The Comparative Biology of Aging Norman S. Wolf Editor The Comparative Biology of Aging 1 3 Editor NormanS.Wolf SeattleWA98195-7470 C-423HealthSciencesBldg. USA [email protected] Coverphotocredits: Human:45yearoldWerner’sSyndromepatient,InternationalRegistryofWernerSyndrome, courtesy of Dru Leistritz.Rhesus monkey: 29 year old male (M. mulatta), NIA Study on Primate Aging, courtesy of Dr. Julie Mattison, primate aging colony, NIA. Dog: “Tiger”, 13yearoldWestHighlandWhiteterrier,courtesyofDanandPatMcCutcheonandJeremy andMicheleWolf.Mouse,BalbCstrain:CourtesyofDr.NormWolflaboratory,Department of Pathology, University of Washington. Fruit fly (D. melanogaster): Courtesy of Dr. Rolf Bodmer,DevelopmentandAging,BurnhamInstitute,LaJolla,Ca.Roundworm(C.elegans): CourtesyofDrs.MattKaeberleinandBrianKennedy,DepartmentofPathology,University ofWashington. ISBN978-90-481-3464-9 e-ISBN978-90-481-3465-6 DOI10.1007/978-90-481-3465-6 SpringerDordrechtHeidelbergLondonNewYork LibraryofCongressControlNumber:2009942696 ©SpringerScience+BusinessMediaB.V.2010 Nopartofthisworkmaybereproduced,storedinaretrievalsystem,ortransmittedinanyformorby anymeans,electronic,mechanical,photocopying,microfilming,recordingorotherwise,withoutwritten permissionfromthePublisher,withtheexceptionofanymaterialsuppliedspecificallyforthepurpose ofbeingenteredandexecutedonacomputersystem,forexclusiveusebythepurchaserofthework. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Contents Introduction: Lifespans and Pathologies Present at Death inLaboratoryAnimals . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 NormanS.WolfandStevenAustad AnimalSize,MetabolicRate,andSurvival,AmongandWithinSpecies 27 StevenN.Austad HormonalInfluencesonAgingandLifespan . . . . . . . . . . . . . . . 43 AdamSpongandAndrzejBartke Exploring Mechanisms of Aging Retardation by Caloric Restriction:StudiesinModelOrganismsandMammals . . . . . . . . . 69 RozalynM.Anderson,RickiJ.Colman,andRichardWeindruch CellReplicationRatesInVivoandInVitroandWoundHealing asAffectedbyAnimalAge,Diet,andSpecies . . . . . . . . . . . . . . . 97 NormanS.Wolf SirtuinFunctioninLongevity . . . . . . . . . . . . . . . . . . . . . . . . 123 DanielL.SmithJr.andJeffreyS.Smith TheRoleofTORSignalinginAging . . . . . . . . . . . . . . . . . . . . 147 MattKaeberleinandLaraS.Shamieh Mitochondria,OxidativeDamageandLongevity:WhatCan ComparativeBiologyTeachUs? . . . . . . . . . . . . . . . . . . . . . . 163 YunShi,RochelleBuffenstein,andHollyVanRemmen ComparativeGenomicsofAging . . . . . . . . . . . . . . . . . . . . . . 191 JanVijg,AnaMariaGarcia,BrentCalder,andMartijnDollé Changes in Lysosomes and Their Autophagic Function in Aging:TheComparativeBiologyofLysosomalFunction . . . . . . . . 201 SamanthaJ.OrensteinandAnaMariaCuervo TelomeresandTelomerase . . . . . . . . . . . . . . . . . . . . . . . . . 227 N.M.V.Gomes,J.W.Shay,andW.E.Wright v vi Contents CardiacAging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259 Dao-FuDai,RobertJ.Wessells,RolfBodmer,and PeterS.Rabinovitch ComparativeSkeletalMuscleAging . . . . . . . . . . . . . . . . . . . . 287 DavidJ.Marcinek,JonathanWanagat,andJasonJ.Villarin AgingoftheNervousSystem . . . . . . . . . . . . . . . . . . . . . . . . 319 CatherineA.Wolkow,SigeZou,andMarkP.Mattson AgingoftheImmuneSystemAcrossDifferentSpecies . . . . . . . . . . 