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P1:OTA/XYZ P2:ABC Color:1C BLBS098-fm BLBS098-Khatib November3,2011 19:17 Trim:244mmX172mm Livestock Epigenetics i P1:OTA/XYZ P2:ABC Color:1C BLBS098-fm BLBS098-Khatib November3,2011 19:17 Trim:244mmX172mm Livestock Epigenetics Edited by Hasan Khatib DepartmentofAnimalSciences UniversityofWisconsin Madison,Wisconsin,USA A John Wiley & Sons, Ltd., Publication iii P1:OTA/XYZ P2:ABC Color:1C BLBS098-fm BLBS098-Khatib November3,2011 19:17 Trim:244mmX172mm Thiseditionfirstpublished2012(cid:2)C 2012byJohnWiley&Sons,Inc. Wiley-BlackwellisanimprintofJohnWiley&Sons,formedbythemergerofWiley’sglobal Scientific,TechnicalandMedicalbusinesswithBlackwellPublishing. Registeredoffice: JohnWiley&Sons,Ltd,TheAtrium,SouthernGate,Chichester, WestSussex,PO198SQ,UK Editorialoffices: 2121StateAvenue,Ames,Iowa50014-8300,USA TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 9600GarsingtonRoad,Oxford,OX42DQ,UK Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowto applyforpermissiontoreusethecopyrightmaterialinthisbookpleaseseeourwebsiteat www.wiley.com/wiley-blackwell. Authorizationtophotocopyitemsforinternalorpersonaluse,ortheinternalorpersonaluseof specificclients,isgrantedbyBlackwellPublishing,providedthatthebasefeeispaiddirectlytothe CopyrightClearanceCenter,222RosewoodDrive,Danvers,MA01923.Forthoseorganizations thathavebeengrantedaphotocopylicensebyCCC,aseparatesystemofpaymentshasbeen arranged.ThefeecodesforusersoftheTransactionalReportingServiceareISBN-13: 978-0-4709-5859-9/2012. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.All brandnamesandproductnamesusedinthisbookaretradenames,servicemarks,trademarksor registeredtrademarksoftheirrespectiveowners.Thepublisherisnotassociatedwithanyproductor vendormentionedinthisbook.Thispublicationisdesignedtoprovideaccurateandauthoritative informationinregardtothesubjectmattercovered.Itissoldontheunderstandingthatthepublisher isnotengagedinrenderingprofessionalservices.Ifprofessionaladviceorotherexpertassistanceis required,theservicesofacompetentprofessionalshouldbesought. LibraryofCongressCataloging-in-PublicationData Livestockepigenetics/editor,HasanKhatib. p.cm. Includesbibliographicalreferencesandindex. ISBN978-0-470-95859-9(hardcover:alk.paper) 1.Livestock–Genetics. 2.Livestock–Embryology. 3.Epigenesis. I.Khatib,Hasan. SF756.5.L582012 636.089’6042–dc23 2011036446 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprint maynotbeavailableinelectronicbooks. Setin9.5/11ptTimesNewRomanbyAptara(cid:2)R Inc.,NewDelhi,India 1 2012 iv P1:OTA/XYZ P2:ABC Color:1C BLBS098-fm BLBS098-Khatib November3,2011 19:17 Trim:244mmX172mm Contents Contributors vii Preface xi 1. EpigeneticsofMammalianGameteandEmbryoDevelopment 3 NelidaRodriguez-Osorio,SuleDogan,andErdoganMemili 2. EpigeneticsofClonedPreimplantationEmbryosofDomesticAnimals 27 X.CindyTianandSadieL.Marjani 3. RolesofImprintedGenesinFertilityandPromisesoftheGenome-Wide Technologies 43 AshleyDriver,WenHuang,andHasanKhatib 4. SheepasanExperimentalModelforHumanART:NovelInsightsonPhenotypic AlterationsinART-DerivedSheepConceptuses 59 PasqualinoLoi,AntonellaD’Agostino,MartaCzernik,FedericaZacchini,PaolaToschi, AntonellaFidanza,andGrazynaPtak 5. TheDLK1-DIO3ImprintedGeneClusterandtheCallipygePhenotype inSheep 