,Fu,Chung, Jang,&Lee,2018; Lietal.,2011; Liuetal.,2015; Mullie,Pizot,& Autier,2016; Soedamah-Muthuetal.,2011).Therearecurrentlyno healthbenefitstoSSBconsumptionbutratherconvincingdatasupporting itsroleinobesitywhichis,inturn,ariskfactorforseveraldiseases

(Maliketal.,2010; Malik,Schulze,&Hu,2006 ).Acaveattoourknowledgepertainingtobeverageconsumptionandhumanhealthisthatmuch ofitisderivedfromepidemiologic alstudieswhichhavelimitations (Rothman,Greenland,&Lash,2008 ; Taubes,1995 ; Willett,1998 ).

Thisreviewfocusesonthegeneticdeterminantsofcommonbeverage consumptionandhowresearchinthisfieldiscontributinginsighttowhat andhowmuchweconsumeandwhythisgeneticknowledgemattersfroma researchandpublichealthperspective.

2.Determinantsofbeverageconsumption

Understandingfactorscontributingtobeverageconsumptionhas importantpublichealthandresearchimplications.Knowledgeofboth externalandinternalcuesforbeverageintakemayinformthecausalrole eachbeveragehasinhealth(research)andthepotentialpopulationsubgroupsmostsusceptibletothehealthconsequencesofitsregularconsumption(publichealth).Thirstisanimportantdeterminantofbeverageintake, butintoday’ssocietytheamountandchoiceofbeverageconsumedis governedbyamultitudeofindividualandsocietalfactorssuchasavailability, mood,socialcontext,healthstatus,education,convenience,cost,cultural influences,andsensoryattributessuchassmellandtaste(Block,Gillman, Linakis,&Goldman,2013; Drewnowski,1997; Drewnowski, Henderson,Levine,&Hann,1999; Glanz,Basil,Maibach,Goldberg,& Snyder,1998; Neumark-Sztainer,Story,Perry,&Casey,1999; Wardle, Carnell,&Cooke,2005).Consumptionofcoffee,forexample,tendsto positivelycorrelatewithage,smoking,andalcoholconsumptionandmay alsobeimpactedbyperceivedhealthconsequencesofthebeverage (Cornelis,2012).Degreeofeconomicdevelopment,religiousandcultural norms,theavailabilityandthelevelandeffectivenessofpoliciesaresocietal factorscontributingtoalcoholconsumptionbehaviorswhileage,gender, socialeconomicstatus(SES)andprioralcoholexposureareimportantindividuallevelfactors(Baboretal.,2010).Theacutepositiveornegativereinforcingpropertiesofalcoholandcaffeineareespeciallyimportantin determiningalcoholicbeverageandcoffeedrinkingpatterns(Cornelis, 2012; Koob&Volkow,2016; Kuntsche,Knibbe,Gmel,&Engels, 2005).Physiologicaleffectsofbeveragesareintrinsicandoftenvarybetween individuals.Geneticsalsoplayaroleinbeverageconsumptionbehaviorand morelikelywithregardstothesephysiologicaleffects.

3.Geneticdeterminantsofbeverageconsumption

Twinstudieshavelargelybeenthestandardapproachestoestimating thegeneticorheritablecomponentofbeverageconsumptionbehaviors whichcanvaryfrom0(notheritable)to1(completelyinherited).Twin studiesestimateheritabilitybycomparingmonozygotictwins,whoshare thecommonenvironmentandhaveidenticalgenetics,todizygotictwins, whoalsosharethecommonenvironmentbutonlyhalftheirgenetics (Neale&Cardon,1994; Turkheimer,D’Onofrio,Maes,&Eaves,2005).