353 JankoNikolich-ŽugichandLukaCˇicˇin-Šain Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Contributors RozalynAnderson WisconsinNationalPrimateResearchCenter,Universityof Wisconsin-Madison,Madison,WI,USA,[email protected] StevenN.Austad DepartmentofCellular&StructuralBiology,BarshopInstitute forLongevityandAgingResearch,UniversityofTexasHealthScienceCenterat SanAntonio,SanAntonio,TX,USA,[email protected] AndrzejBartke GeriatricResearch,DepartmentofInternalMedicine,Southern IllinoisUniversitySchoolofMedicine,Springfield,IL62794,USA;Departmentof Physiology,SouthernIllinoisUniversitySchoolofMedicine,Springfield,IL 62794,USA,[email protected] RolfBodmer BurnhamInstitute,Developmentandagingprogram,LaJolla, CA,USA RochelleBuffenstein DepartmentofPhysiology,CellularandStructuralBiology, BarshopInstituteforLongevityandAgingStudies,UniversityofTexasHealth ScienceCenteratSanAntonio,SanAntonio,TX,USA BrentCalder DepartmentofGenetics,AlbertEinsteinCollegeofMedicine, Bronx,NY10461,USA LukaCicin-Sain VaccineandGeneTherapyInstituteandtheOregonNational PrimateResearchCenter,OregonHealthandScienceUniversity,Beaverton,OR 97006,USA RickiColman WisconsinNationalPrimateResearchCenter,Universityof Wisconsin-Madison,Madison,WI,USA;DepartmentofMedicine,Universityof Wisconsin-Madison,Madison,WI,USA AnaMariaCuervo DepartmentofDevelopmentalandMolecularBiology, MarionBessinLiverResearchCenter,InstituteforAgingResearch,Albert EinsteinCollegeofMedicine,Bronx,NY,USA,[email protected] Dao-FuDai DepartmentofPathology,UniversityofWashington,Seattle, WA,USA vii viii Contributors MartijnDollé NationalInstituteofPublicHealthandtheEnvironment,Bilthoven, TheNetherlands AnaMariaGarcia UniversityofTexasatSanAntonio,SanAntonio,TX,USA N.M.V.Gomes DepartmentofCellBiology,TheUniversityofTexas SouthwesternMedicalCenteratDallas,Dallas,TX75390-9039,USA YujiIkeno BarshopInstituteandUniversityofTexasScienceCenteratSan Antonio,SanAntonio,TX,USA MattKaeberlein DepartmentofPathology,UniversityofWashington,Seattle, WA,USA,[email protected] MarkP.Mattson LaboratoryofNeurosciences,NationalInstituteonAging IntramuralResearchProgram,Baltimore,MD,USA,[email protected] DavidJ.Marcinek DepartmentofRadiology,UniversityofWashington,Seattle, WA98195,USA,[email protected] JulieMattison LaboratoryofExperimentalGerontology,NIA,Baltimore, MD,USA JankoNikolich-Zugich DepartmentofImmunobiologyandtheArizonaCenter onAging,UniversityofArizonaCollegeofMedicine,Tucson,AZ85718,USA; VaccineandGeneTherapyInstituteandtheOregonNationalPrimateResearch Center,OregonHealthandScienceUniversity,Beaverton,OR97006,USA, [email protected] SamanthaJ.Orenstein DepartmentofDevelopmentalandMolecularBiology, MarionBessinLiverResearchCenter,InstituteforAgingResearch,Albert EinsteinCollegeofMedicine,Bronx,NY,USA BeverlyPaigen JAXLaboratories,BarHarbor,ME,USA PeterS.Rabinovitch DepartmentofPathology,UniversityofWashington, Seattle,WA,USA,[email protected] LaraS.Shamieh DepartmentofPathology,UniversityofWashington,Seattle, WA,USA J.W.Shay DepartmentofCellBiology,TheUniversityofTexasSouthwestern MedicalCenteratDallas,Dallas,TX75390-9039,USA YunShi DepartmentofPhysiology,BarshopInstituteforLongevityandAging Studies,UniversityofTexasHealthScienceCenteratSanAntonio,SanAntonio, TX,USA DanielL.Smith DepartmentofBiochemistryandMolecularGenetics,Schoolof Medicine,UniversityofVirginiaHealthSystem,Charlottesville,VA22908,USA Contributors ix JeffreyS.Smith DepartmentofBiochemistryandMolecularGenetics,Schoolof Medicine,UniversityofVirginiaHealthSystem,Charlottesville,VA22908,USA, [email protected] AdamSpong GeriatricResearch,DepartmentofInternalMedicine,Southern IllinoisUniversitySchoolofMedicine,Springfield,IL62794,USA HollyVanRemmen