73 ChristopherA.Bidwell,RossL.Tellam,JolenaN.Waddell,TonyVuocolo, TracyS.Hadfield,andNoelleE.Cockett 6. GenomicImprintingandImprintedGeneClustersintheBovineGenome 89 IkhideG.Imumorin,SundayO.Peters,andMarcosDeDonato 7. ImprintinginGenomeAnalysis:ModelingParent-of-OriginEffectsin QTLStudies 113 SuzanneRowe,StephenBishop,andD.J.deKoning 8. EpigeneticsandAnimalHealth 131 JuanLuo,YingYu,andJiuzhouSong v P1:OTA/XYZ P2:ABC Color:1C BLBS098-fm BLBS098-Khatib November3,2011 19:17 Trim:244mmX172mm vi Contents 9. EpigeneticsandmicroRNAsinAnimalHealth 147 FeiTianandJiuzhouSong 10. NutrientsandEpigeneticsinBovineCells 161 Cong-junLi,RobertW.Li,andTedH.Elsasser Index 179 Colorplatesappearbetweenpages148and149. P1:OTA/XYZ P2:ABC Color:1C BLBS098-bcontrib BLBS098-Khatib October17,2011 15:8 Trim:244mmX172mm Contributors ChristopherA.Bidwell DepartmentofAnimalSciences PurdueUniversity Indiana,USA StephenBishop TheRoslinInstituteandR(D)SVS UniversityofEdinburghRoslinMidlothian Scotland,UK NoelleE.Cockett DepartmentofAnimal DairyandVeterinarySciences UtahStateUniversity Utah,USA MartaCzernik DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy AntonellaD’Agostino DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy SuleDogan DepartmentofAnimalandDairySciences MississippiStateUniversity Mississippi,USA vii P1:OTA/XYZ P2:ABC Color:1C BLBS098-bcontrib BLBS098-Khatib October17,2011 15:8 Trim:244mmX172mm viii Contributors MarcosDeDonato DepartmentofAnimalScience CornellUniversity NewYork,USAand LaboratorioGeneticaMolecular IIBCA UniversidaddeOriente Cumana,Venezuela AshleyDriver DepartmentofDairyScience UniversityofWisconsin-Madison Wisconsin,USA TedH.Elsasser BovineFunctionalGenomicsLaboratory Animal&NaturalResourcesInstitute,ARS,USDA Washington,DC AntonellaFidanza DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy TracyS.Hadfield DepartmentofAnimal DairyandVeterinarySciences UtahStateUniversity Utah,USA WenHuang DepartmentofDairyScience UniversityofWisconsin-Madison Wisconsin,USA IkhideG.Imumorin DepartmentofAnimalScience CornellUniversity NewYork,USA HasanKhatib DepartmentofAnimalSciences UniversityofWisconsin-Madison Wisconsin,USA D.J.deKoning DepartmentofAnimalBreedingandGenetics SwedishUniversityofAgriculturalSciences Uppsala,Sweden P1:OTA/XYZ P2:ABC Color:1C BLBS098-bcontrib BLBS098-Khatib October17,2011 15:8 Trim:244mmX172mm Contributors ix Cong-junLi BovineFunctionalGenomicsLaboratory Animal&NaturalResourcesInstitute,ARS,USDA Washington,DC RobertW.Li BovineFunctionalGenomicsLaboratory Animal&NaturalResourcesInstitute,ARS,USDA Washington,DC PasqualinoLoi DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy JuanLuo DepartmentofAnimalandAvianSciences UniversityofMaryland Maryland,USA SadieL.Marjani DepartmentofGenetics YaleUniversitySchoolofMedicine Connecticut,USA ErdoganMemili DepartmentofAnimalandDairySciences MississippiStateUniversity Mississippi,USA SundayO.Peters DepartmentofAnimalScience CornellUniversity NewYork,USA GrazynaPtak DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy NelidaRodriguez-Osorio GrupoCentauro UniversidaddeAntioquia Colombia SuzanneRowe TheRoslinInstituteandR(D)SVS UniversityofEdinburghRoslinMidlothian Scotland,UK P1:OTA/XYZ P2:ABC Color:1C BLBS098-bcontrib BLBS098-Khatib October17,2011 15:8 