Handbook of Neurochemistry and Molecular Neurobiology BehavioralNeurochemistry,NeuroendocrinologyandMolecularNeurobiology Editor-in-Chief AbelLajtha Director CenterforNeurochemistry NathanS.KlineInstituteforPsychiatricResearch 140OldOrangeburgRoad Orangeburg NewYork,10962 USA VolumeEditor JeffreyD.Blaustein CenterforNeuroendocrineStudies 135HicksWay UniversityofMassachusetts Amherst,MA01003‐9271 USA LibraryofCongressControlNumber:2006922553 ISBN13:978‐0‐387‐30362‐8 Additionally,thewholesetwillbeavailableuponcompletionunderISBN‐13:978‐0‐387‐35443‐9 TheelectronicversionofthewholesetwillbeavailableunderISBN‐13:978‐0‐387‐30426‐7 TheprintandelectronicbundleofthewholesetwillbeavailableunderISBN‐13:978‐0‐387‐35478‐1 (cid:1)2007SpringerScienceþBusinessMedia,LLC. Allrightsreserved.Thisworkmaynotbetranslatedorcopiedinwholeorinpartwithoutthewrittenpermission ofthepublisher(SpringerScienceþBusinessMedia,LLC.,233SpringStreet,NewYork,NY10013,USA),exceptfor briefexcerptsinconnectionwithreviewsorscholarlyanalysis.Useinconnectionwithanyformofinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodologynow knownorhereafterdevelopedisforbidden. Theuseinthispublicationoftradenames,trademarks,servicemarks,andsimilarterms,eveniftheyarenot identifiedassuch,isnottobetakenasanexpressionofopinionastowhetherornottheyaresubjectto proprietaryrights. springer.com Printedonacid‐freepaper SPIN:114177672109–543210 Table of Contents Preface .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. .. v Contributors.. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. . xi 1 NeuroendocrinologyofMaleReproductiveBehavior. .. .. .. . .. .. .. . .. .. .. 1 M.J.Baum 2 NeurochemistryofMaleSexualBehavior . .. .. .. . .. .. .. .. . .. .. .. . .. .. . 37 E.M.Hull.J.M.Dominguez.J.W.Muschamp 3 FeminineSexualBehaviorfromNeuroendocrineandMolecular NeurobiologicalPerspectives .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. . 95 J.D.Blaustein.S.K.Mani 4 TheNeurochemistryofLimbic‐HypothalamicCircuitsRegulatingSexual Receptivity. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 151 PaulMicevych.KevinSinchak 5 NeuroendocrinologyandNeurochemistryofMaternalMotivationand Behavior .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 195 J.S.Lonstein.J.I.Morrell 6 Neuroendocrinology,Neurochemistry,andMolecularNeurobiologyof AffiliativeBehavior . .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 247 E.A.D.Hammock.L.J.Young 7 NeurochemistryandMolecularNeurobiologyofAggressiveBehavior . .. .. 285 K.A.Miczek.S.P.Faccidomo.E.W.Fish.J.F.DeBold 8 NovelMechanismsUnderlyingNeuroendocrineRegulationofAggression: ASynthesisofRodent,Avian,andPrimateStudies .. .. .. .. . .. .. .. . .. .. 337 G.E.Demas.M.A.Cooper.H.E.Albers.K.K.Soma 9 MolecularNeurobiologyofBirdSong. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 373 D.F.Clayton #Springer-VerlagBerlinHeidelberg2007 viii TableofContents 10 TheNeuroendocrinologyandNeurochemistryofBirdsong .. . .. .. .. . .. .. 419 G.F.Ball.J.Balthazart 11 SexDifferencesinNeurotransmittersandBehavior:Development .. . .. .. 459 M.M.McCarthy,PhD 12 SexDifferencesinNeurotransmittersSystems;Vasopressinas anExample .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 487 G.J.deVries 13 NeurochemicalSystemsRegulatingtheHypothalamo–Pituitary– AdrenocorticalAxis . .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 513 J.P.Herman.H.F.Figueiredo.N.K.Mueller.M.M.Ostrander.R.Zhang. M.Tauchi.D.C.Choi.A.R.Furay.N.K.Evanson.E.B.Nelson. Y.M.Ulrich‐Lai 14 NeuroendocrinologyofStress.. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 571 B.S.McEwen.S.Chattarji 15 MaternalProgrammingofGlucocorticoidReceptorExpressionandHPA ResponsestoStressThroughDNAMethylationintheRat. .. . .. .. .. . .. .. 