DepartmentofPhysiology,CellularandStructuralBiology, BarshopInstituteforLongevityandAgingStudies,UniversityofTexasHealth ScienceCenteratSanAntonio;AudieMurphyDivision,SouthTexasVeterans HealthCareSystem,SanAntonio,TX,USA,[email protected] JanVijg DepartmentofGenetics,AlbertEinsteinCollegeofMedicine,Bronx, NY10461,USA,[email protected] JasonJ.Villarin DepartmentofRadiology,UniversityofWashington,Seattle, WA98195,USA JonathanWanagat DivisionofGerontologyandGeriatricMedicine,University ofWashington,Seattle,WA98195,USA RichardWeindruch WisconsinNationalPrimateResearchCenter,Universityof Wisconsin-Madison,Madison,WI,USA;DepartmentofMedicine,Universityof Wisconsin-Madison,Madison,WI,USA;GRECCVAHospitalMadison, UniversityofWisconsin-Madison,Madison,WI,USA RobertJ.Wessells DepartmentofInternalMedicine,UniversityofMichigan, AnnArbor,MI,USA NormanS.Wolf DepartmentofPathology,UniversityofWashington,Seattle, WA91895,USA,[email protected] CatherineA.Wolkow LaboratoryofNeurosciences,NationalInstituteonAging IntramuralResearchProgram,Baltimore,MD,USA,[email protected] W.E.Wright DepartmentofCellBiology,TheUniversityofTexasSouthwestern MedicalCenteratDallas,Dallas,TX75390-9039,USA, [email protected] RongYuan JAXLaboratories,BarHarbor,ME,USA SigeZou LaboratoryofExperimentalGerontology,NationalInstituteonAging IntramuralResearchProgram,Baltimore,MD,USA Introduction: Lifespans and Pathologies Present at Death in Laboratory Animals NormanS.Wolf1andStevenAustad2 1DepartmentofPathology,UniversityofWashington,Seattle,WA 2BarshopInstituteandUniversityofTexasScienceCenteratSanAntonio, SanAntonio,TX Abstract This initial chapter introduces those that follow with a summary of the lifespansandendoflifepathologiesoftheseveralspeciesthatareincludedinlater chapters. It is not presented as a complete coverage, but rather as background for what follows, as gathered from the literature and personal information. Included here is that information on yeast as Saccharomyces cervisiae, round worms as Caenorhabditis. elegans, the fruit fly, Drosophila melanogaster, mice, rats, dogs, primates,andsomebatsandbirds.Thefollowingchapterswillcomparefindingsin theirspecificareaofcoverageforthosespeciesforwhichsuchdataisavailable. Thisbriefintroductionprovidesgenerallyacceptedspecieswildtypelifespansand theendoflifepathologies,alongwithsomespecialattributesthatlendaspeciesto acomparativebiologyapproach. Keywords Aging·Lifespan·Endoflifeconditions·Humans·Dogs·Mice·Rates· Fish·Amphibians·Causeofdeath LaboratoryStrainsofYeast(Saccharomycescervisiae) WildTypeOnly ThelifespanofS.cervisiaevarieswithbothstrainandmediaused(especiallyglu- cosecontent).Thelifespanofyeastmaybedeterminedbytwodifferentmeans:the numberofdivisionsaccomplishedbythemothercell(replicativelifespan),orbythe survival of individual cells in a non-dividing, quiescence-like state (chronological lifespan).Replicativelifespanisdeterminedbyphysicalseparationofdaughtercells from mother cells via micromanipulation, while tallying the number of daughter cellsproducedbyeachmother[1].Chronologicallifespanistypicallydetermined Data Contributors: Julie Mattison, Laboratory of Experimental Gerontology, NIA; Mathew Kaeberlein,DepartmentofPathology,UniversityofWashington,Seattle,WA;BeverlyPaigenand RongYuan,JAXLaboratories,BarHarbor,ME;YujiIkeno,BarshopInstituteandUniversityof TexasScienceCenteratSanAntonio,TX;RochelleBuffenstein,DepartmentofPhysiologyand BarshopInstitute,UniversityofTexasScienceCenteratSanAntonioTX N.S.Wolf(ed.),TheComparativeBiologyofAging, 1 DOI10.1007/978-90-481-3465-6_1,(cid:2)C SpringerScience+BusinessMediaB.V.2010