HANDBOOK OF BEHAVIORAL NEUROSCIENCE VOLUME 17 HANDBOOK OF ANXIETY AND FEAR Other volumes in HANDBOOK OF BEHAVIORAL NEUROSCIENCE (formerly Techniques in the Behavioral and Neural Sciences) Volume1: FeedingandDrinking,byF.ToatesandN.E.Rowland(Eds.),1987,ISBN0-444-80895-7 Volume2: Distribution-freeStatistics:Application-orientedApproach,byJ.Krauth,1988,ISBN0-444-80934-1,PaperbackISBN 0-444-80988-0 Volume3: MolecularNeuroanatomy,byF.W.VanLeeuwen,R.M.Bujis,C.W.PoolandO.Pach(Eds.),1989,ISBN0-444-81014-5, PaperbackISBN0-444-81016-1 Volume4: ManualofMicrosurgeryontheLaboratoryRat,Part1,byJ.J.vanDongen,R.Remie,J.W.RensemaandG.H.J.van Wunnik(Eds.),1990,ISBN0-444-81138-9,PaperbackISBN0-444-81139-7 Volume5: DigitalBiosignalProcessing,byR.Weitkunat(Ed.),1991,ISBN0-444-81140-0,PaperbackISBN0-444-98144-7 Volume6: ExperimentalAnalysisofBehavior,byI.H.IversenandK.A.Lattal(Eds.),1991,Part1,ISBN0-444-81251-2,Paperback ISBN0-444-89160-9,Part2,ISBN0-444-89194-3,PaperbackISBN0-444-89195-1 Volume7: MicrodialysisintheNeuroscience,byT.E.RobinsonandJ.B.Justice,Jr.(Eds.),1991,ISBN0-444-81194-X,Paperback ISBN0-444-89375-X Volume8: TechniquesfortheGeneticsAnalysisofBrainandBehavior,byD.Goldowitz,D.WahlstenandR.E.Wimer(Eds.),1992, ISBN0-444-81249-0,PaperbackISBN0-444-89682-1 Volume9: Research Design and Method in Psychiatry, by M. Fava and J.F. Rosenbaum (Eds.), 1992, ISBN 0-444-89595-7, PaperbackISBN0-444-89594-9 Volume10: MethodsinBehavioralPharmacology,byF.vanHaaren(Ed.),1993,ISBN0-444-81444-2,PaperbackISBN0-444-81445-0 Volume11: Methods in Neurotransmitter and Neuropeptide Research, by S.H. Parvez (Ed.), 1993, Part 1, ISBN 0-444-81369-1, PaperbackISBN0-444-81674-7,Part2,ISBN0-444-81368-3,PaperbackISBN0-444-81675-5 Volume12: Neglected Factors in Pharmacology and Neuroscience Research, by V. Claassen (Ed.), 1994, ISBN 0-444-81871-5, PaperbackISBN0-444-81907-X Volume13: HandbookofMolecular-GeneticTechniquesforBrainandBehaviorResearch,byW.E.CrusioandR.T.Gerlai(Eds.), 1999,ISBN0-444-50239-4 Volume14: ExperimentalDesign.AHandbookandDictionaryforMedicalandBehavioralResearch,byJ.Krauth(Ed.),2000,ISBN 0-444-50637-3,PaperbackISBN0-444-50638-1 Volume15: Handbook of Stress and the Brain, by T. Steckler, N.H. Kalin and J.M.H.M. Reul (Eds.), 2005, Part 1, ISBN 0-444-51173-3,Part2,ISBN0-444-51823-1 Volume16: HandbookofMicrodialysis.Methods,ApplicationsandPerspectives,byB.H.C.WesterinkandT.I.F.H.Cremers(Eds.), 2007,ISBN978-0-444-52276-4 HANDBOOK OF BEHAVIORAL NEUROSCIENCE Series Editor J.P. HUSTON Du¨sseldorf VOLUME 17 HANDBOOK OF ANXIETY AND FEAR Edited by ROBERT J. BLANCHARD DepartmentofPsychology,UniversityofHawaiiatManoa,Honolulu,HI,USA D. CAROLINE BLANCHARD PacificBiosciencesResearchInstitute,andDepartmentofGeneticsandMolecularBiology,JohnA.Burns SchoolofMedicine,UniversityofHawaiiatManoa,Honolulu,HI,USA GUY GRIEBEL Sanofi-Aventis,PsychopharmacologyDepartment,Bagneux,France DAVID NUTT PsychopharmacologyUnit,UniversityofBristol,Bristol,UK AMSTERDAM–BOSTON–HEIDELBERG–LONDON–NEWYORK–OXFORD PARIS–SANDIEGO–SANFRANCISCO–SINGAPORE–SYDNEY–TOKYO ACADEMIC