595 M.J.Meaney.I.C.G.Weaver.T.Wu.I.Hellstrom.J.Diorio.MosheSzyf 16 EnergyBalanceandFeeding . .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 619 S.C.Benoit.D.J.Clegg.S.C.Woods 17 TheNeuroendocrinology,NeurochemistryandMolecularBiologyof ThirstandSaltAppetite .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 641 A.K.Johnson.R.L.Thunhorst 18 Neurochemistry/NeuropharmacologyofFearandFearConditioning . . .. .. 689 G.E.Schafe.J.E.LeDoux 19 NeurochemistryandMolecularNeurobiologyofMemory . .. . .. .. .. . .. .. 709 P.Dash.A.N.Moore 20 NeurochemistryandMolecularNeurobiologyofReward .. .. . .. .. .. . .. .. 739 J.B.Becker.R.L.Meisel 21 NeuroendocrinologyofMemoryandCognitiveFunction.. .. . .. .. .. . .. .. 775 V.N.Luine 22 TheMammalianCircadianSystem:fromGenestoBehavior . . .. .. .. . .. .. 801 B.H.Miller.E.L.McDearmon.J.S.Takahashi 23 NeuroendocrinologyofBehavioralRhythms .. .. . .. .. .. .. . .. .. .. . .. .. 835 T.M.Lee.L.Smale TableofContents ix 24 NeurochemistryofSleep .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 869 T.Porkka‐Heiskanen.L.Alanko.D.Stenberg 25 NeuroendocrinologyofSleep .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 895 A.Steiger Index . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. . .. .. .. .. . .. .. .. . .. .. 939 Preface Behavioral neuroscience was not covered extensively in the second edition of the Handbook of Neurochemistry, published in 1983. That was nearly a decade before the formation of the international society,whichnameditselfafterthisdiscipline,theInternationalSocietyforBehavioralNeuroscience,and itwasevenlongerbeforetheinceptionoftheSocietyforBehavioralNeuroendocrinology,whichfocuseson a subfield of behavioral neuroscience. The progress that has been made in the study of the cellular and molecular underpinnings of behavior was almost unimaginable in 1983. The field has prosperedthanks to development in novel drugs, genetic models, and related molecular techniques, neuroanatomical techniques, including in situ hybridization histochemistry, new immunocytochemical techniques, real timePCR,microarrays,andmoresophisticatedbehavioralanalysis. This volume is filled with a tremendous amount of history that documents the coming of age of behavioral neuroscience. Learning the history and following the development of a field are often an importantpartofunderstandinganareaofscience,andmanyoftheauthorshaveelaboratedextensively onthehistoryoftheirfield.Thoughbehavioralneurosciencehasadvancedtremendouslyinrecentyears, impedimentstoprogressstillremaininthisfield.Forexample,behaviorstilloccasionallytakesabackseat tothestudyofsimplerphysiologicalendpoints,suchasthecontrolofovulation.Yet,itisthemorecomplex regulationofbehaviorandtheinteractionsoftheenvironmentonitthatallowforfertilization,without which,ovulationwouldbeirrelevant.Inhismemorablebiographyofthefieldofhormonesandbehavior, FrankBeach(1981)explainedsomeofthebackgroundforbiasagainststudiesofbehavior.Healsoprovided whatisperhapsoneofthemostnoteworthyexamplesofthisbiasagainststudyingbehavioralendpoints.In 1935,EdwardDempsey,workinginWilliamC.Young’sgroup,madethehereticproposalthatsequential exposure to estradiol and progesterone was necessary to induce the expression of estrous behaviors in female guinea pigs. This made no sense at the time since it was widely believed that the source of progesteronewasthecorpusluteum,mostdefinitelyformedafterovulation,longaftertheanimal’sestrous behavior had commenced. Beach recounts that Edgar Allen, a co‐discoverer of estradiol, suggested that Young would be ‘‘well‐advised to give up behavior and return to his more promising early studies on physiologyoftheepididymis.’’Itwastobeanother30yearsbeforenovelbiochemicalprocedureswouldbe