Suppressing the Mind Anesthetic Modulation of Memory and Consciousness Edited by A H NTHONY UDETZ R P OBERT EARCE Suppressing the Mind Forfurthervolumes: http://www.springer.com/series/7678 · Anthony Hudetz Robert Pearce Editors Suppressing the Mind Anesthetic Modulation of Memory and Consciousness Editors AnthonyHudetz RobertPearce DepartmentofAnesthesiology DepartmentofAnesthesiology MedicalCollegeofWisconsin UniversityofWisconsin,Madison B6/319ClinicalScienceCenter 8701WatertownPlankRoad 600HighlandAvenue MilwaukeeWI53226 MadisonWI53792 USA USA [email protected] [email protected] ISBN978-1-60761-463-0 e-ISBN978-1-60761-462-3 DOI10.1007/978-1-60761-462-3 LibraryofCongressControlNumber:2009938710 ©HumanaPress,apartofSpringerScience+BusinessMedia,LLC2010 Allrightsreserved.Thisworkmaynotbetranslatedorcopiedinwholeorinpartwithoutthewritten permissionofthepublisher(HumanaPress,c/oSpringerScience+BusinessMedia,LLC,233Spring Street,NewYork,NY10013,USA),exceptforbriefexcerptsinconnectionwithreviewsorscholarly analysis.Useinconnectionwithanyformofinformationstorageandretrieval,electronicadaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. Theuseinthispublicationoftradenames,trademarks,servicemarks,andsimilarterms,eveniftheyare notidentifiedassuch,isnottobetakenasanexpressionofopinionastowhetherornottheyaresubject toproprietaryrights. Whiletheadviceandinformationinthisbookarebelievedtobetrueandaccurateatthedateofgoing topress,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityforany errorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,withrespect tothematerialcontainedherein. Printedonacid-freepaper springer.com Foreword Anestheticsproduceareversiblestateofunconsciousnessaccompaniedbyantero- gradeamnesia.Thisremarkablephenomenonbringsgreatrelieftosurgicalpatients and wonder to clinicians and scientists. To date, we do not fully understand the mechanismsbywhichanestheticsablateconscioussensationandmemory.Weare, however,makingprogress. This book presents original results as well as overviews of the current state of knowledge of the problem. It is authored by investigators who know the field well; their research at a number of levels has contributed substantially to our cur- rent understanding of anesthetic modulation of memory and consciousness. Most of the contributors were presenters at two workshops organized by Dr. Pearce and Dr. Hudetz at the 40th Annual Winter Conference on Brain Research, held at Snowmass Village, Colorado, from January 27 through February 2, 2007. One workshopfocusedonanestheticmodulationofconsciousnessandanotheronanes- theticmodulationofmemory.Sevenofthechaptersarebasedonmaterialpresented atthesesymposia–appropriatelyupdatedwithnewrelevantfindings.Thisinforma- tionissupplementedbychaptersonanesthesiaandsleep,computationalanalysisof the state of anesthesia, and the clinical phenomenon of “anesthesia awareness,” a topic that has recently received much public attention. With these three additional contributions,thebookthusincludes10chapters. Several excellent books on consciousness and memory have been published in recent years, but none of these has presented a systematic compilation of studies onanestheticmodulationofmemoryandconsciousness–atleastinaunifiedview of the subject matter. Likewise, several texts have been written about fundamen- tal anesthetic mechanisms, focusing on pharmacological, cellular, and molecular changes. However, no volume has bridged molecular, cellular, integrative, and systems-level effects, as we believe will be necessary to address the core issues ofanestheticmechanisms.Thisbookisintendedtofillthisneed.Wehopethatby buildingthesebridgesbetweenbenchandclinicalresearch,newideasandtestable hypotheses will emerge, so that future work will ultimately lead to an integrative theoryofanesthetic-inducedunconsciousnessandamnesia. There is a long history of interest in unraveling the mechanisms of anesthesia. Withtherecentintroductionofseveralnewinvestigativemethodologies,andasnew hypotheses have emerged, there has been a surge in interest from the traditional, v vi Foreword pharmacological, and clinical neurosciences, as well as newer fields, such as cog- nitiveandcomputationalneurosciences.Consideringtheincrediblesignificanceof understanding the neurobiological basis of consciousness and memory, we expect that