Revised 17 November 2013

Cognition and Working Memory in Brain Injury and Autism

Alan Challoner M A (Phil) MChS Working memory can be seriously affected by brain injury both In terms of the structures that are damaged and the failure of some developmental processes that follow, when the injury occurs early in a child s life! It is of importance for there to be a better understanding of autistic spectrum disorder so that consideration can be given to the possibilities that the condition may arise as a conse"uence of early brain damage! #, $, % A review of literature may help to understand better these issues especially to consider research that indicates how working memory affects our cognitive processes and how the deficits following brain injury affect cognition and communication as a result of damage to the prefrontal corte& and its connections to other brain areas! In the normal brain, the very complicated span of neural activity which we call working memory goes on without us having to think about it consciously! 'owever, with subjects who have damage to some of the areas involved, it can be seen why there is a failure of certain cognitive and social skills and it is important to portray these difficulties as being influenced by brain damage and not by lack of intellect! Attentional focus is important for many cognitive processes including problem solving and its differential effects on analytic and creative activity! (ne of the main ways in which working memory capacity benefits analytic problem solving seems to be that it helps to control attention, resist distraction and narrow the search through a problem space! Conversely, several lines of recent research show evidence that too much focus here can actually harm performance on creative problem)solving tasks! *

Working Memory

+Working memor ! is a "orm o" shor#$#erm memor #ha# is o"#en e%em&li"ied b #he #imes 'hen in"orma#ion is being held on line "or #he &(r&ose o" &er"orming )om&(#a#ions on i# (analogo(s #o a men#al s)ra#)h &ad)* +(s#er ,-. &rovided #he "irs# de#ailed a))o(n# o" #he role o" 'orking memor in &re"ron#al &ro)esses* /e des)ribed #he role o" #he &re"ron#al )or#e% as #ha# o" in#egra#ing #em&orall dis#rib(#ed in"orma#ion- a )om&le% &ro)ess 'hi)h he a##rib(#ed &ar#l #o shor#$#erm 'orking memor * Con#ras#ing #his vie' 'i#h s(&ervisor a##en#ional s s#em (SAS)$like e%e)(#ive a))o(n#s o" &re"ron#al

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h##&011'''*s)ribd*)om1do)12.3,34.1Abnormal$5rain$S#r()#(re$and$A(#ism$ h##&011'''*s)ribd*)om1do)13..,7401A(#ism$and$Abnormali#ies$in$#he$5rain h##&011'''*s)ribd*)om1do)1124032.715rain$6amage$Ca(sed$b $7a))ina#ion ,ennifer Wiley, ,! - ,aros., A/! Working Memor Ca&a)i# - A##en#ional +o)(s- and Problem Solving* C(rren# 6ire)#ions in Ps )hologi)al S)ien)e A(g(s# 2012 vol* 21 no* 4 2,3$2.2* /uster ,M! The Prefrontal Cortex: Anatomy, Physiology, and Neuropsychology of the Frontal Lobe. Ne' 8ork0 Raven Press- 1230* /uster ,M! 9he &re"ron#al )or#e% and #em&oral in#egra#ion- in :ones ;<- Pe#ers A (eds)0 Cerebral Cortex: 7ol 4* Association and Auditory Cortices. Ne' 8ork0 Raven Press= 123,*

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Revised 17 November 2013 "(n)#ion7- he 'ro#e0 >9he &re"ron#al )or#e% 'o(ld no# s(&erim&ose a s#eering or dire)#ing "(n)#ion on #he remainder o" #he nervo(s s s#em- b(# ra#her- b e%&anding #he #em&oral &ers&e)#ives o" #he s s#em- i# 'o(ld allo' i# #o in#egra#e longer- ne'er- and more )om&le% s#r()#(res o" behavio(r*? <oldman$Raki)3 has also &ro&osed a 'orking$memor a))o(n# o" "ron#al lobe "(n)#ion on #he basis o" e%#ensive resear)h 'i#h non$h(man &rima#es* 5(ilding on #he 'ell$es#ablished rela#ionshi& be#'een #he &re"ron#al )or#e% and dela ed$res&onse #asks- she arg(es #ha# #he &re"ron#al )or#e% is res&onsible "or main#aining in"orma#ion (>re&resen#a#ional memor ?) #ha# is la#er (sed #o g(ide a)#ion* +(nahashi and )olleag(es2-10 )arried o(# a varie# o" lesion s#(dies and single$(ni# re)ording s#(dies #o es#ablish #he role o" &re"ron#al )or#e% in 'orking memor - &rimaril 'i#h monke s #rained #o &er"orm s&a#ial 'orking memor #asks* Ne(ro$imaging s#(dies in h(mans11-12-13-14 s(gges# #ha# similar 'orking$memor &ro)esses are lo)a#ed &re$"ron#all in #he h(man brain* <oldman$Raki) (1237- idem) has &(# "or'ard a vie' #ha# #he asso)ia#ion be#'een &re"ron#al )or#e% and 'orking memor )an- in &rin)i&le- e%&lain a range o" h(man )ogni#ive im&airmen#s "ollo'ing "o)al "ron#al lesions as 'ell as o#her non$"o)al &a#hologies a""e)#ing &re"ron#al )or#e%* 9he &ro&osal #ha# 'orking memor im&airmen# )o(ld (nderlie #he range o" )ogni#ive )hanges seen a"#er &re"ron#al damage "irs# "o(nd dire)# s(&&or# in #he 'ork o" Cohen and Servan$S)hreiber1,*