developed, whichwouldproveDempsey, Young and colleagues correct; progesterone was being secreted fromanothersourcebeforetheformationofthecorpusluteum.Behavioralstudieshadindeedinformed physiology. Throughout this volume, youwill find examples of dogmas that ultimately did not hold water (the timingofprogesteronesecretionduringtheestrouscyclejustdiscussedisbutoneexample).Timeandtime again, throughout the history of science, scientists who have questioned dogma have been subjected to ridiculeorderisionfortheiractions.Winorlose,thedogmafightisalwaysworthwaging,anditisgood science.Ifthedogmarepresentstruth,itwillstand;ifnot,itwilleventuallytopple,butusuallynotwithout significantbattle.Idedicatethisvolumetoallscientistswhohaveatonepointoranotherchallengeddogma intheirwork. Inthisvolume,Ihavecollectedinoneplacetheexpertiseofnumerousauthoritiesinthediversefieldof behavioralneuroscience. Avolumeofthis sizedoes notallowfor anexhaustivetreatmentoftheneuro- chemistry,neuroendocrinology,andmolecularneurobiologyofbehavior;ratheritisasamplingofsome fascinatingareaswithintherealmofbehavioralneuroscience.Moreover,becauseofmypersonalbias,these vi Preface areasofbehavioralneuroscienceoftenhaveanimportant,well‐developed,endocrineslant.Toappreciate howmuchspaceacomprehensivetreatmentoftheentirefieldwouldrequire,considerthatacomprehen- sive coverage of the subfield, the relatively narrow field of behavioral neuroendocrinology, was recently accomplished admirably in a discipline‐defining, five‐volume, and nearly 4,000 page work, which was editedbyDonaldW.Pfaffetal. Itisimpossibletoacknowledgehereeachoftheauthorsandalloftheimportantfindingsofthefields thattheyrepresent.Sufficeittosaythattremendousprogresshasbeenmadeinthestudyofreproductive behaviors,affiliativeandaggressivebehaviors,birdsong,sexdifferences,thehypothalamo‐pituitary‐adrenal axis,stress,ingestivebehaviors,fear,cognitivefunction,reward,rhythms,andsleep.Thebookstartsatthe beginningwithreproductionandendsappropriatelywithsleep.Betweenthosetwobasic,life‐generating andrestorativeactivities,tremendousprogressinallofthesefieldsisdescribed. Recent changes in the funding climate have affected many areas within behavioral sciences. For example,theNationalInstitutesofHealthhaveshiftedemphasistomoretranslationalresearch;insome cases,attheexpenseofmorebasicresearch.Thisvolumeisfilledwithexamplesofbasicresearchthathave ledtoabetterunderstandingofthehumanbrainandbehavior.Ihopethatitservesastestamenttothe indispensablevalueofbasicresearch,aswellastranslationalresearch,inbehavioralneuroscience. FrankBeachendedhisbiographyofthefieldofbehavioralendocrinologywiththefollowingpassage: ‘‘Scientistswithdoctoratesinpsychologystudydevelopmentofprogesteronereceptorsinneuronsoftherat hypothalamuswhileotherinvestigatorsinitially trainedinpharmacology inventelegantbehavioralmea- suresofsexualmotivationintheestrousfemale.Thesedevelopmentsappeartorepresentmorethanamere borrowing of techniques by one discipline from another. Instead they seem to reflect progress toward recognition of common goals and shared theoretical interests. If such indeed is the case, behavioral endocrinologymaywellbeadisciplineinstatunascendi,’’thatis,adisciplineinastateofbeingborn.He could have said the same about behavioral neuroscience. I submit that the discipline of behavioral neuroscience has become a fully developed discipline with investigators answering questions that truly runthegamutfrommoleculartobehavioralandalllevelsinbetween. Iamimmenselygratefultoallwhocontributedtothisvolume.Writingacomprehensivereviewofa field,evenone’sspecialty,istime‐consumingandlaborious,anditinvariablytakestimeawayfromother worthytasks.Myjobofconvincingcolleaguestocontributetothisvolumewasmaderelativelyeasybecause