Table Of ContentTheN eurboi ologoyfN euraNle tworks
CopyrighMtaetde rial
ComputationNaelu roscience
TerreJn.Sc eej nowasnkdTi o masAo.P oggeidoi,t ors
MethodisnN euronMaold eliFnrgo:m S ynapsteoNs e tworeksd,i tbeyd
ChrisKtoocfah n dI daSne ge1v98,9
NeuraNle tsi nE lectFriischW ,a ltHeeri lige1n9b9e1 rg,
TheC omputatiBornaailPn ,a trSi.cC ihau rchlaanndd
TerreJn.Sc eej now1s99k2i ,
DynamiBci ologNiectawlo rkTsh:eS tomatogaNsetrrviocuS sy steemd, ited
byR onaMl.dH arris-WEavreMr aircdke,Ar l,l Ie.Sn e lverasntdo n,
MauriMcoeu li1n9s92,
TheN eurobioolfoN geyu raNle tworeksd,i tbeyDd a niGealr dn1e9r93,
CopyrighMtaetde rial
TheN eurobioloogfNy e uraNle tworks
edited by
DanieGla rdner
A BradfBoorodk
TheM ITP ress
CambriMdagse,s achusetts
LondoEnn,g land
CopyrighMtaetde rial
© 1993M assachuseItntsst itouftT ee chnology
Allr ighrtess erveNdo. p arotf t hibso okm ay ber eproducienda nyf ormb y anye lec
tronoircm echanicmaela ns( includpihnogt ocopyirnegc,o rdionrgi ,n formatsitoonr age
andr etriewviatlh) oupte rmissiionwn r itifnrgo mt he publisher.
Thisb ookw ass eti nP alatibnyo D anieGla rdnearn dw asp rinteandd b oundi nt he
UniteSdt ateosfA merica.
LibraorfyC ongresCsa taloging-in-PubDlaitcaa tion
I
TheN eurobioloofgn ye uranle tworkse ditebdy DanieGla rdner.
p. cm. - (Computationneaulr oscience)
"A Bradfobrodo k.'·
Includbeisb liograprheifcearle nacnedsi ndex.
ISBN0 -262-07150-9
1.N euracli rcuitr2y.N .e uranle twork(sC omputesrc ience)
3.N eurobiologIy.G. a rdneDra,n ie1l9,4 5- IIS.e ries.
QP363.3.N415959 3
612.8-dc20 93-9125
CIP
CopyrighMtaetde rial
Contents
Contributors
IX
Preface
Xl
1.I ntroductTioowna:r dN euraNle uraNle tworks 1
DanieGla rdner
2.T wo PrincipolfeBs r ainO rganization:
A Challenfgoer A rtificNieaulr aNle tworks 13
CharlFe.sS tevens
3.S tatiDce terminanotfsS ynaptiSct rength 21
DanieGla rdner
4.L earninRgu lesf romN eurobiology 71
DouglAa.s B axtaenrd J ohHn. B yrne
5.R ealistic NeMtowdoerlks o fD istributed Prociens sing
theL eech 107
ShawnR .L ockearnyd TerrJe.Sn ecjel lowski
6.N euraaln dP eripherDayln amicass D eterminanotfs
PatterneMdo torB ehavior 137
HillJe.Cl h iealn dR andaDl.lB eer
7.D ynamicN euraNle tworkM odelso fS ensorimotor
Behavior 165
EberhaEr.dF etz
References 191
Index 219
CopyrighMtaetde rial
SerieFso reword
Computatnieounraols ciisae nan pcper otaouc nhd erstatnhideni fnogr
maticoonn toefnn te ursailg nbaylm so deltihnnege rvosuyss taetm
mandyi ffesrternutc stcuarlaielns c,l utdhibeni go phystihcceia rlc,u it,
andt hsey stelmesv eClosm.p utseirm ulaotfin oenusr oannsdn eural
netwoarrkceso mplemetnott raardyi ttieocnhanlii qnnu eeusr oscience.