AcademicPressisanimprintofElsevier PRESS AcademicPressisanimprintofElsevier Radarweg29,POBox211,1000AEAmsterdam,TheNetherlands LinacreHouse,JordanHill,OxfordOX28DP,UK Firstedition2008 Copyrightr2008ElsevierB.V.Allrightsreserved Nopartofthispublicationmaybereproduced,storedinaretrievalsystem ortransmittedinanyformorbyanymeanselectronic,mechanical,photocopying, recordingorotherwisewithoutthepriorwrittenpermissionofthepublisher PermissionsmaybesoughtdirectlyfromElsevier’sScience&TechnologyRights DepartmentinOxford,UK:phone(+44)(0)1865843830;fax(+44)(0)1865853333; email:permissions@elsevier.com.Alternativelyyoucansubmityourrequestonlineby visitingtheElsevierwebsiteathttp://www.elsevier.com/locate/permissions,andselecting ObtainingpermissiontouseElseviermaterial Notice Noresponsibilityisassumedbythepublisherforanyinjuryand/ordamagetopersons orpropertyasamatterofproductsliability,negligenceorotherwise,orfromanyuse oroperationofanymethods,products,instructionsorideascontainedinthematerial herein.Becauseofrapidadvancesinthemedicalsciences,inparticular,independent verificationofdiagnosesanddrugdosagesshouldbemade LibraryofCongressCataloging-in-PublicationData AcatalogrecordforthisbookisavailablefromtheLibraryofCongress BritishLibraryCataloguinginPublicationData AcataloguerecordforthisbookisavailablefromtheBritishLibrary ISBN:978-0-444-53065-3(thisvolume) ISSN:1569-7339(Series)(formerly0921-0709) ForinformationonallAcademicPresspublications visitourwebsiteatbooks.elsevier.com PrintedandboundinTheNetherlands 08 09 10 11 12 10 9 8 7 6 5 4 3 2 1 List of Contributors JamesL.Abelson(455),DepartmentofPsychiatry,UniversityofMichigan,RachelUpjohnBuilding,4250 Plymouth Road Ann Arbor, MI 48109-5765, USA David S. Baldwin (395), Clinical Neuroscience Division, School of Medicine, University of Southampton, UniversityDepartmentofMentalHealth,RSHHospital,BrintonsTerrace,SouthamptonSO14OYG,UK Catherine Belzung (325), EA3248 Psychobiologie des Emotions, Universite´ Franc-ois Rabelais, UFR Sciences et Techniques, Parc Grandmont, 37200 Tours, France D. Caroline Blanchard (3, 63, 81, 141), Department of Genetics and Molecular Biology, John A. Burns SchoolofMedicine;andPacificBiosciencesResearchCenter,UniversityofHawaiiatManoa,1993East West Road, Honolulu, HI 96822, USA Robert J. Blanchard (3, 63, 81), Department of Psychology, University of Hawaii at Manoa, Gartley Hall 2430 Campus Road, Honolulu, HI 96822, USA MarcoBortolato(303),DepartmentofCardiovascularandNeurologicalSciences,PoliclinicoUniversitario, University of Cagliari, S.S. 554km 4.500, 09042 Monserrato (CA), Italy Christopher K. Cain (103), Center for Neural Science, New York University, 4 Washington Place, Room 809, New York, NY 10003, USA Newton S. Canteras (125, 141), Departamento de Anatomia, Instituto de Cieˆncias Biome´dicas, Universidade de Sa˜o Paulo, Av. Lineu Prestes, 2415 CEP 05508-900, Sa˜o Paulo, SP, Brazil Antonio Pa´dua Carobrez (241), Department of Pharmaoclogy, CCB, Universidade Federal de Santa Catarina, Campus Universita´rio-Trindade, Floriano´polis, SC 88040-900, Brazil John F. Cryan (269), School of Pharmacy, Department of Pharmacology and Therapeutics, Alimentary Pharmabiotic Centre, Cavanagh Pharmacy Building, University College Cork, Cork, Ireland SimonJ.C.Davies(365),AcademicUnitofPsychiatry,UniversityofBristol,CothamHouse,CothamHill, Bristol BS6 6JL, UK Michael Davis (49), Department of Psychiatry and Behavioral Sciences, Center for Behavioral Neuroscience, Emory University, 4th Floor, WMB, 1639 Pierce Drive, Atlanta, GA 30322, USA Kumlesh K. Dev (269), Department of Anatomy, Biosciences Institute, University College Cork, Cork, Ireland Gabriel Esquivel (413), Experimental and Clinical Psychiatry Section, Department of Psychiatry and Neuropsychology,FacultyofHealth,MedicineandLifeSciences,MaastrichtUniversity,Vijverdalseweg 1, k.2.039, Gebouw Concorde, 6226 NB Maastricht, The Netherlands Michael S. Fanselow (29), Psychology and the Brain Research Institute, University of California, 1285 Franz Hall, Los Angeles, CA 90095, USA Berta Garcia de Miguel (365), Psychopharmacology Unit, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK Matthew Garner (395), Clinical Neuroscience Division, School of Medicine, University of Southampton, Southampton SO14 OYG, UK FredericoGuilhermeGraeff(241),DepartmentofNeurology,PsychiatryandMedicalPsychology,Hospital das Clı´nicas da Faculdade de Medicina de Ribeira˜o Preto, Universidade de Sa˜o Paulo, Avenida Bandeirantes 3900, Ribeira˜o Preto, SP 14090-900, Brazil v vi Guy Griebel (3, 325), Sanofi-Aventis, Psychopharmacology Department, 31 Avenue Paul Vaillant- Couturier, 92220 Bagneux, France Eric Griez (413), Experimental and Clinical Psychiatry Section, Department of Psychiatry and Neuro- psychology, Faculty of Health, Medicine and Life Sciences, Maastricht University, Vijverdalseweg 1, k.2.039, Gebouw Concorde, 6226 NB Maastricht, The Netherlands Francisco Silveira Guimara˜es (241), Department of Pharmacology, Faculdade de Medicina de Ribeira˜o Preta, Universidade de Sa˜o Paulo, Avenida Bandeirantes 3900, Ribeira˜o Preto, SP 14090-900, Brazil John M. Hettema (475), Virginia Institute for Psychiatric and Behavioral Genetics, Department of Psychiatry, Virginia Commonwealth University, 800 East Leigh Street, Room 1-116, Richmond, VA 23219, USA Andrew Holmes (355), Section on Behavioral Science and Genetics, Laboratory for Integrative Neuroscience, National Institute on Alcohol Abuse and Alcoholism, 5625 Fishers Lane Room 2N09, Rockville, MD 20852-9411, USA JosephE.LeDoux(103),CenterforNeuralScience,NewYorkUniversity,4WashingtonPlace,Room809, New York, NY 10003, USA Samuel Leman (325), EA3248 Psychobiologie des Emotions, Universite´ Franc-ois Rabelais, UFR Sciences et Techniques, Parc Grandmont, 37200 Tours, France Israel Liberzon (455), Department of Psychiatry, University of Michigan, Rachel Upjohn Building, 4250 Plymouth Road, Ann Arbor, MI 48109-5765, USA YoavLitvin(81),DepartmentofPsychology,UniversityofHawaiiatManoa,2430CampusRoad,Gartley Hall 110, Honolulu, HI 96822, USA Andrea L. Malizia (437), Academic Unit of Psychiatry and Psychopharmacology Unit, University