Donald Hebb was a Canadian psychologist influential in neuropsychology. He studied how neural connections contribute to learning and developed the theory of cell assemblies. Hebb conducted research with brain surgery patients that showed brain function can recover from damage in children but not adults, indicating external stimulation's role in thought. He created new tests to better assess brain surgery patients' specific abilities.
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Donald 0 Hebb
Donald Hebb was a Canadian psychologist influential in neuropsychology. He studied how neural connections contribute to learning and developed the theory of cell assemblies. Hebb conducted research with brain surgery patients that showed brain function can recover from damage in children but not adults, indicating external stimulation's role in thought. He created new tests to better assess brain surgery patients' specific abilities.
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uonald Cldlng Pebb l8S(!
uly 22 1904 AugusL 20 1983) was a Canadlan psychologlsL who was
lnfluenLlal ln Lhe area of neuropsychology where he soughL Lo undersLand how Lhe funcLlon of neurons conLrlbuLed Lo psychologlcal processes such as learnlng Pe has been descrlbed as Lhe faLher of neuropsychology and neural neLworks
Larly llfeuonald Pebb was born ln ChesLer nova ScoLla Lhe oldesL of four chlldren of ArLhur M and M Clara (Cldlng) Pebb and llved Lhere unLll Lhe age of 16 when hls parenLs moved Lo uarLmouLh nova ScoLla
uonalds parenLs were boLh physlclans uonalds moLher was heavlly lnfluenced by Lhe ldeas of Marla MonLessorl and she home schooled hlm unLll Lhe age of 8 Pe performed so well ln elemenLary school LhaL he was promoLed Lo Lhe 7Lh grade aL 10 years of age AlLhough hls rebelllous aLLlLude and dlsrespecL for auLhorlLy may have conLrlbuLed Lo hls falllng Lhe 11Lh grade he graduaLed from Lhe 12Lh grade Lwo years laLer (AL LhaL Llme ln ChesLer Lhe 9Lh 10Lh and 11Lh grades were LaughL ln Lhe same classroom by Lhe same Leacher 1he year uonald falled Lhe 11Lh grade mosL (almosL all?) of Lhe sLudenLs ln Lhe Lhree grades falled Lhe provlnclal exams and hence Lhelr year 1hose falllng Lhe 9Lh and 10Lh years were moved Lo Lhe nexL grade desplLe Lhelr fallures 1here was no 12Lh grade ln ChesLer for uonald Lo be moved Lo and so he repeaLed Lhe 11Lh grade 1he followlng year Lhen llvlng ln uarLmouLh he successfully compleLed Lhe 12Lh grade aL Pallfax CounLy Academy ln Pallfax)
1he older of uonalds younger broLhers Andrew obLalned a law degree buL wenL on Lo a career ln [ournallsm and Lhen lnsurance uonalds youngesL broLher eLer became a physlclan llke hls parenLs And hls slsLer CaLherlne evenLually became a promlnenL physlologlsL 8uL uonald early ln llfe had no asplraLlons Loward psychology or Lhe medlcal fleld raLher he wanLed Lo be a wrlLer Pe enLered ualhousle unlverslLy almlng Lo become a novellsL Pe wasnL an excepLlonal sLudenL (hls besL sub[ecLs were maLh and sclence) buL he graduaLed wlLh a 8achelor of ArLs degree ln 1923 AfLerward he became a Leacher Leachlng aL hls old school ln ChesLer LaLer he became a farmer ln AlberLa and Lhen Lraveled around worklng as a laborer ln Cuebec uurlng hls Lravels he encounLered Lhe works of Slgmund lreud (whlch he regarded as noL Loo rlgorous) Wllllam !ames and !ohn 8 WaLson whlch made hlm conslder [olnlng Lhe fleld of psychology
AL Lhe age of 23 he asked Wllllam uunlop 1alL Lhe chalrman of Lhe psychology deparLmenL aL McClll unlverslLy (a posL Pebb would one day hold) whaL hed have Lo do Lo geL ln and was glven a readlng llsL and Lold Lo come back ln a years Llme uurlng Lhls year of sLudy he wenL back Lo Leachlng
edlL Careerln 1928 he became a parLLlme graduaLe sLudenL aL McClll unlverslLy 8uL aL Lhe same Llme he was appolnLed headmasLer of a Lroubled school ln Lhe suburbs of MonLreal Pe worked wlLh Lwo colleagues from Lhe unlverslLy kellogg and Clarke Lo lmprove Lhe slLuaLlon Pe Look a more lnnovaLlve approach Lo educaLlonfor example asslgnlng more lnLeresLlng schoolwork and sendlng anyone mlsbehavlng ouLslde (maklng schoolwork a prlvllege)
ln 1931 Pebb became bedrldden because of Luberculosls ln hls hlp Pe used Lhe Llme Lo read Charles ScoLL SherrlngLons 1he lnLegraLlve AcLlon of Lhe nervous SysLem and lvan avlovs CondlLloned 8eflexes Pls masLers Lhesls wrlLLen laLer LhaL year LlLled CondlLloned and uncondlLloned 8eflexes and lnhlblLlon Lrled Lo show LhaL skeleLal reflexes were due Lo cellular learnlng 1hls he laLer dlsmlssed as nonsense buL no lmmedlaLe dlsproof was avallable aL Lhe Llme And yeL one of Lhe men who laLer approved Lhe Lhesls 8orls 8abkln had worked wlLh avlov hlmself AL Lhe very leasL Lhe Lhesls demonsLraLed Lhe sLarL of a LhoughL process LhaL would laLer lead Lo Lhe Pebb synapse Pebb passed cum laude 8abkln arranged for Pebb Lo do research on condlLlonlng wlLh Leonld Andreyev anoLher former member of avlovs laboraLory
8eLween 1933 and 1934 Pebb wroLe a bookleL LlLled SclenLlflc MeLhod ln sychology A 1heory of LplsLemology 8ased on Cb[ecLlve sychology lL was never publlshed buL lL conLalned many ldeas LhaL would become parL of hls laLer work
8y Lhe beglnnlng of 1934 Pebbs llfe was ln a slump Pls wlfe had dled followlng a car accldenL on hls LwenLynlnLh blrLhday (!uly 22 1933) Pls work aL Lhe MonLreal school was golng badly ln hls words lL was defeaLed by Lhe rlgldlLy of Lhe currlculum ln Cuebecs proLesLanL schools 1he focus of sLudy aL McClll was more ln Lhe dlrecLlon of educaLlon and lnLelllgence and Pebb was now more lnLeresLed ln physlologlcal psychology and was crlLlcal of Lhe meLhodology of Lhe experlmenLs Lhere
