Encyclopedia of neuroscience Ten vol. set

10798 Pages·2009·3.48 MB·English

by  Larry R Squire

EDITOR IN CHIEF
Larry R. Squire
Departments of Psychiatry, Neurosciences, and Psychology
University of California, San Diego and VA Medical Center
San Diego, CA
USA
SENIOR EDITORS
Thomas D. Albright
Systems Neurobiology Laboratories
Salk Institute for Biological Studies
La Jolla, CA
USA
Floyd E. Bloom
Molecular and Integrative Neuroscience Department
The Scripps Research Institute
La Jolla, CA
USA
Fred H. Gage
Laboratory of Genetics
Salk Institute for Biological Studies
La Jolla, CA
USA
Nicholas C. Spitzer
Neurobiology Section
University of California
La Jolla, CA
USA
ASSOCIATE EDITORS
CarroleeBarlow EdwardM.Callaway
BrainCells, Inc. Systems Neurobiology Laboratory
San Diego, CA Salk Institute for Biological Studies
USA San Diego, CA
USA
M.FlintBeal
WilliamA.Catterall
Department of Neurology and Neuroscience
Department of Pharmacology
Weill Medical College of Cornell University
University of Washington
New York, NY
Seattle, WA
USA
USA
DarwinK.Berg
VerneS.Caviness
Department of Biology
Department of Neurology
University of California
Massachusetts General Hospital
La Jolla, CA
Boston, MA
USA
USA
AndersBjorklund JohnE.Dowling
Department of Neurobiology Department of Neuroscience
Lund University Harvard University
Lund Cambridge, MA
Sweden USA
JamesEberwine
M.ChristianBrown
Department of Pharmacology
Eaton-Peabody Lab
University of Pennsylvania
Massachusetts Eye and Ear Infirmary
Philadelphia, PA
Boston, MA
USA
USA
HowardEichenbaum
GeoffreyBurnstock
Center for Memory and Brain
Autonomic Neuroscience Centre
Boston University
Royal Free and University College Medical School
Boston, MA
London
USA
UK
CalebE.Finch
JohnH.Byrne Department of Neurogerontology
University of Texas Medical School University of Southern California
Houston, TX Los Angeles, CA
USA USA
ii AssociateEditors
CharlesD.Gilbert ChristopherR.Kintner
The Rockefeller University Salk Institute for Biological Studies
New York, NY La Jolla, CA
USA USA
YukikoGoda KeithR.Kluender
MRC Cell Biology Unit Department of Psychology
University College London University of Wisconsin
London Madison, WI
UK USA
LawrenceS.B.Goldstein RichardJ.Krauzlis
Howard Hughes Medical Institute Salk Institute for Biological Studies
UCSD School of Medicine La Jolla, CA
La Jolla, CA USA
USA
WilliamB.Kristan
AntonyW.Goodwin
Division of Biological Sciences
Department of Anatomy and Cell Biology
University of California San Diego
University of Melbourne
La Jolla, CA
Victoria
USA
Australia
JosephE.LeDoux
TomasHokfelt
Center for Neural Science
Department of Neuroscience
New York University
Karolinska Institute
New York, NY
Stockholm
USA
Sweden
GregE.Lemke
LeslieIversen
Molecular Neurobiology Laboratory
Department of Pharmacology
Salk Institute for Biological Studies
University of Oxford
San Diego, CA
Oxford
USA
UK
DavidA.Lewis
EugeneM.Johnson
Departments of Psychiatry and Neuroscience
Department of Neurology
University of Pittsburgh
Washington University School of Medicine
Pittsburgh, PA
St. Louis, MO
USA
USA
PierreJ.Magistretti
EdwardG.Jones
Center for Psychiatric Neuroscience
Center for Neuroscience
Prilly
University of California
Switzerland
Davis, CA
USA
RobertC.Malenka
JonH.Kaas Department of Psychiatry and Behavioral Sciences
Department of Psychology Stanford University School of Medicine
Vanderbilt University Palo Alto, CA
Nashville, TN USA
USA
PeterR.Marler
HelmutKettenmann DepartmentofNeurobiology,PhysiologyandBehavior
Max-Delbru¨ck Center for Molecular Medicine UniversityofCalifornia
Berlin Davis,CA
Germany USA
AssociateEditors iii
BruceS.McEwen CraigE.L.Stark
The Rockefeller University Department of Neurobiology and Behavior
New York, NY University of California
USA Irvine, CA
USA
EarlK.Miller
ThePicowerInstituteforLearning andMemoryand RobertStickgold
Department of Brain and Cognitive Sciences Psychiatry Medical Department
Massachusetts Institute of Technology Harvard Medical School
Cambridge, MA Boston, MA
USA USA
PaulE.Sawchenko PeterL.Strick
Salk Institute for Biological Studies Pittsburgh Veterans Affairs Medical Center and
La Jolla, CA Department of Neurobiology
USA University of Pittsburgh
Pittsburgh, PA
WolframSchultz
USA
Department of Physiology, Development and
Neuroscience EdwardM.Stricker
Cambridge University Department of Neuroscience
Cambridge University of Pittsburgh
UK Pittsburgh, PA
USA
TerrenceJ.Sejnowski
Computational Neurobiology Lab ThomasC.Su¨dhof
Salk Institute for Biological Studies Stanford University of Medicine
La Jolla, CA Palo Alto, CA
USA USA
GordonM.Shepherd FredW.Turek