of Bristol, Cotham House, Cotham Hill, Bristol BS6 6JL, UK Eduard Maron (475), Department of Psychiatry, University of Tartu, Tartu, Estonia; and Research Department of Mental Health, The North Estonian Regional Hospital, Psychiatry Clinic, Tallinn, Estonia Karyn M. Myers (49), Department of Psychiatry and Behavioral Sciences, Center for Behavioral Neuroscience, Emory University, 954 Gatewood Road NE, Atlanta, GA 30329, USA Neil McNaughton (11), Department of Psychology, University of Otago, P.O. Box 56, Dunedin, New Zealand David Nutt (3, 365, 437), Psychopharmacology Unit, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK Nathan S. Pentkowski (81), Department of Psychology, University of Hawaii at Manoa, 2430 Campus Road, Gartley Hall 110, Honolulu, HI 96822, USA Daniele Piomelli (303), Department of Pharmacology and Center for Drug Discovery, 3101 Gillespie Neuroscience Facility, University of California, Irvine, CA 92697-4625, USA Roger L. Pobbe (81), Pacific Biosciences Research Center, 1993 East-West Road, Honolulu, HI 96822, USA Ravikumar Ponnusamy (29), Psychology and the Brain Research Institute, University of California, 1285 Franz Hall, Los Angeles, CA 90095, USA James K.Rowlett(223),HarvardMedical School,New England Primate ResearchCenter,Box 9102, One Pine Hill Drive, Southborough, MA 01772-9102, USA Koen Schruers(413),Experimental and Clinical PsychiatrySection, Department of Psychiatry and Neuro- psychology, Faculty of Health, Medicine and Life Sciences, Maastricht University, Vijverdalseweg 1, k.2.039, Gebouw Concorde, 6226 NB Maastricht, The Netherlands Jakov Shlik (475), Department of Psychiatry, University of Ottawa, Royal Ottawa Health Care Group, 1145 Carling Avenue, Ottawa, Ont. K1Z7K4, Canada vii Thomas Steckler (157), Department of Psychiatry, Research and Early Development Europe, Johnson & Johnson Pharmaceutical Research and Development, Turnhoutseweg 30, 2340 Beerse, Belgium ElizabethA. Young(455),Molecular and Behavioral Neurosciences Institute,205 Zina Pitcher Place, Ann Arbor, MI 48109-5720, USA He´lio Zangrossi Jr. (11), Department of Pharmacology, School of Medicine of Ribeira˜o Preto, University of Sa˜o Paulo, Av. Bandeirantes 3900, 14049-900, Ribeira˜o Preto-SP, Brazil This page intentionally left blank Preface Fearandanxietyconstitutecrucialemotionalbehaviors,manifesttosomedegreeinvirtuallyallchordates andmagnificentlyrepresentedinhumanbehaviorandhistory.Becauseoftheirimportance,theyhavebeen intensively investigated from a number of scientific perspectives. The commonalities and consistencies of theircrossspeciesrepresentation,whilebynomeanstotal,alsosuggestafoundationfortheiranalysisthat is not available for more evanescent, subtle, or idiosyncratic emotions. This volume provides examples of the variety and intensity of scientific attention to fear and anxiety. We hope that it also indicates good progresstowardunderstandingthemechanismsandfunctionsoftheseemotionsandthebehaviorpatterns with which they are associated. R.J. Blanchard D.C. Blanchard G. Griebel D. Nutt (The Editors) ix