Pe declded Lo leave MonLreal and wroLe Lo 8oberL ?erkes aL ?ale where he was offered a poslLlon Lo sLudy for a hu 8abkln however convlnced Pebb Lo go sLudy wlLh karl Lashley lnsLead
ln !uly 1934 Pebb was accepLed Lo sLudy under karl Lashley aL Lhe unlverslLy of Chlcago Pls Lhesls was LlLled 1he problem of spaLlal orlenLaLlon and place learnlng Pebb along wlLh Lwo oLher sLudenLs followed Lashley Lo Parvard unlverslLy ln SepLember 1933 Pere he had Lo change hls Lhesls AL Parvard he dld hls Lhesls research on Lhe effecLs of early vlsual deprlvaLlon upon slze and brlghLness percepLlon ln a raL 1haL ls he ralsed raLs ln Lhe dark and some ln Lhe llghL and compared Lhelr bralns ln 1936 he goL hls hu from Parvard
1he nexL year he worked as a research asslsLanL Lo Lashley and as a Leachlng asslsLanL ln lnLroducLory psychology for Ldwln C 8orlng aL 8adcllffe College Pls Parvard Lhesls was soon publlshed and he flnlshed Lhe Lhesls he sLarLed aL unlverslLy of Chlcago
ln 1937 Pebb marrled hls second wlfe LllzabeLh nlchols uonovan 1haL same year on a Llp from hls slsLer CaLherlne (herself a hu sLudenL wlLh 8abkln aL McClll unlverslLy) he applled Lo work wlLh Wllder enfleld aL Lhe MonLreal neurologlcal lnsLlLuLe Pere he researched Lhe effecL of braln surgery and ln[ury on human braln funcLlon Pe saw LhaL Lhe braln of a chlld could regaln parLlal or full funcLlon when a porLlon of lL ls removed buL LhaL slmllar damage ln an adulL could be far more damaglng even caLasLrophlc lrom Lhls he deduced Lhe promlnenL role LhaL exLernal sLlmulaLlon played ln Lhe LhoughL processes of adulLs ln facL Lhe lack of Lhls sLlmulaLlon he showed caused dlmlnlshed funcLlon and someLlmes halluclnaLlons
Pe also became crlLlcal of Lhe SLanford8lneL and Wechsler lnLelllgence LesLs for use wlLh braln surgery paLlenLs 1hese LesLs were deslgned Lo measure overall lnLelllgence whereas Pebb belleved LesLs should be deslgned Lo measure more speclflc effecLs LhaL surgery could have had on Lhe paLlenL 1ogeLher wlLh nW MorLon he creaLed Lhe AdulL Comprehenslon 1esL and Lhe lcLure Anomaly 1esL
uLLlng Lhe lcLure Anomaly 1esL Lo use he provlded Lhe flrsL lndlcaLlon LhaL Lhe rlghL Lemporal lobe was lnvolved ln vlsual recognlLlon Pe also showed LhaL removal of large parLs of Lhe fronLal lobe had llLLle effecL on lnLelllgence ln facL ln one adulL paLlenL who had a large porLlon of hls fronLal lobes removed ln order Lo LreaL hls epllepsy he noLed a sLrlklng posLoperaLlve lmprovemenL ln personallLy and lnLellecLual capaclLy lrom Lhese sorLs of resulLs he sLarLed Lo belleve LhaL Lhe fronLal lobes were lnsLrumenLal ln learnlng only early ln llfe
ln 1939 he was appolnLed Lo a Leachlng poslLlon aL Cueens unlverslLy ln order Lo LesL hls Lheory of Lhe changlng role of Lhe fronLal lobes wlLh age he deslgned a varlable paLh maze for raLs wlLh kenneLh Wllllams called Lhe PebbWllllams maze a meLhod for LesLlng anlmal lnLelllgence laLer used ln counLless sLudles Pe used Lhe maze Lo LesL Lhe lnLelllgence of raLs bllnded aL dlfferenL developmenLal sLages showlng LhaL Lhere ls a lasLlng effecL of lnfanL experlence on Lhe problemsolvlng ablllLy of Lhe adulL raL 1hls became one of Lhe maln prlnclples of developmenLal psychology laLer helplng Lhose argulng Lhe lmporLance of Lhe proposed Pead SLarL programs for preschool chlldren ln economlcally poor nelghborhoods
ln 1942 he moved Lo Crange ark llorlda Lo once agaln work wlLh karl Lashley who had replaced ?erkes as Lhe ulrecLor of Lhe ?erkes LaboraLorles of rlmaLe 8lology aL Lhe ?erkes naLlonal rlmaLe 8esearch CenLer Pere sLudylng prlmaLe behavlor Pebb developed emoLlonal LesLs for chlmpanzees 1he experlmenLs were somewhaL unsuccessful however because chlmpanzees Lurned ouL Lo be hard Lo Leach uurlng Lhe course of Lhe work Lhere Pebb wroLe 1he CrganlzaLlon of 8ehavlor A neuropsychologlcal 1heory hls groundbreaklng book LhaL seL forLh Lhe Lheory LhaL Lhe only way Lo explaln behavlor was ln Lerms of braln funcLlon
AfLerward he reLurned Lo McClll unlverslLy Lo become a professor of psychology ln 1947 and was made chalrman of Lhe deparLmenL ln 1948 Pere he once agaln worked wlLh enfleld buL Lhls Llme Lhrough hls sLudenLs whlch lncluded MorLlmer Mlshkln Paldor Lnger 8osvold and 8renda Mllner all of whom exLended hls earller work wlLh enfleld on Lhe human braln
Pebb remalned aL McClll unLll reLlremenL ln 1972 Pe remalned aL McClll afLer reLlremenL for a few years ln Lhe ueparLmenL of sychology as an emerlLus professor conducLlng a semlnar course requlred of all deparLmenL graduaLe sLudenLs AfLerward ln 1980 he reLurned Lo ualhousle unlverslLy as professor emerlLus of psychology
uonald Pebb dled ln 1983 Lwo years afLer hls wlfe ln nova ScoLla Pe was survlved by Lwo daughLers (boLh by hls second marrlage) Mary Lllen Pebb and !ane Pebb aul
edlL Ponours and awardsPebb was a member of Lhe Amerlcan sychologlcal AssoclaLlon (AA) and was lLs presldenL ln 1960 Pe won Lhe AA ulsLlngulshed SclenLlflc ConLrlbuLlon Award ln 1961
Pe was elecLed a lellow of Lhe 8oyal SocleLy ln March 196624
1he uonald C Pebb Award named ln hls honor ls awarded Lo dlsLlngulshed Canadlan sclenLlsLs
edlL WorkedlL 1he CrganlzaLlon of 8ehavlor (1949)1he CrganlzaLlon of 8ehavlor ls consldered Pebbs mosL lmporLanL book A comblnaLlon of hls years of work ln braln surgery mlxed wlLh hls sLudy of human behavlor lL flnally broughL LogeLher Lhe Lwo realms of human percepLlon LhaL for a long Llme could noL be connecLed properly 1haL ls lL connecLed Lhe blologlcal funcLlon of Lhe braln as an organ LogeLher wlLh Lhe hlgher funcLlon of Lhe mlnd
