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Coverphotocredit: Meyer,T.,Herrmann-Lingen,C. NatriureticPeptidesinAnxietyandPanicDisorder VitaminsandHormones(2017)103,pp.131-146. AcademicPressisanimprintofElsevier 50HampshireStreet,5thFloor,Cambridge,MA02139,UnitedStates 525BStreet,Suite1800,SanDiego,CA92101-4495,UnitedStates TheBoulevard,LangfordLane,Kidlington,OxfordOX51GB,UnitedKingdom 125LondonWall,London,EC2Y5AS,UnitedKingdom Firstedition2017 Copyright©2017ElsevierInc.Allrightsreserved. Nopartofthispublicationmaybereproducedortransmittedinanyformorbyanymeans, electronicormechanical,includingphotocopying,recording,oranyinformationstorageand retrievalsystem,withoutpermissioninwritingfromthepublisher.Detailsonhowtoseek permission,furtherinformationaboutthePublisher’spermissionspoliciesandour arrangementswithorganizationssuchastheCopyrightClearanceCenterandtheCopyright LicensingAgency,canbefoundatourwebsite:www.elsevier.com/permissions. Thisbookandtheindividualcontributionscontainedinitareprotectedundercopyrightby thePublisher(otherthanasmaybenotedherein). Notices Knowledgeandbestpracticeinthisfieldareconstantlychanging.Asnewresearchand experiencebroadenourunderstanding,changesinresearchmethods,professionalpractices, ormedicaltreatmentmaybecomenecessary. Practitionersandresearchersmustalwaysrelyontheirownexperienceandknowledgein evaluatingandusinganyinformation,methods,compounds,orexperimentsdescribed herein.Inusingsuchinformationormethodstheyshouldbemindfuloftheirownsafetyand thesafetyofothers,includingpartiesforwhomtheyhaveaprofessionalresponsibility. Tothefullestextentofthelaw,neitherthePublishernortheauthors,contributors,oreditors, assumeanyliabilityforanyinjuryand/ordamagetopersonsorpropertyasamatterof productsliability,negligenceorotherwise,orfromanyuseoroperationofanymethods, products,instructions,orideascontainedinthematerialherein. ISBN:978-0-12-811914-3 ISSN:0083-6729 ForinformationonallAcademicPresspublications visitourwebsiteathttps://www.elsevier.com/ Publisher:ZoeKruze AcquisitionEditor:AlexWhite EditorialProjectManager:HeleneKabes ProductionProjectManager:VigneshTamil CoverDesigner:MilesHitchen TypesetbySPiGlobal,India Former Editors ROBERT S. HARRIS KENNETH V. THIMANN Newton, Massachusetts University of California Santa Cruz, California JOHN A. LORRAINE IRA G. WOOL University of Edinburgh Edinburgh, Scotland University of Chicago Chicago, Illinois PAUL L. MUNSON EGON DICZFALUSY University of North Carolina Chapel Hill, North Carolina Karolinska Sjukhuset Stockholm, Sweden JOHN GLOVER ROBERT OLSEN University of Liverpool Liverpool, England School of Medicine State University of New York GERALD D. AURBACH at Stony Brook Stony Brook, New York Metabolic Diseases Branch National Institute of DONALD B. MCCORMICK Diabetes and Digestive and Kidney Diseases Department of Biochemistry National Institutes of Health Emory University School of Bethesda, Maryland Medicine, Atlanta, Georgia CONTRIBUTORS D.C.Aguiar InstituteofBiologicalSciences,UniversidadeFederaldeMinasGerais,BeloHorizonte, Brazil B.Bonaz CliniqueUniversitaired’H(cid:1)epato-Gastroent(cid:1)erologie,CHUdeGrenoble;Universit(cid:1)e GrenobleAlpes,GrenobleInstitutdesNeurosciences,FonctionsC(cid:1)er(cid:1)ebraleset Neuromodulation,INSERM,Grenoble09,France A.P.Borrow ColoradoStateUniversity,FortCollins,CO,UnitedStates M.A.Briscione TraumaRecoveryProgram,MentalHealthServiceLine,AtlantaVeteransAffairsMedical