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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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0% found this document useful (0 votes)
45 views15 pages

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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© Attribution Non-Commercial (BY-NC)
We take content rights seriously. If you suspect this is your content, claim it here.
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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

Pls wlfe LllzabeLh dled ln 1962 ln 1966 Pebb marrled hls Lhlrd wlfe MargareL uoreen WrlghL (nee
Wllllamson) a wldow

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
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unlverslLy ress of Amerlca 1998
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3 Cooper S! uonald C Pebb's learnlng rule and synapse a hlsLory and commenLary neuroscl
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4 Parrls ku neural slgnaLures of cell assembly organlzaLlon naL 8ev neuroscl 20036399
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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
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