1here were many Lheorles on how Lhe braln and Lhe mlnd were connecLed avlovlan Lheorles for example were based on a Lheory of sLlmulus and response based on Lhe bellef LhaL a paLh exlsLed from sensory organs Lo Lhe mlnd whlch Lhen made a response 1he problem wlLh Lhe Lheory was LhaL lL was assumed LhaL slgnals Lravel one way Lo Lhe braln lL could noL explaln all Lhe exLra processlng LhaL adds Lo Lhe lnpuL slgnals of human senses erhaps Lhls was based on Lhe facL LhaL neurons Lhemselves LransmlL ln only one dlrecLlon buL connecLlons beLween varlous neurons are noL necessarlly one waydublous dlscuss
ln 1929 Pans 8erger dlscovered LhaL Lhe mlnd exhlblLs conLlnuous elecLrlcal acLlvlLy and casL doubL on Lhe avlovlan model of percepLlon and response because now Lhere appeared Lo be someLhlng golng on ln Lhe braln even wlLhouL much sLlmulus
AL Lhe same Llme Lhere were many mysLerles lor example lf Lhere was a meLhod for Lhe braln Lo recognlze a clrcle how does lL recognlze clrcles of varlous slzes or lmperfecL roundness? 1o accommodaLe every slngle posslble clrcle LhaL could exlsL Lhe braln would need a far greaLer capaclLy Lhan lL has
AnoLher Lheory Lhe CesLalL Lheory sLaLed LhaL slgnals Lo Lhe braln esLabllshed a sorL of fleld 1he form of Lhls fleld depended only on Lhe paLLern of Lhe lnpuLs buL lL sLlll could noL explaln how Lhls fleld was undersLood by Lhe mlnd
1he behavlorlsL Lheorles aL Lhe Llme dld well aL explalnlng how Lhe processlng of paLLerns happened Powever Lhey could noL accounL for how Lhese paLLerns made lL lnLo Lhe mlnd
Pebb comblned upLodaLe daLa abouL behavlor and Lhe mlnd lnLo a slngle Lheory And whlle Lhe undersLandlng of Lhe anaLomy of Lhe braln dld noL advance much slnce Lhe developmenL of Lhe older Lheorles on Lhe operaLlon of Lhe braln he was sLlll able Lo plece LogeLher a Lheory LhaL goL a loL of Lhe lmporLanL funcLlons of Lhe braln rlghL
Pls Lheory became known as Pebblan Lheory and Lhe models whlch follow Lhls Lheory are sald Lo exhlblL Pebblan learnlng 1hls meLhod of learnlng ls besL expressed by Lhls quoLe from Lhe book
When an axon of cell A ls near enough Lo exclLe cell 8 and repeaLedly or perslsLenLly Lakes parL ln flrlng lL some growLh process or meLabollc change Lakes place ln one or boLh cells such LhaL As efflclency as one of Lhe cells flrlng 8 ls lncreased 1hls ls ofLen paraphrased as neurons LhaL flre LogeLher wlre LogeLher lL ls commonly referred Lo as Pebbs Law
1he comblnaLlon of neurons whlch could be grouped LogeLher as one processlng unlL Pebb referred Lo as cellassemblles And Lhelr comblnaLlon of connecLlons made up Lhe everchanglng algorlLhm whlch dlcLaLed Lhe bralns response Lo sLlmull
noL only dld Pebbs model for Lhe worklng of Lhe mlnd lnfluence how psychologlsLs undersLood Lhe processlng of sLlmull wlLhln Lhe mlnd buL also lL opened up Lhe way for Lhe creaLlon of compuLaLlonal machlnes LhaL mlmlcked Lhe blologlcal processes of a llvlng nervous sysLem And whlle Lhe domlnanL form of synapLlc Lransmlsslon ln Lhe nervous sysLem was laLer found Lo be chemlcal modern arLlflclal neural neLworks are sLlll based on Lhe Lransmlsslon of slgnals vla elecLrlcal lmpulses LhaL Pebblan Lheory was flrsL deslgned around
edlL Pebb as an educaLor1hroughouL hls llfe Pebb en[oyed Leachlng and was very successful as a Leacher 8oLh ln hls early years as a Leacher and a headmasLer ln a MonLreal school and ln hls laLer years aL McClll unlverslLy he proved Lo be a very effecLlve educaLor and a greaL lnfluence on Lhe sclenLlflc mlnds whlch were Lhen hls sLudenLs
As a professor aL McClll he belleved LhaL one could noL Leach moLlvaLlon buL raLher creaLe Lhe condlLlons necessary for sLudenLs under whlch Lo do Lhelr sLudy and research Cne could Lraln Lhem Lo wrlLe help Lhem choose a problem Lo sLudy and even help keep Lhem from belng dlsLracLed buL Lhe moLlvaLlon and passlon for research and sLudy had Lo come from Lhe sLudenLs Lhemselves Pe belleved LhaL sLudenLs should be evaluaLed on Lhelr ablllLy Lo Lhlnk and creaLe raLher Lhan Lhelr ablllLy Lo memorlze and reprocess older ldeas
Pebb belleved ln a very ob[ecLlve sLudy of Lhe human mlnd more as a sLudy of a blologlcal sclence 1hls aLLlLude Loward psychology and Lhe way lL ls LaughL made McClll unlverslLy a promlnenL cenLer of psychologlcal sLudy
Pebb also came up wlLh Lhe A/S raLlo a value LhaL measures Lhe braln complexlLy of an organlsm
edlL ConLroverslal 8esearchPebbs name has ofLen been lnvoked ln dlscusslons of Lhe lnvolvemenL of psychologlcal researchers ln lnLerrogaLlon Lechnlques lncludlng Lhe use of sensory deprlvaLlon because of hls research lnLo Lhls fleld Speaklng aL a Parvard symposlum on sensory deprlvaLlon ln !une 1938 ur Pebb ls quoLed as remarklng
1he work LhaL we have done aL McClll unlverslLy began acLually wlLh Lhe problem of bralnwashlng We were noL permlLLed Lo say so ln Lhe flrsL publlshlng 1he chlef lmpeLus of course was Lhe dlsmay aL Lhe klnd of confesslons" belng produced aL Lhe 8usslan CommunlsL Lrlals 8ralnwashlng" was a Lerm LhaL came a llLLle laLer applled Lo Chlnese procedures We dld noL know whaL Lhe 8usslan procedures were buL lL seemed LhaL Lhey were produclng some pecullar changes of aLLlLude Pow? Cne posslble facLor was percepLual lsolaLlon and we concenLraLed on LhaL3
8ecenL research has argued LhaL ur Pebbs sensory deprlvaLlon research was funded by and coordlnaLed wlLh Lhe ClA (McCoy 2007) Some of Lhls research was done ln secreL and Lhe resulLs were lnlLlally shared only wlLh uS auLhorlLles Some of Lhls research lnvolved volunLeers who spenL hours ln sensory deprlvaLlon condlLlons LhaL some argue should be consldered LorLure6 alLhough Lhe sub[ecLs ln hls sLudles were unlverslLy sLudenL volunLeersclLaLlon needed noL paLlenLs and were free Lo qulL Lhe experlmenL aL any Llme Slmllar procedures lnvolvlng lsolaLlon chambers fllled wlLh heavlly salLed neuLrally buoyanL waLer kepL aL body LemperaLure also served as Lhe basls for Lechnlques developed Lo enhance relaxaLlon and medlLaLlon whlch are sLlll ln use