Working Memor and ;arl 5rain @nA(r

9reble et al have "o(nd #ha# de"i)i#s in 'orking memor are a )ommon )onseB(en)e o" &aedia#ri) #ra(ma#i) brain inA(r and are believed #o )on#rib(#e #o di""i)(l#ies in a range o" )ogni#ive and a)ademi) domains* 9heir resear)h has sho'n #ha# red()ed in#egri# o" #he )or&(s )allos(m (CC) a"#er #ra(ma#i) brain inA(r ma disr(&# #he )onne)#ivi# be#'een bila#eral "ron#o$&arie#al ne(ral ne#'orks (nderl ing 'orking memor * 9he (sed di""(sion #ensor imaging (69@) #ra)#ogra&h #o e%amine eigh# )allosal s(bregions (CC1$CC3) in rela#ion #o meas(res o" verbal and vis(os&a#ial 'orking memor in 74 )hildren s(s#aining #ra(ma#i) brain inA(r and 42 # &i)all develo&ing )om&arison )hildren*

0upervisory attentional system C a loosel de"ined )olle)#ion o" brain &ro)esses #ha# are res&onsible "or &lanning- )ogni#ive "le%ibili# - abs#ra)# #hinking- r(le a)B(isi#ion- ini#ia#ing a&&ro&ria#e a)#ions and inhibi#ing ina&&ro&ria#e a)#ions- and sele)#ing relevan# sensor in"orma#ion* 1oldman)2akic 30! Cir)(i#r o" &rima#e &re"ron#al )or#e% and reg(la#ion o" behavior b re&resen#a#ional memor - in Pl(m +- Mo(n#)as#le 7 (eds)0 andboo! of Physiology, The Ner"ous #ystem: $. 5e#hesda- M60 Ameri)an Ph siologi)al So)ie# - 1237* /unahashi 04 Bruce C, - 1oldman)2akic 30* Mnemoni) )oding o" vis(al s&a)e in #he monke Ds dorsola#eral &re"ron#al )or#e%* % Neurophysiol &':(('342-1232* /unahashi 04 Bruce C, - 1oldman)2akic 30! 6orsola#eral &re"ron#al lesions and o)(lomo#or dela edres&onse &er"orman)e0 ;viden)e "or mnemoni) Es)o#omas!* % Neurosci 1301472$ 1427- 1223* Cohen ,4 /orman 0 - Braver 5, et al! A)#iva#ion o" &re"ron#al )or#e% in a nons&a#ial 'orking memor #ask 'i#h "(n)#ional MR@* um )rain *apping 10223$304- 1224* 678sposito M4 0hin 29 - 6etre ,A, et al! FbAe)# and s&a#ial 'orking memor a)#iva#es dorsola#eral &re"ron#al )or#e%0 A "(n)#ional MR@ s#(d * #oc Neurosci Abstr 2101423- 122,* 678sposito M4 6etre , - Alsop 6, et al! 9he ne(ral basis o" #he )en#ral e%e)(#ive s s#em o" 'orking memor * Nature 3730272$231- 122,* ,onides ,4 0mith 8 - 9oeppe 2, et al. S&a#ial 'orking memor in h(mans as revealed b P;9* Nature 3.30.23$.2,- 1223* Cohen ,6 - 0ervan)0chreiber 6* Con#e%#- )or#e%- and do&amine0 A )onne)#ionis# a&&roa)h #o behavior and biolog in s)hiGo&hrenia* Psychol +e" ,,:-.77-1222*