Trim:244mmX172mm x Contributors JiuzhouSong DepartmentofAnimalandAvianSciences UniversityofMaryland Maryland,USA RossL.Tellam CSIROLivestockIndustries St.Lucia,Queensland Australia X.CindyTian DepartmentofAnimalScience CenterforRegenerativeBiology UniversityofConnecticut Connecticut,USA FeiTian DepartmentofAnimalandAvianSciences UniversityofMaryland Maryland,USA PaolaToschi DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy TonyVuocolo CSIROLivestockIndustries St.Lucia,Queensland Australia JolenaN.Waddell DepartmentofAnimalSciences PurdueUniversity Indiana,USA YingYu DepartmentofAnimalandAvianSciences UniversityofMaryland Maryland,USA FedericaZacchini DepartmentofComparativeBiomedicalSciences UniversityofTeramo Teramo,Italy P1:OTA/XYZ P2:ABC Color:1C BLBS098-Preface BLBS098-Khatib November10,2011 10:55 Trim:244mmX172mm Preface Inrecentyears,wehavewitnessedremarkablepositivechangesintheperceptionofthevalue oflivestockgeneticandepigeneticresearchamongfederalandnonfederalagenciesandinthe scientificcommunitymoregenerally.First,itisbecomingwidelyacceptedthathumanhealth benefitsfromthepromotionofanimalhealth.Humanconsumptionofhealthymeat,eggs,dairy, andotheranimalproductsnecessitateshealthyanimals.Second,thewiderangeofphenotypic andgenotypicdiversityinlivestockpopulationsfacilitatestheuseofthesespeciesasmodels forcertainhumandiseasesandhealthtraits.Thereareplentifulhistoricandcontemporaryex- amplesofdomesticanimalsbeingusedinbiomedicalresearch,includingvaccinations(cowpox virus),xenotransplantations(heartvalvesofpigs),reproductivebiologyandcloning(Dolly,the firstclonedmammal),developmentalbiology(limbpatterninginchicken),metabolicdiseases (malignanthyperthermiainthepig),andneurodegenerativediseases(prionswerefirstisolated fromsheepandgoat)(http://www.adsbm.msu.edu/). At present, epigenetic studies are focused on understanding the mechanisms of actions of chromatinmodificationsthatregulatethemachineryofgeneexpression.AlthoughDNAmuta- tionsarecrucialinthedevelopmentofdiseasesandotherphenotypes,itisbecomingincreasingly evidentthatthesemutationsarenotsufficienttofullyexplaindiseaserisksandthat“additional” factorsareequallyimportant.Amajorelementofthe“additional”factorsisepigenetics,which playsakeyroleinphenotypicvariations,someofwhichappeartobetransgenerationallyinher- itedinalmostallorganisms.Theaccelerationofepigeneticresearchoverthelastfewyearsis impressive.Indeed,itisnoteasytofindabiologicalquestionthatdoesnotengageanepigenetic component. Thereisampleandaccumulatingevidenceoftheinfluenceofenvironment,includingdiet, ongeneactionsthroughthemanipulationoftheepigenome.Currently,thebestknownexample ofanepigeneticallysensitivegeneaffectedbyanenvironmentalfactorisAgoutiviableyellow and the maternal diet in the mouse. Interestingly, it was found that methyl supplements in the diet of pregnant mice increased the methylation level of the Agouti gene, consequently causingcoatcolorchangesintheoffspring(Cooneyetal.,2002;Waterlandetal.,2008).Further studies in rats showed that high-fat diet consumption by sires induced increased body and bodyfatweightsandhigherglucosetoleranceandinsulinresistanceintheirfemaleoffspring (Ngetal.,2010).AnotherfascinatingexampleofnutritionaleffectsonDNAmethylationand xi

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