previousversionsoftheHandbookofNeurochemistryhavebeenwellreceived,andhaveoftenservedas landmarkvolumesintheirrespectivefields.Itismyhopethattheauthorswillbewellcompensatedfortheir workwiththesatisfactionofknowingthattheirreviewswillbereadandthat,astheirareasevolve,progress canbeupdatedandfollowedintheelectronicversionoftheHandbook. JeffreyD.Blaustein Amherst,USA April2006 Contributors L.Alanko D.Choi InstituteofBiomedicine/Physiology,Universityof DepartmentofPsychiatry,UniversityofCincinnati, Helsinki,Helsinki,Finland Cincinnati,OH45237‐0506,USA H.E.Albers D.F.Clayton DepartmentsofBiologyandPsychologyandCenter SchoolofMolecularandCellularBiology, forBehavioralNeuroscience,GeorgiaStateUniversity, BeckmanInstitute,andInstituteforGenomic Atlanta,GA30303,USA Biology,UniversityofIllinois,Urbana‐Champaign, IL61801,USA G.F.Ball DepartmentofPsychologicalandBrainSciences, D.J.Clegg JohnsHopkinsUniversity,3400N.CharlesStreet, DepartmentofPsychiatry,UniversityofCincinnati,USA Baltimore,MD21218‐2686,USA M.A.Cooper DepartmentofPsychologyandCenterforBehavioral J.Balthazart Neuroscience,GeorgiaStateUniversity,Atlanta, CenterforCellularandMolecularNeurobiology, GA30303,USA ResearchGroupinBehavioralNeuroendocrinology, UniversityofLie`ge,1Boulevarddel’Hopital(Bat.L36), B‐4000Lie`ge1,Belgium P.Dash TheVivianL.SmithCenterforNeurologicResearch DepartmentofNeurobiologyandAnatomyThe M.J.Baum UniversityofTexasMedicalSchoolatHouston, Dept.ofBiologyBostonUniversityBoston, Houston,Texas77030,USA MA02215,USA G.J.deVries J.B.Becker CenterforNeuroendocrineStudiesandDepartment DepartmentofPsychology.NeuroscienceProgramand ofPsychologyUniversityofMassachusetts, ReproductiveSciencesProgram,UniversityofMichigan, Amherst,MA01003,USA 525EastUniversityAve.,AnnArbor,MI48109,USA J.F.DeBold S.C.Benoit DepartmentofPsychology,TuftsUniversity,Medford DepartmentofPsychiatry,UniversityofCincinnati,USA andBoston,Massachusetts,USA J.D.Blaustein G.E.Demas CenterforNeuroendocrineStudiesandPsychology DepartmentofBiology,PrograminNeuralScience Department,UniversityofMassachusetts, andCenterfortheIntegrativeStudyofAnimal Amherst,MA,USA Behavior,IndianaUniversity,Bloomington, IN47405,USA S.Chattarji NationalCentreforBiologicalSciences,TataInstitute J.Diorio ofFundamentalResearch,Bangalore560065,India DouglasHospitalResearchCentre,Montreal,Canada #Springer-VerlagBerlinHeidelberg2007 xii Contributors J.M.Dominguez T.M.Lee DepartmentofPsychologyFloridaStateUniversity DepartmentofPsychologyandNeuroscience Tallahassee,FL32306‐1270,USA Program,UniversityofMichigan,AnnArbor, 48109‐1043,USA N.K.Evanson DepartmentofPsychiatry,UniversityofCincinnati, J.S.Lonstein Cincinnati,OH45237‐0506,USA PrograminNeuroscienceandDepartmentof Psychology,GiltnerHall,MichiganStateUniversity, S.P.Faccidomo EastLansing,MI48824,USA DepartmentofPsychology,TuftsUniversity,Medford andBoston,Massachusetts,USA V.N.Luine DepartmentofPsychology,HunterCollegeofC.U.N.Y., H.F.Figueiredo 695ParkAve.,NewYork,NY10021,USA DepartmentofPsychiatry,UniversityofCincinnati, Cincinnati,OH45237‐0506,USA S.K.Mani DepartmentofMolecularandCellularBiology,Baylor CollegeofMedicine,Houston,TX,USA E.W.Fish DepartmentofPsychology,TuftsUniversity,Medford M.M.McCarthy,PhD andBoston,Massachusetts,USA DepartmentsofPhysiologyandPsychiatry,University ofMaryland,Baltimore–SchoolofMedicine.655 A.Furay W.BaltimoreSt.,Baltimore,MD21201,USA DepartmentofPsychiatry,UniversityofCincinnati, Cincinnati,OH45237‐0506,USA E.L.McDearmon HowardHughesMedicalInstitute E.A.D.Hammock DepartmentofPsychiatryandBehavioralSciences, B.S.McEwen EmoryUniversity,Atlanta,GA30329.Centerfor HaroldandMargaretMillikenHatchLaboratory