Center,Decatur;EmoryUniversitySchoolofMedicine,Atlanta,GA,UnitedStates P.Chanana UniversityInstituteofPharmaceuticalSciences,UGCCentreofAdvancedStudy,Panjab University,Chandigarh,India U.Doboszewska FacultyofPharmacy,JagiellonianUniversityMedicalCollege,Krakow,Poland M.Gaweł FacultyofPharmacy,JagiellonianUniversityMedicalCollege,Krakow,Poland F.V.Gomes UniversityofPittsburgh,Pittsburgh,PA,UnitedStates F.S.Guimara˜es MedicalSchoolofRibeira˜oPreto,UniversityofSa˜oPaulo;CenterforInterdisciplinary ResearchonAppliedNeurosciences(NAPNA),UniversityofSa˜oPaulo(USP),Ribeira˜o Preto,Brazil K.Gulati VallabhbhaiPatelChestInstitute,UniversityofDelhi,Delhi,India R.J.Handa ColoradoStateUniversity,FortCollins,CO,UnitedStates S.M.J.Hemmings FacultyofMedicineandHealthSciences,DepartmentofPsychiatry,Stellenbosch University,CapeTown,SouthAfrica C.Herrmann-Lingen UniversityofGo€ttingenMedicalCentre;GermanCentreforCardiovascularResearch, UniversityofGo€ttingen,Go€ttingen,Germany F.Huang YiyangMedicalCollege,Yiyang,Hunan,PRChina xi xii Contributors T.Jovanovic EmoryUniversitySchoolofMedicine,Atlanta,GA,UnitedStates A.Kumar UniversityInstituteofPharmaceuticalSciences,UGCCentreofAdvancedStudy,Panjab University,Chandigarh,India C.-Q.Li SchoolofBasicMedicalSciences,CentralSouthUniversity,Changsha,Hunan,PRChina S.F.Lisboa MedicalSchoolofRibeira˜oPreto,UniversityofSa˜oPaulo;CenterforInterdisciplinary ResearchonAppliedNeurosciences(NAPNA),UniversityofSa˜oPaulo(USP),Ribeira˜o Preto,Brazil S.Malan-Mu€ller FacultyofMedicineandHealthSciences,DepartmentofPsychiatry,Stellenbosch University,CapeTown,SouthAfrica T.Meyer UniversityofGo€ttingenMedicalCentre;GermanCentreforCardiovascularResearch, UniversityofGo€ttingen,Go€ttingen,Germany V.Michopoulos EmoryUniversitySchoolofMedicine;YerkesNationalPrimateResearchCenter,Atlanta, GA,UnitedStates K.Młyniec FacultyofPharmacy,JagiellonianUniversityMedicalCollege,Krakow,Poland F.A.Moreira InstituteofBiologicalSciences,UniversidadeFederaldeMinasGerais,BeloHorizonte, Brazil S.D.Norrholm TraumaRecoveryProgram,MentalHealthServiceLine,AtlantaVeteransAffairsMedical Center,Decatur;EmoryUniversitySchoolofMedicine,Atlanta,GA,UnitedStates G.Nowak FacultyofPharmacy,JagiellonianUniversityMedicalCollege,Krakow,Poland S.Pellissier LaboratoireInteruniversitairedePsychologie,Personnalit(cid:1)e,Cognition,ChangementSocial, Universit(cid:1)eSavoieMont-Blanc,Chamb(cid:1)ery,France N.Rai VallabhbhaiPatelChestInstitute,UniversityofDelhi,Delhi,India A.Ray VallabhbhaiPatelChestInstitute,UniversityofDelhi,Delhi,India Contributors xiii L.B.M.Resstel MedicalSchoolofRibeira˜oPreto,UniversityofSa˜oPaulo;CenterforInterdisciplinary ResearchonAppliedNeurosciences(NAPNA),UniversityofSa˜oPaulo(USP),Ribeira˜o Preto,Brazil G.Starowicz FacultyofPharmacy,JagiellonianUniversityMedicalCollege,Krakow,Poland A.L.B.Terzian MedicalSchoolofRibeira˜oPreto,UniversityofSa˜oPaulo;CenterforInterdisciplinary ResearchonAppliedNeurosciences(NAPNA),UniversityofSa˜oPaulo(USP),Ribeira˜o Preto,Brazil Z.Yang YiyangMedicalCollege,Yiyang,Hunan,PRChina PREFACE While occasional episodes of anxiety are normal life experiences, anxiety disorderischronicandseemstobeinherited.Panicattacksandsocialanxiety aretwoconstituentsofanxietydisorder.Somedisorderthatincludesanxiety intheUnitesStatesalonemayinvolveabout40millionpersons.Theprob- abilityof developingany anxietydisorderapproaches 30%, andfemales are 60% more likely to have anxiety disorders than males. The non-Hispanic white population is more susceptible