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Revised 17 November 2013 Rela#ive #o #he )om&arison gro(&- )hildren 'i#h #ra(ma#i) brain inA(r demons#ra#ed &oorer vis(os&a#ial 'orking memor - b(# )om&arable verbal 'orking memor * Mi)ros#r()#(re o" #he CC 'as signi"i)an#l )om&romised in brain$inA(red )hildren- 'i#h lo'er "ra)#ional aniso#ro& (+A) and higher a%ial and radial di""(sivi# me#ri)s in all )allosal s(bregions* @n bo#h gro(&s o" )hildren- lo'er +A and1or higher radial di""(sivi# in )allosal s(bregions )onne)#ing an#erior and &os#erior &arie#al )or#i)al regions &redi)#ed &oorer verbal WM- 'hereas higher radial di""(sivi# in )allosal s(bregions )onne)#ing an#erior and &os#erior &arie#al- as 'ell as #em&oral- )or#i)al regions &redi)#ed &oorer vis(os&a#ial 'orking memor * 69@ me#ri)s- es&e)iall radial di""(sivi# - in &redi)#ive )allosal s(bregions a))o(n#ed "or signi"i)an# varian)e in 'orking memor over and above remaining )allosal s(bregions* Red()ed mi)ros#r()#(ral in#egri# o" #he CC- &ar#i)(larl in s(bregions )onne)#ing &arie#al and #em&oral )or#i)es- ma a)# as a ne(ro&a#hologi)al me)hanism )on#rib(#ing #o long$ #erm WM de"i)i#s* 9he "(#(re )lini)al (se o" ne(roana#omi)al biomarkers ma allo' "or #he earl iden#i"i)a#ion o" )hildren a# highes# risk "or 'orking memor de"i)i#s and earlier &rovision o" in#erven#ions "or #hese )hildren* 1. /ardan et al "o(nd #ha# areas o" #he an#erior s(b$regions o" #he )or&(s )allos(m 'ere smaller in #hose 'i#h a(#ism #han in individ(all ma#)hed )on#rols* 17 9his ma s(&&or# a vie' #ha# some brain inA(ries )an res(l# in a(#is#i) s&e)#r(m disorder i" #he o))(r earl eno(gh in li"e* Information 3rocessing - Brain Injury: A )ommon de"i)i# arising "rom a brain inA(r and more )ommonl "rom a head inA(r - is #he inabili# #o &ro)ess in"orma#ion a# #he normal ra#e* Whereas #he )hild ma be able #o )arr o(# a varie# o" men#al #asks- #he s&eed a# 'hi)h #hese are )om&le#ed in a brain$inA(red )hild ma be signi"i)an#l slo'er #han i# 'o(ld be "or a normal )hild* 13 9he de"i)i# ma involve #he s&eed a# 'hi)h #he )hild )an (nders#and #he #ask involved- learn #he ma#erial- re#rieve i# "rom memor and #hen )arr o(# #he men#al &ro)esses involved* 6e"i)i#s in #his area )an severel limi# #he abili# o" #he )hild #o "(n)#ion in man ever da si#(a#ions and s()h de"i)i#s )an arise "ollo'ing even a mild head inA(r * 12 @n s&e)i"i) de#ail#he )hild "inds di""i)(l# )om&rehending in"orma#ion a# #he normal ra#e- "orm(la#ing #heir #ho(gh#s and #hen )arr ing o(# #he reB(ired a)#ions* 9he ma "ind i# di""i)(l# #o (nders#and i" #oo m()h in"orma#ion is &resen#ed a# an one o))asion* 9he ma #here"ore ini#iall s#ar# o"" (nders#anding 'ha# is going on b(# ra&idl lose #heir 'a as #he amo(n# o" in"orma#ion a))(m(la#es* 9his is &ar#i)(larl #he )ase i" #he level o" in"orma#ion be)omes more )om&le% and i# )an o))(r in bo#h #he )lassroom and in so)ial si#(a#ions* 5orod "o(nd brain$inA(red )hildren #o be less )om&e#en# a# )om&rehending emo#ional in"orma#ion* @n #he #ea)hing si#(a#ion #his o"#en sho's i#sel" in #he )hildDs inabili# #o )om&le#e 'ri##en assignmen#s or ans'er B(es#ions in #he allo##ed #ime- and #here"ore

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5reble A4 'asan 9M4 Iftikhar A4 0tuebing 994 9ramer <A4 Co& C0 ,r4 0wank 32 - 8wing)Cobbs <* Working memor and )or&(s )allos(m mi)ros#r()#(ral in#egri# a"#er &edia#ri) #ra(ma#i) brain inA(r 0 a di""(sion #ensor #ra)#ogra&h s#(d * % Neurotrauma. 2013 F)# 1=30(12)01.02$12* doi0 10*10321ne(*2013* 2234* 'ardan A>4 Minshew ?, - 9eshavan M0* Cor&(s Callos(m SiGe @n A(#ism* Neurology 2000 F)# 10=,,(7)01033$.* 5rooks, ?! Cogni#ive de"i)i#s a"#er head inA(r * @n0 Closed ead /n0ury: Psychosocial #ocial and Family Conse1uences (ed* N* 5rooks)- &&* 44$73* F%"ord0 F%"ord Comm(ni# Press= 1234* Wrightson, 3!, Mc1inn, @!, - 1ronwall, 6! Mild head inA(r in &re$s)hool )hildren H eviden)e #ha# )an be asso)ia#ed 'i#h a &ersis#ing )ogni#ive de"e)#* %ournal of Neurology Neurosurgery and Psychiatry ,2- 37,$30= 122,*