Department of Neurobiology Center for Sleep and Circadian Biology
Yale University Northwestern University
New Haven, CT Evanston, IL
USA USA
ClarkeR.Slater StephenC.Woods
Institute of Neuroscience Department of Psychiatry
Newcastle University University of Cincinnati
Newcastle upon Tyne Cincinnati, OH
UK USA
FOREWORD
Whatisanencyclopedia?Thetermisderivedfromtwo of selective lesions, and biochemists, who no longer
Greek words: enkuklios, which means cyclical, and were relegated to monitoring neurotransmitters in
paideia,whichmeanseducation.Intheearlysixteenth homogenates of the whole brain. Advances in the
century, copyists of Latin manuscripts combined the 1970s in receptor biochemistry, their localization by
twowordsintoaLatindesignationwhichcomestous autoradiography, and neuropeptide immunohisto-
inEnglishwiththesamespellingandwiththemean- chemistry further enhanced discourse among neuro-
ing ‘general course of instruction’ (The American physiologists, neuroanatomists, neurochemists, and
HeritageDictionary,2000,Boston:HoughtonMifflin, neuropharmacologists. In the past two decades, the
p. 589). The original Greek provides a dynamic toolsofmolecularbiologyhavefurtheredthedialogue.
connotation, an effort to approach knowledge in a The explosion of the neurosciences can also be
probing,integratedfashion.InLatinandthenEnglish, documented through the chronicles of the Society
the term has generally been applied to reference for Neuroscience (SFN). The SFN was founded in
volumesaddressingbroadareasofknowledge. 1970 with Vernon Mountcastle as the first elected
The Greek emphasis on creative approaches to president, and its inaugural annual meeting in
informationisparticularlyaptfortheneurosciences. Washington, DC hosted a few hundred researchers.
The term ‘neuroscience’ is of remarkably young vin- When I served as president in 1980, SFN numbered
tage. Francis Schmitt, in his foreword to the first 7000 members. One of my key tasks was to combat
edition of this encyclopedia, related his process of attacks on the raison d’eˆtre of the society. Some
concatenating scientists from disparate fields into an argued, “We have too many scientists in this organi-
invisiblecollegewhoseworkshopsattackedthebrain’s zation.Let’ssplitintotwosocieties,the‘Wets’andthe
mostrecalcitrantpuzzles.Hedubbedtheorganization ‘Drys.’” Instead, to emphasize the integrated nature
theNeuroscienceResearchProgram(NRP).Theearly ofthefield,welaunchedtheJournalofNeuroscience.
NRP‘associates’includedgiantssuchasMelvinCal- Also, we argued that growth might plateau and that
vin, whose Nobel Prize honored his work on photo- careful meeting organization would prevent indivi-
synthesis; the Nobel laureate physical chemist duals from getting ‘lost in a crowd.’ My prediction
Manfred Eigen; the biochemist Albert Lehninger; aboutaplateauwasoffthemark.Asofthiswriting,
andMarshallNirenberg,thenayoungmolecularbiol- May 2007, SFN numbers about 38000 active mem-
ogistinthethroesofbreakingthegeneticcode. bers,withupto35000peopleattendingeachannual
WhenSchmittestablishedtheNRP,neuroscienceas meeting, dwarfing any other biomedical research
anintegratedendeavorhardlyexisted.Theinvention society.
ofthe microelectrodewas permittingneurophysiolo- Most would agree that neuroscience is the most
gists to record from single cells, but characterizing integrated scientific discipline. As such, the concept
them biochemically was impossible. A major step of an encyclopedia, in the original Greek sense of a
forward was the emergence in the mid-1960s of the circleoflearning,isnotablyappropriateyetimmensely
Falck-Hillarp fluorescence microscopic techniques, challenging.Thecurrentedition,likeearlierones,suc-
which permitted selective visualization of catechol- ceeds by careful attention to organization and, most
amine and serotonin neurons. Mapping the aminer- importantly,totheselectionofthefinestresearchersas
gic pathways soon led to collaborative efforts of AssociateEditorsforindividualtopics.TheAssociate
behaviorists, who could examine the consequences Editorsareallseasonedveteransyetactiveresearchers
ii Foreword
whosevisionremainsattheforefrontoftheirfield.All a lucid, reader-friendly format. Such a publication
areasofimportancearecovered,from‘souptonuts.’ can be provocative and invigorating to the most
Emphasisiselegantlybalancedbetweenmolecularand sophisticated professionals. At the same time, the
systemsneuroscience. entries are presented in such an inviting fashion that
In this era of rapid advances, one can question theencyclopediaservesfornovicesastheidealentre´e
whether an encyclopedia, comprising a snapshot in intotheworldofthenervoussystem.