to having anxiety disorder than any other group. Anxietydisordercanbedissectedintoatleastfivesubgroups.Thus,there areabout6.8millionpersonsintheUnitedStateswithgeneralizedanxiety disorder,2.2millionpersonshaveobsessivecompulsivedisorder,7.7million persons have posttraumatic stress disorder, 15 million persons have social phobia,and19million personshave aspecificothertypeof phobia.Obvi- ously,thesestatisticsequatetoamajorhealthproblem,requiringinvestiga- tionsintothemechanismsinvolved.Neuroendocrineinvolvementisclearly indicated. This volume reviews the roles that hormones and other factors play in anxiety and stress. Intheopeningchapter,A.Ray,K.Gulati,andN.Raiexamine“Stress, anxiety, and immunomodulation: a pharmacological analysis.” Next, A.P. Borrow and R.J. Handa report on “Estrogen receptors and modulation ofanxiety-likebehavior.”Followingthis,“Neuroendocrineunderpinnings of increased risk for posttraumatic stress disorder in women” is the topic reviewedbyM.A.Briscione,V.Michopoulos,T.Jovanovic,andS.D.Nor- rholm. The ever-enlarging role of RNAs in disease is exemplified by the chapterbyS.Malan-Mu€llerandS.M.J.Hemmingson“Thebigroleofsmall RNAsinanxietyandstress-relateddisorders.”T.MeyerandC.Herrmann- Lingen then describe “Natriuretic peptides in anxiety and panic disorder.” Two chapters deal with the relatively new gaseous hormone, nitric oxide. The first is “The role of nitric oxide in stress-induced anxiety: from path- ophysiologytotherapeutictarget”byA.KumarandP.Chanana.Thesec- ond is “Nitric oxide and anxiety” by K. Gulati, N. Rai, and A. Ray. S.F. Lisboa, F.V. Gomes, A.L.B. Terzian, D.C. Aguiar, F.A. Moreira, L.B.M. Resstel, and F.S. Guimara˜es report on “The endocannabinoid system and anxiety.”“Themelatonergicsysteminanxietydisordersandtheroleofmel- atonininconditionalfear”isthesubjectofF.Huang,Z.Yang,andC.-Q.Li. xv xvi Preface “The role of elements in anxiety” is examined by K. Młynec, M. Gaweł, U. Doboszewska, G. Starowicz, and G. Nowak. In the final chapter, S. PellissierandB.Bonazdescribe“Theplaceofstressandemotionsintheirri- table bowel syndrome.” TheillustrationonthecoverisareproductionofFig.3fromChapter5 by T. Meyer and C. Herrmann-Lingen entitled: “Natriuretic peptides in anxiety and panic disorder.” (A) Crystal structure of the extracellular ligand-bindingdomainoftheatrialnatriuretic(ANP)receptordimercom- plexed with rat ANP7-27. The receptor structure is shown as a ribbon model with carbohydrate structures depicted in magenta, while the receptor-boundhormoneisdepictedinaspace-fillingmodelwithnitrogen atomsmarkedinblue,oxygenatomsinred,andsulfuratomsinyellow.The figurewasdrawnwithPyMOL(DeLanoScientific)usingdatafromthePro- teinDatabasefile1T34(Ogawaetal.,2004).(B)Overviewofthepleiotro- pic functions of cGMP-mediated ANP signaling including its effect on anxiety. Publication of this book was facilitated by Helene Kabes of Elsevier, Oxford, UK, and by Vignesh Tamilselvvan, Elsevier, S&T Book Produc- tion, Chennai, India. GERALD LITWACK July 18, 2016 North Hollywood, California CHAPTER ONE Stress, Anxiety, and Immunomodulation: A Pharmacological Analysis A. Ray1, K. Gulati, N. Rai VallabhbhaiPatelChestInstitute,UniversityofDelhi,Delhi,India 1Correspondingauthor:e-mailaddress:[email protected] Contents 1. Introduction 2 2. MechanismofStressResponse 4 3. Stress,CNS,andBehavior 5 4. StressandAnxiety 7 5. StressandVariousAnxiolyticDrugs 9 6. StressandImmunity 11 7. Stress,Anxiety,andImmunomodulation 