BehavioralNeuroscience,Atlanta,GA30329. ofNeuroendocrinology,TheRockefellerUniversity, YerkesNationalPrimateResearchCenter,Atlanta, NewYork,NY10021,USA GA30329,USA M.J.Meaney I.Hellstrom McGillProgramfortheStudyofBehavior,Genesand McGillProgramfortheStudyofBehavior,Genesand Environment,McGillUniversity,MontrealCanada. Environment,McGillUniversity,Montreal,Canada DouglasHospitalResearchCentre,6875boul.LaSalle, Montreal,Que´bec,CanadaH4H1R3 J.P.Herman DepartmentsofPsychiatryandCellBiology, R.L.Meisel NeurobiologyandAnatomy,UniversityofCincinnati, DepartmentofPsychologicalSciencesand Cincinnati,OH45237‐0506,USA NeuroscienceProgram,PurdueUniversity,703ThirdSt., WestLafayette,IN47907,USA E.M.Hull DepartmentofPsychologyFloridaStateUniversity P.Micevych Tallahassee,FL32306‐1270,USA DeptofNeurobiology,Laboratoryof NeuroendocrinologyoftheBrainResearchInstitute, A.K.Johnson DavidGeffenSchoolofMedicineatUCLA, DepartmentsofPsychology,Pharmacologyand LosAngeles,CA90095‐1763,USA ExerciseScience,andtheCardiovascularCenter, UniversityofIowa,IowaCity,IA52242,USA K.A.Miczek DepartmentofPsychology,Psychiatry,Pharmacology, J.E.LeDoux andNeuroscienceTuftsUniversityMedfordandBoston, CenterforNeuralScience,NewYorkUniversity,USA Massachusetts,USA Contributors xiii B.H.Miller A.Steiger TheScrippsResearchInstitute,Jupiter,FL33458,USA MaxPlanckInstituteofPsychiatry,Munich, Germany A.N.Moore TheVivianL.SmithCenterforNeurologicResearch D.Stenberg DepartmentofNeurobiologyandAnatomyThe InstituteofBiomedicine/Physiology,University UniversityofTexasMedicalSchoolatHouston, ofHelsinki,Helsinki,Finland Houston,Texas77030,USA M.Szyf J.I.Morrell CenterforMolecularandBehavioralNeuroscience, McGillProgramfortheStudyofBehavior,Genes andEnvironment,McGillUniversity,Montreal, 197UniversityAve,RutgersUniversity,Newark, Canada.DepartmentofPharmacology, NJ07102,USA McGillUniversity,MontrealCanada N.K.Mueller DepartmentofPsychiatry,UniversityofCincinnati, J.S.Takahashi Cincinnati,OH45237‐0506,USA HowardHughesMedicalInstitute,Departmentof Neurobiology&Physiology,NorthwesternUniversity, J.W.Muschamp Evanston,IL60208,USA DepartmentofPsychology,FloridaStateUniversity Tallahassee,FL32306‐1270,USA M.Tauchi DepartmentofPsychiatry,UniversityofCincinnati, E.B.Nelson Cincinnati,OH45237‐0506,USA DepartmentofPsychiatry,UniversityofCincinnati, Cincinnati,OH45237‐0506,USA R.L.Thunhorst M.M.Ostrander DepartmentsofPsychology,Pharmacologyand DepartmentofPsychiatry,UniversityofCincinnati, ExerciseScience,andtheCardiovascularCenter, Cincinnati,OH45237‐0506,USA UniversityofIowa,IowaCity,IA52242,USA T.Porkka‐Heiskanen Y.M.Ulrich‐Lai InstituteofBiomedicine/Physiology,Universityof DepartmentofPsychiatry,UniversityofCincinnati, Helsinki,Helsinki,Finland Cincinnati,OH45237‐0506,USA G.E.Schafe I.C.G.Weaver DepartmentofPsychology&Interdisciplinary McGillProgramfortheStudyofBehavior,Genes NeuroscienceProgram,YaleUniversity.Department andEnvironment,McGillUniversity,Montreal, ofPsychology,YaleUniversity,2HillhouseAvenue, Canada.DouglasHospitalResearchCentre, NewHaven,CT06520,USA Montreal,Canada.DepartmentofPharmacology, K.Sinchak McGillUniversity,Montreal,Canada DeptofPsychology,CenterforNeuroendocrine StudiesandNeuroscienceandBehaviorProgram, S.C.Woods UniversityofMassachusetts,Amherst,MA,USA DepartmentofPsychiatry,Universityof Cincinnati,USA L.Smale DepartmentofPsychology,MichiganStateUniversity, T.Wu E.Lansing,MI48824,USA McGillProgramfortheStudyofBehavior,Genes andEnvironment,McGillUniversity,Montreal K.K.Soma Canada.DouglasHospitalResearchCentre, DepartmentsofPsychologyandZoology,Graduate Montreal,Canada.DepartmentofPharmacology, PrograminNeuroscience,UniversityofBritish McGillUniversity,Montreal,Canada Columbia,Vancouver,BCCanadaV6T1Z4
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