Inanearliertwinstudyby deCastro(1993),theheritabilityestimatefor amountofdrinkingwaterconsumedwas0.43,whichwasslightlyhigher thanthe0.37estimatefortotalwater(foodandbeverages).Heritabilityestimatesforalcohol,soda,milk,coffeeandfruitjuicewere0.45,0.54,0.04, 0.69and0.12,respectively. Hasselbalchetal.(2010) reportedlowerestimatesforsoda:0.26and0.30formenandwomen,respectively,while Teucheretal.(2007) reportedamuchhigherestimateforfruitjuice (0.70).Heritabilityestimatesforself-reportedtotalcaffeineintake(derived fromcaffeine-containingcoffee,teaandsoda)rangedbetween0.30and 0.58,withhigherestimatesreportedforheavyuse(upto0.77)(Yang, Palmer,&deWit,2010).Studiesthatseparatedheritabilityestimatesbycaffeinesourcereporthigherheritabilityforcoffeerelativetoothersources (Luciano,Kirk,Heath,&Martin,2005; Teucheretal.,2007; Vink, Staphorsius,&Boomsma,2009).Twinstudiesaswellasfamilyandadoption studiesofhabitualalcoholconsumptionoralcoholismhavelargelyfocused onmalesandreportheritabilityestimatesbetween0.30and0.60(McGue, 1999; Prescott&Kendler,1999; Reedetal.,1994; Teucheretal.,2007). HeathandMartin(1988) proposethatthedecisiontoabstainfromdrinking isnotgeneticallydetermined,buttheonsetandamountconsumedoncethat decisionismadeareinfluencedbygeneticfactors.

Twinstudiesarepowerfulepidemiologicalapproachestomeasuring thecontributionofgeneticstoagiventraitbutareoftenunderpowered andsubjecttobiaswhichlikelyaddtobetweenstudydifferencesinestimates ofheritability(Kendler,1993; Zaitlen&Kraft,2013; Zuk,Hechter, Sunyaev,&Lander,2012).Furthermore,althoughresultsaboveprovide compellingevidenceforageneticinfluenceonbeverageintakeandchoice, theydonotindicatethespecificgenesthatincreaseordecreasedrinking behaviors.

3.1Earlyapproaches:Linkageandcandidategeneanalysis

Theearliesteffortsingene-traitmappinginvolvedgeneticlinkageandcandidategeneanalysis.Geneticlinkagestudiesdetermineinheritanceofa binaryorquantitativetraitamongfamilymembersinextensivepedigrees byevaluatingwhetheroneormoregeneticmarkersspacedacrossthe 23chromosomessegregatewiththetrait(Cantor,2014).Thisapproach roughlylocatesabroadchromosomalregion,whichmaycontain10sof 100sofgenes,thatco-segregatewiththetrait.Otherstrategiessuchasfine mappingandtargetedassociationanalysismustbeusedtofurtherrefinethe linkedregionandidentifythegeneofinterest(Cantor,2014).

Lactoseintolerance(orlactasenon-persistence)isacommonautosomal recessiveconditionresultingfromthephysiologicaldeclineinactivityof lactase-phlorizinhydrolase(LPH)inintestinalcellsafterweaningandhas asignificantimpactonmilkdrinkingbehavior.Theageofonsetvaries betweenpopulationsandinsomepopulationslactaseactivitypersistsata highlevelthroughoutadultlife(Sahi,1994; Sahi,Isokoski,Jussila,& Launiala,1972; Swallow,2003; Wangetal.,1998).Althoughthesequence ofthelactasegene(LCT,encodingLPH)hadbeenknownsince1991(Boll, Wagner,&Mantei,1991),thecausativemechanismforlactasepersistence remainedelusiveuntil2002whenalinkageanalysisofnineFinnishfamilies withhypolactasiaidentifiedavariantupstreamoftheinitiationcodonof LCT (LCT-13910C > T)whichdemonstratedcompleteassociationwith lactasepersistence(Enattahetal.,2002).

Thefirstlinkagestudiesonalcoholdependence(AD)fromthefamilybasedCollaborativeStudyontheGeneticsofAlcoholism(COGA)(Reich etal.,1998)andasib-pairstudyfromaSouthwestAmericanIndiantribe (Longetal.,1998)reportedabroadrisklocusonchromosome4qthatcontainsthegenesthatencodetheisoformsofalcoholdehydrogenase(ADH). Hilletal.(2004) reportedsupportforADlociatchromosome1q23.3-q25.1 inagenome-widelinkageanalysisofdoubleprobands.Thisregionwas followedupusingfine-mappinggenotypingandconfirmedsinglenucleotidepolymorphism(SNP)-ADassociationswithin ASTN1 (Hill,Weeks, Jones,Zezza,&Stiffler,2012).AdditionalADstudieshaveimplicatedother chromosomalregions(Dicketal.,2010; Edenberg&Foroud,2014; Wang etal.,2004).Thegeneticlinkageapproach,totheauthor’sknowledge,has notbeenappliedto habitual beverageconsumption.