14 References 20 Abstract Stressandstressfuleventsarecommonoccurrencesinourdailylivesandsuchaversive situationsbringaboutcomplexchangesinthebiologicalsystem.Suchstressresponses influence the brain and behavior, neuroendocrine and immune systems, and these responsesorchestratetoincreaseordecreasetheabilityoftheorganismtocopewith such stressors. The brain via expression of complex behavioral paradigms controls peripheralresponsestostressandabidirectionallinkexistsinthemodulationofstress effects.Anxietyisacommonneurobehavioralcorrelateofavarietyofstressors,andboth acuteandchronicstressexposurecouldprecipitateanxietydisorders.Psychoneuroim- munologyinvolvesinteractionsbetweenthebrainandtheimmunesystem,anditis nowbeingincreasinglyrecognizedthattheimmunesystemcouldcontributetothe neurobehavioralresponsestostress.Studieshaveshownthatthebrainanditscomplex neurotransmitternetworkscouldinfluenceimmunefunction,andtherecouldbeapos- sible link between anxiogenesis and immunomodulation during stress. Physiological andpharmacologicaldatahavehighlightedthisconcept,andthepresentreviewgives anoverviewoftherelationshipbetweenstress,anxiety,andimmuneresponsiveness. VitaminsandHormones,Volume103 #2017ElsevierInc. 1 ISSN0083-6729 Allrightsreserved. http://dx.doi.org/10.1016/bs.vh.2016.09.007 2 A.Rayetal. 1. INTRODUCTION Stressisaresponsetoaversivestimuli,externalorinternal,capableof altering the physiological homeostasis and the ability to cope with such stressful situation is a crucial determinant of health and disease. Exposure to adverse conditions initiates a series of adaptive response organized to defendthestabilityoftheinternalenvironmentandenhanceanorganism’s survival. This orchestrated process usually referred to as “stress response” involves various mechanisms that allow the body to make necessary physi- ological and metabolic adjustments required to cope with the demands of homeostatic challenge (Gold & Chrousos, 2002). Such changes may occur on the physiological (emotional and cognitive), behavioral, and biological level(alteredautonomicandneuroendocrinefunction).Thecentralnervous system (CNS) plays a crucial role in the regulation of stress responses, and complex neurochemical pathways have been proposed (Carrasco & Van de Kar, 2003). Intheearly19thcentury,theFrenchphysiologistClaudeBernardintro- duced a theory suggesting that, as organisms become more independent of their surroundings, they develop more complex ways of stabilizing their internalenvironmentstocounterthechangesintheirexternalenvironment. The importance of adaptive mechanisms was thus recognized as the con- stancy of “milieu interieur,” which would be the condition of a free and independentexistence(Goldstein&Kopin,2007).Intheearly1900,Walter Cannonexpandedthistheoryandcoinedtheterm“homeostasis.”Hedem- onstrated in several seminal experiments that the sympathoadrenal system was responsible for coordinating the “fight and flight” response necessary to meet external challenges (Cannon, 1929; LeMoal, 2007). Cannon was able to show that both emotional and physical disturbances triggered the same response from the organism. In addition, he proposed that there was a “critical” level of stress, in terms of magnitude and duration, against whichthehomeostaticmechanismsfailandtheorganismperishes.Cannon believedthatanindividualorganism’ssusceptibilitytothiscriticalstressvar- iedunderdifferentgeneralconditionsandduringthenormalandpathologic