time,servesameaningfulfunction.Mightnotallthe
information in such an enterprise be obsolete soon SolomonH.Snyder
after publication? An effective encyclopedia, exem- DistinguishedServiceProfessor,
plifiedinthesevolumes,integratesdisparateareasin JohnsHopkinsUniversity
PREFACE
During the second half of the twentieth century, the programs,teachersinthelifesciences,clinicians,and
studyofthenervoussystemmovedfromaperipheral sciencewriters.
position within the biological and psychological The inaugural edition of this encyclopedia, which
sciences to become an interdisciplinary field called was the first comprehensive reference work for the
neuroscience. The new discipline brought bioche- field,waspublishedin1987undertheableleadership
mists, cell biologists, anatomists, physiologists, psy- ofGeorgeAdelman.Itincludedsome700entriesand
chologists,neurologists,andpsychiatrists–scientists appearedintwovolumes.Thesecondedition,edited
and clinicians from diverse backgrounds, all drawn by George Adelman and Barry Smith, appeared in
tothe promise and excitement ofstudyingthe brain. 1999 in two volumes and included more than 800
They aimed to discover the mechanisms of neuro- entries. A CD-ROM version of this edition was also
nalfunction,elucidatetheneuralsubstratesofbehav- published. A third edition, published only in elec-
ior and cognition, and learn about the diseases of tronicform,appearedin2004.
the nervous system. The development of the dis- TheEncyclopediaofNeuroscience includesnearly
cipline was catalyzed in 1969 by the formation of 1500 entries. The full work will be published online
the Society for Neuroscience, which now has nearly at Science Direct, which can be accessed with sub-
37000members. The first academic training pro- scriptionatwww.sciencedirect.com.Toassemblethe
grams for neuroscience were established in medical entries,theSeniorEditorsidentified46majorareasof
schools(theDepartmentofNeurosciencesattheUni- the discipline and then invited 46 Associate Editors,
versity of California, San Diego in 1965 and the all experts in their field, to survey the content of
Department of Neurobiology at Harvard University neurosciencewithineachoftheseareas.EachAssoci-
in 1966). The first undergraduate training programs ate Editor then invited 30 to 40 authors to prepare
in neuroscience were established in 1972at Amherst articles on specific topics, with the objective of
CollegeandatOberlinCollege,almamaterofNobel- obtaining complete coverage for each area. Many of
istRogerSperryandthreePast-PresidentsoftheSoci- the authors are the recognized leaders in their field.
etyforNeuroscience.Today,therearemorethan300 The result is a compendium of expert articles repre-
neuroscience departments and programs around the senting the currentworld ofneuroscience–the most
world. importantresearch,themostpowerfultools,andthe
The Encyclopedia of Neuroscience is intended to mostpromisingapplications.
catalog andexplicatethe rich, diverse subjectmatter Most of the entries are self-contained reviews
of the discipline and to facilitate communication that can be read as independent articles. Extensive
amongitssubspecialties.Itismeanttobeanauthori- cross-listing at the conclusion of each entry directs
tativesourceofinformationforallareasofneurosci- readers to articles on related topics. The principal
ence. It will hopefully make neuroscience more organizationoftheEncyclopedialiesinthealphabeti-
accessible to a wide range of readers, from students callyarrangedlistofentries.Inaddition,thecompre-
makingtheirfirstacquaintancewiththefieldtogen- hensive subject classification will help readers find
eralreadersseekinginformationaboutspecifictopics. related topics and appreciate the structure of the
Itshouldalsoserveasausefulreferenceforworking discipline.
neuroscientists and be useful as well to undergrad- Whilenosinglereferenceworkinneurosciencecan
uate and graduate students in neuroscience training claimtoincludeeverynotableideaandfactaboutthe
ii Preface
brain, the Senior Editors hope that these volumes De Souza and Richard Berryman, and the Editorial
provide a summary of contemporary neuroscience Assistants, Caroline Phipps, Afandi Mohamed and
thatisbothcomprehensiveandinstructive.Neurosci- Nicky Carter, for capably managing the formid-
ence is still a developing field, but the Encyclopedia able task of assembling and organizing the contents
will have succeeded if it conveys the considerable oftheEncyclopedia.AndrewLoweandLauraJackson,
promise that neuroscience offers for conquering the the Project Managers, diligently brought the project
diseasesthataffectthenervoussystemandforunder- throughitsseveralstagesofproduction.
standingthebrain,themind,andourselves.
The Senior Editors are grateful to the Develop- LarryR.Squire
mental Editors at Elsevier, Michael Bevan, Joanna Editor-in-Chief