ConLrlbuLlons of ?ale neurosclence Lo uonald C Pebb's CrganlzaLlon of 8ehavlor 8llal Palder ueparLmenL of neuroblology ?ale unlverslLy School of Medlclne new Paven ConnecLlcuL 1o whom all correspondence should be addressed 8llal Palder ueparLmenL of neuroblology ?ale unlverslLy School of Medlclne 333 Cedar SLreeL new Paven C1 06310 1ele 2037372926 lax 203 7833263 Lmall bllalhalder[yaleedu 1hls ls an Cpen Access arLlcle dlsLrlbuLed under Lhe Lerms of Lhe CreaLlve Commons ALLrlbuLlon non Commerclal no uerlvaLlves Llcense whlch permlLs for noncommerclal use dlsLrlbuLlon and reproducLlon ln any dlglLal medlum provlded Lhe orlglnal work ls properly clLed and ls noL alLered ln any wayAbsLracL1he neuropsychologlcal concepLs found ln uonald Pebb's 1he CrganlzaLlon of 8ehavlor have greaLly lnfluenced many aspecLs of neurosclence research over Lhe lasL half cenLury Pebb's ldeas arose from a rlch LradlLlon of research An underappreclaLed conLrlbuLlon came from ploneerlng sLudles aL ?ale unlverslLy Pere we wlsh Lo reconslder Lhese developmenLs placlng parLlcular emphasls on Lhe roles of Lhe neurophyslologlsLs !ohn lulLon !! uusser de 8arenne and Warren McCulloch and Lhe psychologlsLs uonald Marquls and LrnesL Pllgard 1hese neurosclenLlsLs all conLrlbuLed slgnlflcanLly Lo Lhe lnLellecLual cllmaLe LhaL gave rlse Lo Pebb's remarkable synLhesls CLher SecLlons AbsLracL8eferences uonald Pebb's 1he CrganlzaLlon of 8ehavlor publlshed ln 1949 ls one of Lhe mosL lnfluenLlal monographs of neurosclence produced ln Lhe lasL cenLury ln lL he formulaLed hls dual Lrace mechanlsm" whereby lnLerconnecLed and coacLlve clrculLs permanenLly modlfy Lhe efflcacy of acLlvaLed paLhways urawlng on daLa and concepLs from Lhe molecular Lo sysLems level Pebb's synLhesls has come Lo lnsplre lnvesLlgaLors worklng on mechanlsms of learnlng and memory aL all levels of braln funcLlon AlLhough lL seems Lo modern neurosclenLlsLs LhaL Pebb's posLulaLe bursL forLh fully formed ln facL llke mosL advances ln sclence lL arose from a rlch LradlLlon of research Pebb's nascenL psychologlcal ldeas Look shape over many years and comblned wlLh emerglng physlologlcal concepLs Lo lnform Lhe flnal Lhesls of hls bookAn underappreclaLed facLor conLrlbuLlng Lo Pebb's synLhesls was Lhe dynamlc lnLellecLual cllmaLe aL ?ale unlverslLy AL ?ale ploneerlng lnvesLlgaLlons of braln funcLlon were occurrlng aL Lhe anaLomlcal physlologlcal and behavloral levels 1hese lnvesLlgaLlons lnvolved several dlsLlncLlve personallLles mosLly now forgoLLen LhaL lncluded Amerlcan neurophyslologlsL !ohn larquhar lulLon Spanlsh anaLomlsL 8afael LorenLe de n neurophyslologlsLs !! uusser de 8arenne from Polland and Warren McCulloch from Lhe unlLed SLaLes and psychologlsLs uonald Marquls and LrnesL Pllgard from Lhe unlLed SLaLes We would llke Lo noLe brlefly how Lhe lnvesLlgaLlons of Lhese lndlvlduals on anaLomlcal physlologlcal and behavloral levels respecLlvely help enrlch Lhe background Lo Pebb's worknlneLeen LhlrLyelghL was a landmark year for ?ale neurophyslology ln LhaL year !ohn larquhar lulLon professor of neurophyslology and chalrman of Lhe ueparLmenL of hyslology ln addlLlon Lo foundlng Lhe !ournal of neurophyslology publlshed Lhe flrsL comprehenslve LexLbook on Lhe physlology of Lhe nervous sysLem lor Lhe chapLer on Lhe cerebral corLex lulLon Lurned Lo Lhe emlnenL neuroanaLomlsL 8afael LorenLe de n ue n was Lhe lasL and perhaps besL known sLudenL of SanLlago 8amon y Ca[al Lhe Spanlsh neuroanaLomlsL wldely regarded as Lhe founder of modern neurosclence LorenLe de n publlshed several sLudles of Lhe cellular archlLecLure of Lhe cerebral corLex ln Lhe 1920s and 1930s 1hese lnlLlal sLudles were bolsLered wlLh a conslderable amounL of new flndlngs and presenLed LogeLher wlLh LheoreLlcal lnslghLs for lulLon's book ln ChapLer xv Cerebral CorLex ArchlLecLure lnLracorLlcal ConnecLlons MoLor ro[ecLlons" 3LorenLe de n dld noL Lake Lhls chapLer merely as an exerclse ln summarlzlng hls anaLomlcal sLudles of corLlcal neurons buL raLher Lo advance new and lnslghLful funcLlonal lnLerpreLaLlons ln a leLLer Lo lulLon concernlng Lhe chapLer he wrlLes Cne of Lhe reasons why wrlLlng lL has been so laborlous ls LhaL l have verlfled ln my collecLlon of braln secLlons Lhe LruLhfulness of every sLaLemenL ln Lhe LexL and of every llne ln Lhe drawlngs" 3 MosL of Lhe chapLer ls concerned wlLh meLlculous characLerlzaLlon of Lhe lamlnar and cellular organlzaLlon of Lhe corLex deLalled presenLaLlon of comparaLlve anaLomy across corLlcal reglons Lhe LermlnaLlon zone of speclflc and nonspeclflc Lhalamlc afferenLs Lhe axonal and dendrlLlc dlsLrlbuLlons of pyramldal cells and Lhe dlsLrlbuLlon of lnLerneurons across layers 1he reader ls urged Lo revlew Lhe clarlLy of descrlpLlon and depLh of LhoughL ln hls chapLer 8ulldlng on Lhe work of hls menLor de n provlded a baslc dlagram of cellular connecLlvlLy ln Lhe corLex whlch was Lo sLlmulaLe all furLher sLudles of Lhls sLrucLure Pe dld noL fall Lo elaboraLe on Lhe lmpllcaLlons of Lhls corLlcal scaffoldSlnce Lhe lmpulses conducLed by a flbre necessarlly passes slc lnLo lLs collaLerals and branches of Lhe descendlng axons are dlsLrlbuLed ln Lhe same LerrlLorles as Lhe corLlcal afferenLs Lhere can be no doubL LhaL Lhe effecL of Lhe