upsanddownofexistenceinanordinarylifestyle(LeMoal,2007).In1930s Hans Selye began research on stress and thus helped in understanding the connection between stress and health (Selye, 1936). His definition of stress asbeingthe“nonspecificresponseofthebodytoanydemanduponit”was so persuasive that it persisted and remains widely used even today. By Stress,Anxiety,andImmunomodulation 3 “nonspecific,”Selyemeantasetofsharedelementsofresponses—regardless of the nature of the causative agent, or stressor. Selye proposed three uni- versal stages of coping with a stressor—the “General Adaptation Syn- drome”—an initial “alarm reaction,” analogous to Cannon’s “fight or flight” response, a stage of adaptation associated with resistance to the stressor, and eventually a stage of exhaustion and organismic death. It was laterdemonstratedthatthesechangesareassociatedwith,andtosomeextent resultfrom,activationofthehypothalamic–pituitary–adrenocortical(HPA) axis.Steroidsreleasedintothecirculationfromadrenalcortexcontributeto resistancebutmayalsoberesponsibleforpathologicalchanges.Selye’scon- ceptthatprolongedstresscanproducephysicaldiseaseandmentaldisorders isnowwidelyaccepted(Perdrizet,1997).MasoncriticizedSelye’stheoryof nonspecificity. Anxiety and fear were understood as the important factors that contribute to nonspecific responses on exposure to several stressors (Mason, 1971). More modern concepts view stress as a consciously or unconsciously sensedthreattohomeostasis,inwhichtheresponsehasadegreeofspecific- ity, depending among other things, on the particular challenge to homeo- stasis, the organism’s perception of the stressor and the perceived ability to cope with it. Although homeostasis suggests constancy of values for vari- ables,rangesofacceptablevaluesarenowrecognizedtobedecidedlyincon- stant (Korte, Koolhaas, Wingfield, & McEwen, 2005). There are diurnal variations in bodytemperature,heart rate, andblood pressure.In addition, adaptations to different stressors include alterations in acceptable levels for monitored variables. Sterling and Eyer (1988) introduced the term, “allostasis,” to describe the attainment of stability by natural alterations in acceptable ranges of variables. The brain is the site at which effects of stressorsaresensedandappropriatecoordinatedbehavioralandneuroendo- crine responses initiated. Adaptations involving allostasis to cope with real, simulated,orimaginedchallengesaredeterminedbygenetic,developmen- tal, and previous experimental factors. While they may be effective for a shortinterval,overtime,thealterationsmayhavecumulativeadverseeffects. For instance, chronic elevation of blood pressure ensures adequate blood flow to the brain eventually lead to atherosclerosis and stroke or coronary occlusion. Risk of such adverse effects is termed “allostatic load” (McEwen,1998).Theemergingconceptofallostaticloadprovidesameans ofaugmentingtheconceptofstressasaconceptualandmethodologicalbasis for elucidating the behavioral and physiological mechanisms by which genes,earlylifeexperiences,livingandworkingenvironment,interpersonal

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Anxiety, the latest volume in the Vitamins and Hormones series first published in 1943, and the longest-running serial published by Academic Press, provides up-to-date information onthe roles that hormones and other factors play in anxiety and stress.Each volume focuses on a single molecule or disea
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