Asecondapproachtoisolatinggeneticdeterminantsofatraitinvolves geneticassociationinpopulationorfamily-basestudiesandisessentiallya

formoflinkagemappingbutisallele-basedratherthanlocus-basedandis oftenhypothesisdriven.EffortsfocusonpotentiallyfunctionalSNPsin geneswithbiologicalplausibilityorinregionsidentifiedbylinkage. Inthecontextofbeverageconsumptionbehaviorahighlyimplicated pathwaypertainstotaste.Tasteisoneoftheprimarymeansofdetermining theacceptabilityofafoodandmighthavebeencriticaltothesurvivalof earlyhumansubjects(Tepper,2008).Theperceptionofsweet,umami andbittertastesisallmediatedviaG-coupledproteinreceptors,encoded bythe TAS1R and TAS2R tastereceptorgenefamilies,whilesaltyandsour tastesaretransducedviaionchannels(Wise,Hansen,Reed,&Breslin, 2007).Thereislittleknownregardinggeneticvariationinsaltyand sourtastes(Wiseetal.,2007).Incontrast,bittertastequalityisaffected byvariantsin TAS2R16,TAS2R38,TAS2R43 and TAS2R44 while variantsin TAS1R1 and TAS1R3 impactumamiandsweet(Drayna, 2005; Feeney,O’Brien,Scannell,Markey,&Gibney,2011; Kimetal., 2003; Mainland&Matsunami,2009; Shigemura,Shirosaki,Sanematsu, Yoshida,&Ninomiya,2009).Themoststudiedis TAS2R38,inparticular itsthreeSNPs,whichresultintwocommonhaplotypesthatarenamedfor theiraminoacidsubstitutions:PAV(proline,alanine,andvaline)andAVI (alanine,valine,andisoleucine)(Kim,Breslin,Reed,&Drayna,2004). TAS2R38 diplotypeinfluencestheabilitytotasteafamilyofbittercompoundsincludingphenylthiocarbamide(PTC)and6-n-propylthiouracil (PROP).Althoughthesecompoundsarenotfoundinfoodstuffsnaturally, PTC/PROP-relatedcompoundsarepresentinseveralbittertastingfruits andvegetables.PAVhomozygotesandheterozygotesperceivegreaterbitternessthanAVIhomozygotesthatperceivelittleornobitterness(Kim etal.,2004).Thislocushasbeensubjecttonumerousassociationstudies offoodandbeveragepreferences.Variationin TAS2R38 hasbeenlinked tocoffee,beer,spirits,greentea,sugarcontentofbeveragesandtotalalcohol ordrinkingstatus(Beckettetal.,2017; Choi,Lee,Yang,&Kim,2017; Duffyetal.,2004; Hayesetal.,2011; Mennella,Pepino,&Reed,2005; Ongetal.,2018; Ooi,Lee,Law,&Say,2010; Pernaetal.,2018; Ramos-Lopezetal.,2015; Wangetal.,2007).Variationinother TASR2 membershasalsobeenlinkedtobitterbeverageconsumption(Cornelis, 2012; Hayesetal.,2011; Ongetal.,2018; Pirastuetal.,2014; Wang etal.,2007).Mostofthesefindings,however,lackreplication.Although bitternessiswidelyclaimedtobeanevolutionarilyimportantindicatorof toxicity(Behrens&Meyerhof,2016; Drewnowski&Gomez-Carneros, 2000)notallbitterstimuliaretoxic(Glendinning,1994; Nissim,

Dagan-Wiener,&Niv,2017).Coffeeandbeerareprimeexampleswhereby theinnateeversiontobittertastedoesnothold.Indeed,variantsin TAS2R43 linkedtoincreasedperceptionofcaffeine,abittercompound, associatewithincreasedcoffeeconsumptionandlikingaccordingto candidate-SNPanalysis(Ongetal.,2018; Pirastuetal.,2014).Although coffeebitternessiseasilyoffsetbyadditives,someindividualsmayalsolearn toassociatethissensorycuewithsocialcontextorpostingestivesignals elicitedbybiologicallyactiveconstituentsofcoffee.Variationinthe TAS1R sweetandumamireceptorfamilyhasalsobeenlinkedtoalcoholconsumptionbehavior(Choietal.,2017),winedrinking(Choietal.,2017)and vodkaliking(Pirastuetal.,2012).