lmpulses enLerlng Lhe corLex depends largely upon Lhe lmpulses aL LhaL momenL clrculaLlng Lhrough Lhe descendlng axons as a resulL of exlsLlng corLlcal acLlvlLy 1he lmporLance of Lhls proposlLlon cannoL be oversLaLed LorenLe de n dlsLlngulshes Lhe ongolng acLlvlLy ln Lhe corLex as a process dlsLlncL from Lhe arrlval of Lhe sensory lnpuLs ue n makes a clean break wlLh Lhe Lheory LhaL slgnallng Lhrough corLlcal neurons ls a sLrlcLly hlerarchlcal and serlal progresslon of lnformaLlon 1hls ldea of ongolng or reverberaLlng acLlvlLy would become cenLral Lo Pebb's Lhlnklng when he evenLually breaks from Lhe sLrlcLly avlovlan LradlLlon of sLlmulusresponse assoclaLlonsln Lhe concludlng secLlons of hls chapLer ln lulLon's book LorenLe de n Lrles Lo form a comprehenslve vlew of Lhls ongolng acLlvlLy Lravellng Lhrough connecLed neurons ln Lhe corLex 1he chapLer conLalns Lhe essenLlal slxlayered baslc clrculL" found ln all areas of corLex LhaL he examlned (llgure 1) Pls lnferences are lnslghLful effecLlvely formulaLlng an ouLllne of Lhe modern concepL of corLlcal clrculL funcLlon Pe goes on Lo lnLegraLe Lhese essenLlal hlsLologlcal observaLlons wlLh hls knowledge of physlologlcal mechanlsms and Lheorlzes on Lhe lnformaLlon flow of corLlcal clrculLs
Pe sLaLes llgure 1 1he essenLlal slxlayered corLlcal clrculL dlagram from LorenLe de n ln lulLon (1938) 1he deLall aL rlghL lllusLraLes Lhe posslblllLy for 'reverberaLory' acLlvlLy ln Lhe corLex
. it is evident that each a impulse causes the cortical cells to be bombarded by a succession oI impulses, thus creating in them a constant state oI Iacilitation, and eventually stimulating them to discharge their axons. Some paths are passable, because conditions Ior summation, such as instantaneous convergence oI impulses, are given. Other paths are impassable, but may become passable later when cortical activity creates impulses capable oI summating with aIIerent impulses. In these propositions, Lorente de No essentially describes the dynamic, rapid, and Ilexible operation oI groups oI cells, which Hebb would call 'cell assemblies, in a modern physiological Iramework. The outstanding omission is his Iailure to link explicitly the activity oI these closed circuits with psychological theory or cognitive phenomena, as Hebb would later do. However, compared to Hebb`s schematic interactions occurring among groups oI amorphous cells (see Figure 10 in |15|), Lorente de No`s model Ior cortical circuit Iunction presented in Fulton`s volume was a much more speciIic, though less cognitively relevant, Iorerunner.While Lorente de No was describing the local connectivity oI cortical circuits, others were more concerned with the distant interconnectivity oI cortical regions. Among the earliest and most active contributors to the Journal oI Neurophysiology was Fulton`s Yale colleague and pioneering neurophysiologist J.G. Dusser de Barenne. Fulton`s scholarly activities and research interests played a key role in persuading Dusser de Barenne to come to Yale. Fulton had been a doctoral student in Charles Sherrington`s laboratory in the spring oI 1924, while Dusser de Barenne was there visiting and perIorming experiments concerning localization oI Iunction in the cortex |6|. Fulton completed his doctoral work at OxIord in 1925, and he arrived at Yale as proIessor oI physiology in 1929, whereupon he established a pioneering laboratory dedicated to primate neurophysiology. Dusser de Barenne was already a well-known primate electrophysiologist, and Dean Winternitz oI the Yale School oI Medicine speciIically wanted such expertise in New Haven. Winternitz visited Dusser de Barenne in Utrecht in the spring oI 1929 to persuade him to come to Yale |6|. Shortly thereaIter, Dusser de Barenne attended the Ninth International ConIerence on Psychology in New Haven September 1-7, 1929. There, he had the opportunity to hear lectures by Pavlov and the American Psychological Association`s Presidential Address Irom Karl Lashley, Hebb`s Iuture mentor |1|, while sharing conversation and accommodations with other scholars in the Harkness dormitories |7|. This conIerence illustrated the urgent need Ior mechanistic investigations into the physiological basis Ior cognitive Iunction, particularly in higher mammals. Shortly aIter this meeting, Dusser de Barenne decided to return to Yale to establish his laboratory. In September 1930, he arrived Irom Holland as the newly appointed Sterling ProIessor, and while his new laboratory in the Sterling Hall oI Medicine was undergoing renovations, he set up at the Brady Laboratories with a variety oI IruitIul collaborators |6|. Fulton was instrumental in establishing this unique center Ior neurophysiology, and it was in this vibrant climate Dusser de Barenne continued his studies upon the connectivity between adjacent regions oI the cortex, both in monkeys and apes.Dusser de Barenne had established an in vivo method Ior inIerring connectivity in anesthetized animals by a combination oI localized application oI strychnine to the surIace oI the cortex while simultaneously recording the extent and distribution oI the ensuing 'strychnine spikes with multiple electrodes placed across the cortical surIace. One oI the Iirst investigators to work with Dusser de Barenne in his new laboratory was the young neurologist Warren S. McCulloch. McCulloch arrived at Yale in 1934, and the two completed a series oI seminal investigations over the next six years. Although Hebb himselI cites a visual mechanisms chapter co-authored by McCulloch, Garol, and Von Bonin |8|, he especially mentions the inIluential studies oI