this interest will continue to grow. This book should appeal to anesthesiolo- gists, neurologists, psychologists, scientists, and anyone interested in anesthesia, consciousness, or memory. We hope that it serves as a reference for the scientific community and provides a useful perspective for future treatments of the subject. As a summary of the current state of knowledge, it should serve as a useful text for graduate students and researchers who wish to engage in anesthesia research. Althoughasignificantpartoftheinformationincludedhereistechnical,itiswrit- ten in a style that we hope makes it accessible to a wider audience than simply scientistswhoarecurrentlyengagedinresearchinthefield. The editors would like to express their sincere thanks to all contributors for their outstanding work. They appreciate the reviewers’ suggestions for the inclu- sion of additional specific topics. Special thanks are due to Patrick J. Marton and MatthewGiampoalaatSpringerUSfortheinvitationtopreparethisbook,andfor theeditorialassistanceofMarnieFilstein. MilwaukeeandMadison,WI AnthonyHudetz,DBM,PhDand RobertPearce,MD,PhD Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 AnthonyHudetzandRobertPearce 2 Molecular Targets of General Anesthetics intheNervousSystem . . . . . . . . . . . . . . . . . . . . . . . . . 11 HughC.Hemmings,Jr. 3 A Neurochemical Perspective on States ofConsciousness . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 ChristopherJ.Watson,HelenA.Baghdoyan,andRalphLydic 4 AnestheticModulationofAuditoryPerception:Linking Cellular,Circuit,andBehavioralEffects . . . . . . . . . . . . . . . 81 MatthewI.Banks 5 Cortical Disintegration Mechanism ofAnesthetic-InducedUnconsciousness . . . . . . . . . . . . . . . 99 AnthonyHudetz 6 AnesthesiaandtheThalamocorticalSystem . . . . . . . . . . . . . 127 MichaelT.Alkire 7 Anesthesia-Induced State Transitions inNeuronalPopulations . . . . . . . . . . . . . . . . . . . . . . . . 139 JamieSleigh,MoiraSteyn-Ross,AlistairSteyn-Ross, LoganVoss,andMarcusWilson 8 Anesthesia Awareness: When the Mind IsNotSuppressed. . . . . . . . . . . . . . . . . . . . . . . . . . . . 161 GeorgeA.Mashour 9 Loss of Recall and the Hippocampal Circuit Effects ProducedbyAnesthetics . . . . . . . . . . . . . . . . . . . . . . . . 175 M.BruceMacIver vii viii Contents 10 ModulationoftheHippocampalθ-RhythmasaMechanism forAnesthetic-InducedAmnesia . . . . . . . . . . . . . . . . . . . 193 MishaPerouanskyandRobertPearce 11 PropofolAmnesia–WhatisGoingonintheBrain?. . . . . . . . . 215 RobertA.VeselisandKaneO.Pryor SubjectIndex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 Contributors MichaelT.Alkire DepartmentofAnesthesiologyandPerioperativeCare, UniversityofCalifornia,IrvineMedicalCenter,Orange,CA,USA HelenA.Baghdoyan DepartmentofAnesthesiology,UniversityofMichigan, AnnArbor,MI,USA MatthewI.Banks DepartmentofAnesthesiology,UniversityofWisconsin, Madison,WI,USA HughC.Hemmings,Jr. DepartmentofAnesthesiology,WeillCornellMedical College,NewYork,NY,USA AnthonyHudetz DepartmentofAnesthesiology,MedicalCollegeofWisconsin, Milwaukee,WI,USA RalphLydic DepartmentofAnesthesiology,UniversityofMichigan,AnnArbor, MI,USA M.BruceMacIver DepartmentofAnesthesia,StanfordUniversitySchoolof Medicine,Stanford,CA,USA GeorgeA.Mashour DepartmentsofAnesthesiologyandNeurosurgery, UniversityofMichiganMedicalSchool;AnnArbor,MI,USA RobertPearce DepartmentofAnesthesiology,UniversityofWisconsin,Madison, WI,USA MishaPerouansky DepartmentofAnesthesiology,UniversityofWisconsin, Madison,WI,USA KaneO.Pryor DepartmentofAnesthesiologyandCriticalCareMedicine, MemorialSloan-KetteringCancerCenter,NewYork,NY,USA;Departmentof Anesthesiology,WeillCornellMedicalCollege,NewYork,NY,USA JamieSleigh DepartmentofAnaesthesiology,UniversityofAuckland,Hamilton, NewZealand AlistairSteyn-Ross UniversityofWaikato,Hamilton,NewZealand ix x Contributors MoiraSteyn-Ross UniversityofWaikato,Hamilton,NewZealand RobertA.Veselis DepartmentofAnesthesiologyandCriticalCareMedicine, MemorialSloan-KetteringCancerCenter,NewYork,NY,USA;Departmentof Anesthesiology,WeillCornellMedicalCollege,NewYork,NY,USA LoganVoss DepartmentofAnaesthesiology,UniversityofAuckland,Hamilton, NewZealand ChristopherJ.Watson DepartmentofAnesthesiology,UniversityofMichigan, AnnArbor,MI,USA MarcusWilson UniversityofWaikato,Hamilton,NewZealand