Besidestaste-relatedgenes,thecandidateapproachforalcohol-related traitshasadditionallyfocusedonalcoholmetabolismgenesincluding ADH, CYP2E1 and ALDH (Fig.1),aswellasgenemembersofseveralneurotransmittersystems: GIRK1,GABA-A,DRD2,SLC6A3,SLC6A4, TPH1,COMT,CHRM2 and OPRM1 (Edenberg&Foroud,2006; Matsuoetal.,2006; Reilly,Noronha,Goldman,&Koob,2017; Tawa, Hall,&Lohoff,2016).Forcoffeeandteadrinking,candidategenesinvolved incaffeinemetabolism(CYP1A2)andcaffeine’stargetofaction (ADORA2A,DRD2)havebeenexamined(Cornelis,2012; Cornelis, El-Sohemy,&Campos,2007).Genemembersofthebrainrewardsystem, particularly DRD2,havebeentestedforassociationswithSSB(Baik,2013; Eny,Corey,&El-Sohemy,2009; Ramos-Lopez,Panduro,RiveraIn ˜ iguez,&Roman,2018).

3.2Recentapproaches:Genome-wideanalysis

Withthehumangenomesequencedintheearly2000’sandmappedfrequenciesandpatternsofassociationamongmillionsofcommonSNPsin diversepopulations,theprimaryapproachtoidentifyinggeneticvariants forcomplextraitshasquicklytransitionedtogenome-wideassociation

Fig.1 Alcoholmetabolism.

studies(GWAS).GWASisbasedonthepremisethatacausalvariantis locatedonahaplotype,andthereforeamarkeralleleinlinkagedisequilibrium(LD)withthecausalvariantshouldpresentwithanassociationwitha traitofinterest(Hirschhorn&Daly,2005).Thisapproachisunbiasedwith respecttogenomicstructureandpreviousknowledgeofthetraitetiology, whichcontrastswithearlierapproaches,andthereforehasgreaterpotential torevealnovelgene-traitassociations.

Table1 providesdescriptionsofGWASlistedintheGWAScatalog (November26,2018)reportingsignificantSNP-drinkingbehaviorassociations.GWASdesignsincludedsinglesample,2-stage(i.e.,discoveryand replication)andmeta-analysis. Table2 presentsthoselocidefinedassignificantaccordingtoauthors’apriorithreshold.ResultsofGWASofADand beverage“liking”werealsoincluded.

3.2.1Alcohol

SuccessfulGWASofalcohol-relatedtraitshavebeenundertakenin European,AfricanAmerican,AsianandHispanicLatinopopulations.Most ofthesetargetedADandincludedstudysampleswithahighproportionof individualswithcomorbidpsychiatricdisordersand/orco-occurringdrug dependence.Severaleffortsreportnullfindings(Heathetal.,2011; Kapooretal.,2014; Lydalletal.,2011; Mbareketal.,2015; McGue etal.,2013; Zuoetal.,2012),butinabilitytoreplicatesomelocimaybe afunctionofbothcaseandcontrolascertainment(Franketal.,2012; Mailmanetal.,2007).MostrobustareassociationswithpotentiallyfunctionalSNPsthatalteralcoholmetabolism(Fig.1).Forexample,the ADH1B rs1229984Tallele(48His)resultsina40-to100-foldhigherrateofalcohol toacetaldehydemetabolism(i.e.,ethanoloxidation)(Edenberg,2000; Hurley,Bosron,Stone,&Amzel,1994).The ALDH2 rs671Aallele (504Lys)reduces ALDH2 activityandthusdecreasesacetaldehydetoacetate metabolism(i.e.,acetaldehydeoxidation)(Bosron&Li,1986; Enomoto, Takase,Yasuhara,&Takada,1991; Harada,Misawa,Agarwal,& Goedde,1980; Quillenetal.,2014).Acetaldehydeisatoxicsubstance whoseaccumulationleadstoahighlyaversivereactionthatincludesfacial flushing,nausea,andtachycardia.The ADH1B Hisand ALDH2 Lysvariants influencealcoholdrinkingbehaviorbyelevatingbloodacetaldehydelevels uponalcoholdrinkingwhichultimatelyreducessusceptibilitytodeveloping alcoholdrinkingproblems(Macgregoretal.,2009; Pengetal.,2010; Takeuchietal.,2011; Yokoyamaetal.,2008).

Table1 Genome-wideassociationstudiesofbeverageconsumptionreportingsignificantloci.