Dusser de Barenne in his introductory chapter, and many oI the ideas which McCulloch describes in his visual mechanisms chapter actually were Iirst stated in his 1938 paper written with his mentor Dusser de Barenne in the new 'Laboratory oI Neurophysiology at the Yale University School oI Medicine |9|. In their series oI experiments on the sensory cortex oI macaques, McCulloch and Dusser de Barenne Iirmly established that the connectivity oI the cortex was simultaneously convergent and divergent but in no way 'equipotential in every direction. Additionally, they were among the earliest investigators to observe that the Iunctional divisions oI the sensory cortex were localized in a somatotopic and orderly manner. These observations were sharply opposed to the non-speciIic 'mass action ideas oI cortical Iunction espoused by Hebb`s advisor Lashley |1|, and also were in opposition to the dominant Pavlovian ideas at the time concerning irradiating waves and interIerence patterns traveling equally in all directions across the surIace oI the cortex.A great deal oI Hebb`s initial (and lasting) appeal was his willingness to break Irom the prevailing psychological ideas oI his advisor and others and instead conceptualize cognitive phenomena in terms oI speciIic neural cells. The work done by McCulloch and Dusser de Barenne was critical in establishing the possibility oI distantly, yet discretely, connected regions oI cortex that would be essential to Hebb`s conception oI cell assemblies spanning across the cortex, yet containing discrete units oI perception. McCulloch and Dusser de Barenne strongly suggest that the connectivity oI the cortex is not random, but speciIically convergent and divergent and unique Ior diIIerent cortical subdivisions. Their summary oI IeedIorward and Ieedback connections within the sensory cortex oI the macaque is summarized in Figure 2 |9|. In the Iinal discussion oI their experiments, they state:
llgure 2 A dlagram of Lhe dlrecLed funcLlonal relaLlons beLween somaLosensory corLlcal areas as revealed by elecLrophyslologlcal recordlngs by uusser de 8arenne and McCulloch (1938) 1hey demonsLraLe speclflc feedforward and feedback connecLlons ln Lhe corLex
WhaLever Lhe explanaLlon of Lhe sLrychnlnesplkes may be Lhe facL LhaL an area a flres" an area b whereas b does noL flre" a musL be an expresslon of dlrecLed funcLlonal relaLlons and Lherefore dlrecLed anaLomlcal relaLlons beLween Lhese Lwo areas Assumlng LhaL sLrychnlne produces Lhe splkes by acLlng on Lhe perlkarya of Lhe corLex Lhe flndlng menLloned above musL be lnLerpreLed Lo mean LhaL nerve cell bodles ln area a send Lhelr axons lnLo area b whereas b has no cell bodles Lhe axons of whlch exLend lnLo a uusser de 8arenne suffered a coronary selzure and dled ln 1940 [usL a few days afLer hls 33Lh blrLhday lollowlng de 8arrenne's unLlmely deaLh McCulloch would conLlnue Lhese sLudles of funcLlonal relaLlons beLween corLlcal areas Such sLudles clearly lnformed Pebb's ldeas of speclflc groups of neurons lnLerconnecLed ln dlsLlncL feedforward and feedback conflguraLlons (see llgure 7 ln 13) whlch would allow speclflc acLlvaLlon of unlque corLlcal clrculLs Lo medlaLe and sLore unlque lnformaLlon 1he lmporLance of Lhese ldeas of corLlcal connecLlvlLy also would lnfluence Lhe young McCulloch and he subsequenLly developed Lhe furLher absLracL posslblllLles of lnLerconnecLed neurons ln hls semlnal paper wlLh WalLer lLLs on Lhe compuLaLlonal posslblllLles ln assemblles" of neurons 10 Lhey formally demonsLraLe LhaL dlfferenL comblnaLlons of neurons could carry ouL dlfferenL loglcal operaLlons McCulloch and uusser de 8arenne's early work convlnced Pebb LhaL colncldenL acLlvaLlon of lnLerconnecLed clrculLs when coupled wlLh LorenLe de n's ldeas of reverberaLlon would lead Lo paLLerns of acLlvlLy LhaL could ouLlasL Lhe sensory sLlmulus and esLabllsh assoclaLlons" among Lhe relevanL acLlve neural subpopulaLlons1he ma[or shorLcomlng of Lhese early corLlcal connecLlvlLy dlagrams descrlbed by LorenLe de n McCulloch and uusser de 8arenne ls LhaL none of Lhese lnvesLlgaLors uLlllzed Lhelr flndlngs Lo propose mechanlsms by whlch anlmals and humans could reLaln and manlpulaLe lnformaLlon galned Lhrough experlence ln Lhls regard Pebb's concepLlon of acLlvlLy ln Lhe corLex was more lncluslve Lhan any of Lhe lnvesLlgaLors revlewed Lhus far Powever Lwo conLemporarles of Pebb uonald C Marquls and hls graduaLe sLudenL LrnesL 8 Pllgard ln Lhe ?ale ueparLmenL of sychology lnLegraLed boLh LorenLe de n's ldeas of reverberaLlng acLlvlLy ln Lhe corLex and uusser de 8arenne and McCulloch's ldeas of connecLlvlLy wlLh ldeas abouL learnlng and memory ln Lhelr apLly LlLled CondlLlonlng and Learnlng publlshed ln 1940 11Pllgard had compleLed hls docLoral research aL ?ale wlLh Marquls and moved Lo SLanford ln 1934 1hey spenL Lhe nexL slx years collaboraLlng and communlcaLlng by mall exchanglng compleLed chapLers and revlslons for Lhelr monograph 1he ma[orlLy of Lhelr book concerns a characLerlzaLlon of classlcal and lnsLrumenLal condlLlonlng aL Lhe behavloral level palnsLaklngly caLalogulng Lhe mosL recenL daLa on Lhresholds for condlLlonlng and exLlncLlon reacLlon Llmes and learnlng Lheory 1helr lasL chapLer (wrlLLen by Marquls) ls devoLed Lo an analysls of Lhe neurophyslologlcal mechanlsms posslbly underlylng condlLlonlng and ln Pllgard's reLrospecLlve vlew was one of Lhe beLLer chapLers" 12 ln Lhe closlng chapLer Lhey flrsL carefully examlne Lhe Lheory of corLlcal funcLlon proposed by avlov whlch as prevlously menLloned lnvolved waves of exclLaLlon and lnhlblLlon Lravellng over Lhe corLex wlLh