Thibso oske riweesl comceosn tribtuhtailtoi nntskh eoresttiucdaile s
witehx perimaepnptraola tcouh nedse rstainndfionrgmp artoicoens sing
int hnee rvosuyss tAerme.a asn dt opiocfps a rticiunltaeirrn ecsltu de
biophymseicchaaln ifsocmros m putaitnni eounr ocnosm,p ustiemru
latioofnn esu rcailr cmuoidtesol,fls e arnrienpgr,e senotfsa etnisoonrsy
informiatnni eounrn aelt worskyss,t emmosd eolfss ensory-imnotor
tegraatnidco onm,p utatainoanlayolsfp i rso blienbm iso losgeincsailn g,
motocro ntraonldp, e rception.
TerreJn.Sc eej nowski
TomasPoo ggio
CopyrighMtaetde rial
Contributors
DouglAa.Bs a xter
DepartmoefNn etu robiaonldoA gnya tomy
TheU niverosfTi etxyaM se dicSaclh ool
HoustToenx,a s
JohHn. B yrne
DepartmoefNn etu robiaonldoA gnya tomy
TheU niverosfTi etxyaM se dicSaclh ool
HoustToenx,a s
HilJl.eC lh iel
DepartmoefBn ito logy
andD epartmoefNn etu roscience
Case WeRsetseerrUnvn ei versity
ClevelOahnido,
RandaDl.Bl e er
DepartmoefCn otm puter EnginSeceireinncge and
andD epartmoefBn ito logy
CasWee steRrens erUvnei versity
ClevelOahnido,
EberhEa.rF de tz
DepartmoefPn hty sioalnodB giyo physics
andR egioPnrailm aRtees eaCrecnht er
UniverosfWi atsyh ington
SeatWtalseh,i ngton
DaniGealr dner
DepartmoefPn hty siology
andD epartmoefNn etu roscience
CornUenlilv erMseidtiycC aoll lege
NewY orNke,w Y ork
CopyrighMtaetde rial
ShawRn.L ockery
ComputatNieounraolb iLoalboogrya tory
TheS alIkn stiftouBrti eo logSitcuadli es
SanD iegCoa,l ifornia
TerreJn.Sc eej nowski
ComputatNieounraolb iLoalboogrya tory
TheS alIkn stiftouBrti eo logSitcuadli es
SanD iegCoa,l ifornia
CharFle.Ss t evens
HowarHdu gheMse dicIanls titute
TheS alIkn stiftouBrti eo logSitcuadile s
SanD iegCoa,l ifornia
x Contributors Copyrighted Material
Preface
TheN eurobioolfoN gye uraNle tworikssi ntenndoeotdn laysa r eference
fosrt udeanntdis n vestibguaatts ao s rpsu,ar n dag uitdoew adredv el
opmenotft hnee xgte neratiotnh-igtrehdne e ratinoenu-rnoaeflt
wormko delIsti. sa s erioefvs a riatoinao s nisn gtlhee mteh:ab to th
artifniecuirnaaellt woarnkdsn eurobiwoilloblge yn ebfyia tp artial
synthoefts hiesdi irs panraattuerI ensd.i vidualw ecrhceao pmtmeirss
sionferdos me lecatuetdhw oirtsth h giosa ilnm indt;hw iosr ikns o tt he
proceedoifan s gpse cmiefeitci onrag n a dh occ olleocftu inorne lated
workEsa.cc hh apetxearm inneeusrn aelt woarsks umptainodmn osd els
froamn eurobiopleorgsipceacpltr ievsee,na tc irnigtr iecvailoe ftw h e
experimeevnitdaeblno ctceho nsiasntdie nncto nswiisttnhee nutrn aelt
worcko mponemnetcsh,a nairscmh,i teacntdcu arpea,b iClhiatpiteesr.s
ardee sigannedwd r itstoet nh arte adwehrosa rfea milwiiatnrhe uro
biolobguytn otn etwomrokd elosrt, h ree vercsaena, p prectihaet e
synthTeose insh.a nrceea dabeialccihht ayp,it sse erl f-conrteasiunletd,
inignp araolrcl oemlp lemevniteaowrfssy i mitloapriF crse.q ucernots s
referebnectewsec ehna pteemrpsh astihzueen iotfyt hteh emaen,da
singcloem binbeidb liogermapphhays tihzueen si otfyp urpose.
Neural nenteweiodnr pkusat n,dt hfei eolfnd e urnaelt wonrekesd s
continiunipnfugrt o bmo tohft htew og roufposwr h omt hivso lume
isi ntenndeeudr:o bioalnodng eitswtosmr okd elers.