ReferencePhenotypePhenotypedefinedStudydesignNRace

Baik,Cho,Kim, Han,andShin (2011)

Alcohol drinking

Self-reportedalcoholconsumptionamongever drinkers

Questionnaire,alcoholg/dayinthepastmonth, derived

Population-basedmales

Takeuchietal. (2011)

AlcoholUse Disorder (NULL)

Alcohol drinking

AlcoholUseDisorderIdentificationTest (AUDIT)score: <8or 8score Replication 1113

Population-basedmales

Schumannetal. (2011)

Alcohol drinking

Self-reported“ever”vs“non-drinkers”alcohol consumption

Questionnaire,alcoholgou/wkalsoderived

Yangetal.(2013)

Alcohol drinking

Self-reportedalcoholconsumption

Self-reportedalcoholconsumptionamongever drinkers(sexstratified)

Questionnaire,alcoholg/dayperkg(nodetail)

Discovery733caJapanese

Population-based729co

Replication2794ca

Healthcenter-based1351co

Self-reported“drinkers”( 12drinksinpast year)vs“non-drinkers”alcoholconsumption In-personinterviews.

DiscoveryChinese

1population-based1420ca

1community-based3590co

Fordrinkers,alcoholg/dalsoderived(NULL)Replication4896ca

1population-based13,293co Continued

Table1 Genome-wideassociationstudiesofbeverageconsumptionreportingsignificantloci. cont’d

ReferencePhenotypePhenotypedefinedStudydesignNRace

Kapooretal. (2013) Alcohol drinking

Maximumnumberofdrinksconsumedamong drinkers

In-personinterview:“Whatisthelargest numberofdrinksyouhaveeverhadina24-h period”?(standardservingsizesperbeverage)

Meta-analysis

COGA:ADprobandsrecruited throughalcoholtreatmentprograms; relativesoftheprobandsand comparisonfamilies

SAGE:non-overlappingCOGA, FamilyStudyofCocaineDependence (FSCD),andCollaborativeGenetic StudyofNicotineDependence (COGEND)

4915EUR

Quillenetal. (2014) Alcohol drinking Maximumnumberofdrinksconsumedamong drinkers

In-personinterview,AD:DSM-IV

“Whatisthelargestnumberofalcoholicdrinks youhaveeverconsumedinaday?”and “Doesyourfaceflushafterdrinkingalittle alcohol?”

OneSample

MaleADprobandsandpedigreesfrom isolatedregion

272Chinese

Xuetal.(2015) Alcohol drinking

Maximumnumberofalcoholicbeverages consumedinanylifetime24-hperiod (MAXDRINKS)in-personinterview

Discovery

Yale-UPenn(recruitedforstudiesof geneticsofdrugorAD,familiesand unrelated) 2328EUR 3215AFR

Replication2736EUR

SAGE1276AFR

Adkinsetal.(2015) Alcohol drinking

Meanalcoholintakeoverdurationoffollow-up (i.e.transitionfromadolescentstoadulthood)

Meta-analysis

GreatSmokyMountainStudy(US, children)

Trajectoryestimateofalcoholconsumption (drinksperweek)acrossadolescenceandearly adulthood

ChristchurchHealthandDevelopment Study(CHDS):birthcohortchildren (NewZealand,1977)

2126EUR

In-personinterviews

Schumannetal. (2016) Alcohol drinking Alcoholg/damongdrinkers

Heavyvslight/nodrinking(NULL)

Self-reportedquestionnaires/diaries In-personinterviews

Jorgensonetal. (2017) Alcohol drinking

Individualswhoreporteddrinking 1dayper weekand 1drinkperdayweredefinedas “drinkers,”whereasthosewhoprovided negativeanswers(“nodays”and“none”)were consideredas“non-drinkers.”

Foralcoholdrinkers,theregularquantityof alcoholdrinksconsumedperweekwas calculatedbymultiplyingthetwoanswers

Self-reportedquestionnaire

VirginiaTwinStudyonAdolescent BehavioralDevelopment:twincohort US1974–1983

DiscoveryEUR

Meta-analysis70,460

Population-based74,711

Replication

Meta-analysis31,021

Population-based35,438

Meta-analysis

GeneticEpidemiologyResearchin

AdultHealthandAging(GERA)

cohort:membersoftheKaiser

PermanenteMedicalCarePlan, NorthernCaliforniaRegion(KPNC)

23,104coEUR

47,967caHisp/Latino EastAsian 2673coAA 4374ca 3288co 2746ca 1165co 1310ca

Continued