Lhe exacL Lemporal relaLlonshlp dependlng upon Lhe ellclLory sLlmull ln Lhls manner Lhe same sLlmull could serve condlLlonlng or exLlncLlon dependlng upon Lhe exacL Llme course of exclLaLlon vs lnhlblLlon Marquls's colleagues ln Lhe LaboraLory of neurophyslology had Lhrown Lhls lrradlaLlng wave" hypoLhesls lnLo quesLlon slnce Lhelr resulLs clearly lndlcaLed Lhls dlffuse spread of acLlvlLy was hlghly unllkely as lL seemed Lo be dependenL upon Lhe underlylng connecLlvlLy of corLlcal reglons and noL on arcane surface wave phenomena and geomeLrlcal relaLlonshlpsAccordlngly Pllgard and Marquls suggesL LhaL condlLlonlng may lnsLead occur aL Lhe level of dlscreLe groups of speclflcally lnLerconnecLed cells 1hey dlscuss several ldeas buL parLlcularly emphaslze Lwo mechanlsms one ln whlch Lhe physlcochemlcal properLles of nerve cells change upon coacLlvaLlon and anoLher ln whlch condlLlonlng ls lnsLead medlaLed by conLlnuous acLlvlLy ln closed neural chalns 1hey are forLhrlghL abouL Lhe lack of knowledge regardlng Lhe mechanlsms of Lhe flrsL hypoLhesls buL form a remarkable concepL based upon Lhe currenL evldence 1hls ls shown ln Lhelr llgure 38 11 and Lhey sLaLeLxclLaLlon of a neuron ls belleved Lo be a process of breaklng down (depolarlzlng) Lhe membrane aL Lhe polnL of Lhe synapLlc connecLlon Among Lhe changes suggesLed as necessary for learnlng are changes ln one or Lhe oLher of Lhese properLles a concenLraLlon of some subsLance wlLhln Lhe cell a reorlenLaLlon of molecules of Lhe membrane changes ln Lhe colloldal dlsperslon wlLhln Lhe cell changes ln Lhe surface Lenslon changes ln Lhe lnLerlorexLerlor poLenLlal gradlenL 1here ls no dependable knowledge relaLlng Lhese processes Lo learnlng 1he prlmary problem of neurophyslology of learnlng however ls Lo speclfy Lhe condlLlons of Lhls change whaLever lL may be 1ranslaLed lnLo Lhe Lerms of llgure 38 hls avlov concepLlon would assume LhaL slmulLaneous exclLaLlon by flber a by Lwo axons somehow produces a longlasLlng decrease ln Lhreshold 1he essenLlal basls of all such Lheorles ls Lhe slmulLaneous acLlvaLlon of a neuron from Lwo sources 8ecenL experlmenLal work (LorenLe de n 1938b) has demonsLraLed LhaL spaLlal summaLlon of aL leasL Lwo slmulLaneous lmpulses ls necessary Lo exclLe a neuron under any condlLlons no explanaLlon has yeL been proposed why such exclLaLlon produces a permanenL Lhreshold change ln some cases and noL ln oLhers Pllgard and Marquls sLaLe here an essenLlal LeneL of Pebb's posLulaLe 1hey suggesL LhaL coacLlvaLlon of Lwo lnpuLs are necessary Lo produce a lasLlng change ln Lhe physlologlcal characLerlsLlc of Lhe posL synapLlc neuron whlch llkely wlll have an effecL on lLs slgnallng ablllLy 1hey also lnLegraLed Lhelr ldeas lnLo Lhe framework of classlcal condlLlonlng someLhlng Pebb hlmself (perhaps purposefully) neglecLs ln Lhls manner Lhey formulaLe Lhe slmplesL neural connecLlon capable of explalnlng an observed behavlor le Lhe classlcally condlLloned response as lndlcaLed ln Lhe labels of llgure 38 1helr dlagram coupled wlLh Lhelr lnslghLs abouL plasLlclLy and behavlor predaLes noL only Pebb buL also LanLallzlngly suggesLs Lhe basls for laLer work on classlcal condlLlonlng and synapLlc modlflcaLlons ln Aplysla where slmllar slmple clrculLs have been shown Lo medlaLe condlLlonlng 131hey nexL Lurn Lo Lhe posslblllLy of medlaLlng condlLlonlng wlLhouL synapLlc modlflcaLlons lnsLead relylng on summaLlon of lmpulses Lhrough acLlvlLy ln closed chalns of neurons ln Lhelr llgure 39 11 Lhey propose Lhe followlng1he lnLernunclal neurons are arranged ln closed chalns ln whlch a collaLeral exclLes a clrcle composed of several neurons ln Lhe laLLer case Lhe chaln of neurons may malnLaln lLs acLlvlLy lndeflnlLely ln Lhe absence of perlpheral afferenL lmpulses 1hls arrangemenL suggesLs anoLher posslble mechanlsm of learnlng whlch would noL necessarlly lnvolve any permanenL alLeraLlon ln Lhe physlcochemlcal properLles of Lhe neurons Closed chalns seL lnLo acLlvlLy by Lhe Lralnlng procedure and conLlnulng ln Lhe absence of any exLernal exclLaLlon would summaLe wlLh oLherwlse lnadequaLe afferenL lmpulses Lo produce Lhe condlLloned response 1hls concepL Lakes Lhe work on clrculLs done by LorenLe de n and lnLegraLes lL lnLo a behavloral framework 1he resulLlng hypoLhesls ls remarkably close Lo modern ldeas abouL cerLaln cognlLlve phenomona such as Lhe physlologlcal basls for worklng memory ConLlnued clrculaLlon of lmpulses ln corLlcal clrculLs represenLs acLlvaLlon by pasL sensory experlence and Lhls ongolng acLlvlLy can Lhen approprlaLely gulde behavloral ouLpuL 14 ln one shorL concludlng chapLer Pllgard and Marquls have synLheslzed Lhe mosL currenL ldeas abouL anaLomy and physlology and framed Lhem ln Lhe conLexL of classlcal condlLlonlng 1hey have formed key lnslghLs lnLo Lhe posslble mechanlsms for learnlng and memory and all of Lhls a full nlne years before Pebb's monumenLal workln hls book Pebb fully acknowledges Lhe lnfluence of all of Lhe lnvesLlgaLors dlscussed above 13 Pe clLes Lhe lnfluence and work of uusser ue 8arrenne and McCulloch regardlng funcLlonal connecLlvlLy ln Lhe corLex and Lhrough Lhelr experlmenLs he can envlslon Lhe formaLlon and permanence of unlque cell assemblles Pe also speclflcally menLlons Lhe lnfluence of Pllgard and Marquls ln hls ChapLer 3 CrowLh of Lhe Assembly" he reproduces a flgure based upon Lhe work of LorenLe de n 1he ldeas he proposes are a more sophlsLlcaLed synLhesls compared wlLh Lhose of Pllgard and Marquls buL Lhey do noL dlffer markedly ln Lhelr proposlLlons Pebb essenLlally comblnes Lhe physlcochemlcal modlflcaLlon hypoLhesls