Fonre urobiotlhobego iospktr se,s easn etlsf -coanntaaliyonsfeti dhs e
methodosltorgeyn,ag ntdwh esa knesasnedss i,g nifoifca arntcief icial
neural nmeotdweoalrnskdt heaiprp licatboni eluirtoys cBieecnacues.e
nos ingvloel ucmaebn e s uffitchieearnretpe ,l entrieffuelr etnotc hees
neurnaelt wolrikt ercaotnucreen,t ornat thiceno gm putatipoonwally
erfbuulti nadequnaetuerloym ornpehtdisec v elowpietdht ihnle a st
decadTeh.r oughtohupete ,r speicstn ievuer obiolaolcglhi acpatle;r s
havoen eo rm oraeu thowrhsoa rper actiacswi enlgal sp rofessing
neurobioulsoigsniigsn tgsl,e o-riu nntirta creellcuolradvrio nlgt,ao gre -
patch-clanmepuirnogc,h eomrmi oclaelc ubliaorl ogtieccahln iAq ues.
majgoora ilst oe ncournaeguer obiotlopo agritsitcisi ncp oantset ructing
thnee xgte neraotfni eounrn aelt woarskw se,la lst oe xaminneeu robiol
ogicdaalt a afn reotmw oprekr spective.
CopyrighMtaetde rial
Fonre urnaelt womrokd eleesrpse,c tihaolwslieyt hoabu atc kground
int hlei sfcei entchebeso ,o ckr itiecxaalmliyan sepse ocftn se twork
modetlhsaa trl ei kteolb ye bcootmhp utatisoinganlilfayin cdca onmt
paratbool uecr u rrveinetow fr enaelu raonnssdy napasnetdsh nee tworks
thefyo rBme.c anuoss ei ngvloel ucmaebn e s uffitchieearnretpe ,l entiful
referetnotc herese levant ilnin teeurraotbuircoeol nocgeyn,t orna ting
signifdiecvaenlto pmiennn etusr obniaolp hyssyincasp,pt liacs ticity,
andi nterneuronal woirtghtaihnnlei a zsdatet ciaodAnel .t hotuhgeh
perspeicsnt eiuvreo biolaolcglhi acpatlhe,ar vsoe n eo rm oraeu thors
whoh avceo mteon eurobiforlootmgh pyeh ysisccaile nocree nsg ineer
ingo,rw hoh avber oadetnheedpi err spebcyta id veet oiunrtt oh ese
fielAd msa.j ogro ailst oe ncournaeguern aelt womrokd elteorp sa r
ticiipnca otnes trutchnteei xngtge neraotfni eounrn aelt woarskw se,l l
ast oe xaminneet wodreks iagnndp erformfarnoacmm e o rnee urobiol
ogicpaelr spective.
Speciafcikcn owledgemeinntdsi vfiocdlhulaaoplwt a enrdis n,c lude
a larnguem beorfd iveprusbel aincdp rivfautned iangge nctihees :
OffiocfNe a vaRle seatrhcNeha ,t ioInnaslt iotfHu etaelstt hhNe,a tional
InstiotfMu etnet Haela ltthhAe,i Fro rOcfef iocfSe c ienRteisfeiacr ch,
thNea tioSncaileF nocuen datthiHeoo nw,a rHdu ghMeesd ical Institute,
TheB anokf A merica-GFioaunnndianftioi Mroe nd icRaels eaTrhceh ,
MatheFrosu ndatainodtn hC,el eveland AMdavnaunfcaecdtP urroing
gramA.l thoeuagchah g enhcaysa d istmiinscsti tohnse,u ppoeratc h
hapsr ovitdote hdi esm ergfiinegil sadt ribnuototen ltyot huen iver
saloiftt hye isnev estigbauattli stoonot s hv,ei sioofmn a npye erre view
ers and program officers.
Onef inaaclk nowledigseg meenentrtoa t lhb eo oyke,st p ectiofi itcs
editIto hra:Dn ekb oraanhdB ena,n des pecEisatlhlfeyort r,h esiurp port
duritnhgme a nyi teraatnidro encsu rsnieoendtseo dt unteh iwso rtko
thdee sioruetdp puatt tern.
DaniGealr dner
NewY orCki ty
ThanksgDiavyi1,n9 g92
xii Preface CopyrighMtaetde rial