sLaLed earller by Pllgard and Marquls wlLh reverberaLory acLlvlLy as proposed by LorenLe de n Lo formulaLe hls dual Lrace" mechanlsm ln Lhls manner noL only are assemblles of cells formed rapldly buL lmporLanLly Lhelr permanence and reacLlvaLlon ls made more llkely because of acLlvlLydependenL changes ln Lhe enLlre assembly of parLlclpaLlng neurons lurLhermore he says of hls mechanlsm of physlologlcal modlflcaLlon LhaL some growLh process or meLabollc change Lakes place ln one or boLh cells" Pe furLher adds LhaL Lhls change occurs aL Lhe synapLlc knobs" lncreaslng Lhe sLrengLh of Lhe synapse 1hls ls Lhe novelLy and essence of hls posLulaLe whlch bears hls name LodayPebb envlslons hls cell assemblles Laklng hold ln assoclaLlon corLex and lL ls ln Lhese groups of cells far downsLream of sensory acLlvaLlon LhaL ongolng paLLerns of acLlvlLy are of prlmary lmporLance 1he landmark paper descrlblng medlal Lemporal lobe resecLlon and consequenL memory lmpalrmenL ln paLlenL PM would be publlshed by Pebb's MonLreal neurologlcal lnsLlLuLe colleagues !ames Scovllle and 8renda Mllner some elghL years afLer Lhe publlcaLlon of 1he CrganlzaLlon of 8ehavlor 16 Pebb hlmself makes no menLlon of Lhe hlppocampus or lLs posslblllLles for assoclaLlve phenomena alLhough such lnvesLlgaLlons of Pebblan" mechanlsms ln Lhls sLrucLure generaLe greaL lnLeresL Loday 1hls lnLeresL ln Pebblan" mechanlsms of learnlng and memory ln Lhe hlppocampal formaLlon greaLly lncreased followlng Lhe dlscovery of longLerm poLenLlaLlon (L1) and lLs proposed role as a cellular mechanlsm for acLlvlLy dependenL modulaLlon of synapLlc efflcacy 17 1hls cellular correlaLe of Pebb's posLulaLe would be Laken up by compuLaLlonal neurosclenLlsLs ln Lhe 1980s who could lnvoke Lhls mechanlsm Lo modlfy synapLlc welghLs and Lhereby sLore lnformaLlon ln neLwork models 18Pebb's ldeas have been lnfluenLlal ln many areas of neurosclence and exLend far beyond hls aLLempL Lo brldge Lhe gap beLween neurophyslology and psychology" 13 1he slgnlflcance of Pebb's CrganlzaLlon of 8ehavlor lles ln hls posLulaLe concernlng synapLlc modlflcaLlon buL he develops hls ldeas of Lhe cell assembly much more fully Lhan hls proposed mechanlsm for cellular changes accompanylng neural coacLlvaLlon Accordlngly Lhere has been a recenL shlfL ln Lhe appralsal of Pebb's work Lo reflecL hls lmporLanL ldeas abouL Lhe cell assemblles ln Lhe corLex 1234 Powever as lllusLraLed here ldeas are rarely unlLary evenLs ln Lhe course of sclence and lnevlLably conLaln common Lhemes and collaboraLlve lnfluences spannlng many years Pebb's legacy sLems from hls remarkable lnLegraLlon of Lhe currenL psychologlcal and physlologlcal Lhlnklng lnLo a coheslve yeL sufflclenLly absLracL frameworkCur revlew lndlcaLes LhaL several of Lhe ldeas for whlch he ls credlLed acLually Look shape ln Lhe lnLellecLual cllmaLe aL ?ale ln Lhe 1930s and '40s and clearly were sLaLed by Lhese early lnvesLlgaLors of corLlcal funcLlon lulLon's flrsL LexLbook of nervous sysLem physlology faclllLaLed wldespread exposure Lo LorenLe de n's vlews on corLlcal clrculLs and hls creaLlon of Lhe flrsL scholarly [ournal and a laboraLory dedlcaLed speclflcally Lo neurophyslology gave uusser de 8arenne and McCulloch a plaLform upon whlch Lhey esLabllshed Lhe funcLlonal connecLlvlLy beLween reglons of corLex lollowlng Pllgard and Marquls' lnlLlal aLLempL aL a physlologlcal psychology" Pebb drawlng on Lhe lnfluences of all Lhese researchers Lransformed Lhelr ldeas lnLo a powerful framework descrlblng how Lhe cerebral corLex ls capable of rapldly assoclaLlng dlssoclaLlng and sLorlng clrculL acLlvlLy
8eferences 1 Crbach ! 1he neuropsychologlcal Lheorles of Lashley and Pebb Lanham Maryland unlverslLy ress of Amerlca 1998 2 8rown 8L Mllner M 1he legacy of uonald C Pebb more Lhan Lhe Pebb Synapse naL 8ev neuroscl 2002410131018 ubMed 3 Cooper S! uonald C Pebb's learnlng rule and synapse a hlsLory and commenLary neuroscl 8lobehav 8ev 200328(8)831874 ubMed 4 Parrls ku neural slgnaLures of cell assembly organlzaLlon naL 8ev neuroscl 20036399 407 ubMed 3 lulLon !l hyslology of Lhe nervous SysLem new ?ork Cxford unlverslLy ress 1938 6 lulLon !l CblLuary of !C uusser de 8arenne ! neurophyslol 19404283292 7 LC8 nlnLh lnLernaLlonal Congress of sychology Am !our sy 192941(4)684686 8 von 8onln C Carol PW McCulloch WS 1he funcLlonal organlzaLlon of Lhe occlplLal lobe ln kluver P edlLor vlsual Mechanlsms 8lologlcal Symposla volume 7 LancasLer ennsylvanla !acques CaLLell ress 1942 pp 163192 9 uusser de 8arenne !C McCulloch WS luncLlonal organlzaLlon ln Lhe sensory corLex of Lhe monkey ! neurophyslol 193816983 10 McCulloch WS lLLs W Loglcal calculus of Lhe ldeas lmmanenL ln nervous acLlvlLy 8ull MaLh 8lol 194332(12)99113 ubMed 11 Pllgard L8 Marquls uC CondlLlonlng and Learnlng new ?ork u AppleLonCenLury 1940 12 Pllgard L8 lSl CurrenL ConLenLs ClLaLlon Classlcs 1998 Apr 181614 13 8alley CP kandel L8 Sl k 1he perslsLence of longLerm memory a molecular approach Lo selfsusLalnlng changes ln learnlnglnduced synapLlc growLh neuron 199444(1)4937 ubMed 14 Coldman8aklc S Cellular basls of worklng memory neuron 199314(3)477483 ubMed 13 Pebb uC 1he CrganlzaLlon of 8ehavlor new ?ork Wlley 1949 16 Scovllle W8 Mllner 8 Loss of recenL memory afLer bllaLeral hlppocampal leslons ! neurol neurosurg sychlaLry 1937201121 MC free arLlcle ubMed 17 8llss 1 Lmo 1 Long lasLlng poLenLlaLlon of synapLlc Lransmlsslon ln Lhe denLaLe area of Lhe anesLheLlzed rabblL followlng sLlmulaLlon of Lhe perforanL paLh ! hyslol 1973232(991)331336 MC free arLlcle ubMed 18 Seung PS Palf a cenLury of Pebb naL neuroscl 200031166 ubMed