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Revised 17 November 2013 never being able #o ans'er a B(es#ion dire)#ed a# #he )lass- be)a(se b #he #ime #he raise #heir hand someone else has alread ans'ered be"ore #hem* 20 Children 'ho are signi"i)an#l a""e)#ed )an "ind #hemselves severel disadvan#aged a# a so)ial level* Whereas ad(l#s 'ill give a )hild a s m&a#he#i) look and #he ne)essar #ime #o )olle)# #heir #ho(gh#s- a more )om&e#i#ive adoles)en# gro(& 'ill no# be so kind or so #oleran#* @n #he &la gro(nd- #he )onversa#ion moves a# #he &a)e o" #he gro(& and "or #hose #oo slo' #o res&ond- #he "reB(en#l "ind #hemselves le"# "ar behind= some#imes i# is easier no# #o even #r - and #he )hildren ma "ind #hemselves be)oming in)reasingl isola#ed "rom #heir &eers* 21 9he abili# #o (nders#and- learn and #hen re#rieve in"orma#ion involves a range o" abili#iesin)l(ding a##en#ion- shor#$#erm memor and #he abili# #o mani&(la#e #he in"orma#ion in order #o &la)e i# in#o a more &ermanen# memor s s#em* ;ssen#ial #o #his- is #he abili# #o organiGe ma#erial in#o a meaning"(l manner so #ha# i# )an be re)alled and s(bseB(en#l &la)ed 'i#hin some e%is#ing s)heme or order* As 9rom& and M(lder 22 demons#ra#eda))ess #o #he memor s s#em a""e)#s #he s&eed a# 'hi)h in"orma#ion is &ro)essed* @n"orma#ion #here"ore has #o be organiGed and )a#egoriGed so #ha# i# )an be s#ored in some &ermanen# memor s s#em* 9o do #his- i# is also ne)essar "or #he )hild #o be able #o dis#ing(ish #he )ore in"orma#ion and main ideas "rom an irrelevan# in"orma#ion* @# is a&&aren# #ha# a range o" s#ra#egies )an be (sed a# a sim&lis#i) level* 9he ra#e- amo(n# and )om&le%i# o" #he ma#erial ma - ho'ever- e%)eed 'ha# #he )hild )an )om"or#abl )o&e 'i#h a# a sim&lis#i) level* 9o hel& #he )hild )o&e 'i#h s()h &ro)essing &roblems- i# is im&or#an# #ha# &aren#s and #ea)hers are a'are #ha# #he )hild ma have s()h di""i)(l#ies* @" &eo&le are so a'are- #he &roblem )an be &ar#l resolved b al#ering #he ra#e a# 'hi)h 'ork is &resen#ed* @# is also ver val(able i" #he )hild )an be o""ered some basi) s#ra#egies and &rom&# him #o sa C)an o( &lease e%&lain #ha# again C)an o( go #hro(gh #ha# a li##le slo'er- &lease C or- C )an o( le# me #hink abo(# #ha# "or a min(#e be"ore @ ans'er* 9hese sor#s o" res&onses- i" a&&roved- )an give #he )hild a se)ond o&&or#(ni# #o ans'er or resolve #he &roblem and hel& #o &reven# an an%ie# $&rod()ing si#(a#ion* @# is &ossible #o give )hildren more general s#ra#egies in order #o hel& #o s&eed (& #he &ro)essing &roblem 'i#hin i#s o'n righ#* 9he )hild )an be en)o(raged #o (se vis(al imager - &rovided #ha# #he have in#a)# vis(al and &er)e&#(al skills* 9his vis(al imager #e)hniB(e has been 'ell e%&lained b 5(Gan 23 (sing Dmind ma&sD 'hereb #he )hild )an be en)o(raged #o (se diagrams as )(es #o hel& &rom&# #he memor &ro)ess* 9his #e)hniB(e )an also be (sed #o enhan)e bo#h shor#$#erm and 'orking memor * @# )an sho' ho' ne' learning )an be a##a)hed #o older more es#ablished and more in#a)# areas o" memor and #here"ore #he )hild )an be sho'n on 'hi)h Ehooks! #o hang #he ne' in"orma#ion* Cortical visual processing begins in #he &rimar vis(al area lo)a#ed in #he o))i&i#al lobe (#he rear$mos# &ar# o" #he )or#e%)* 9his area re)eives vis(al in"orma#ion "rom #he vis(al #halam(s- &ro)esses i#- and #hen dis#rib(#es i#s o(#&(#s #o a varie# o" o#her )or#i)al regions* Al#ho(gh #he )or#i)al vis(al s s#em is enormo(sl )om&le%-24 #he ne(ral &a#h'a s res&onsible "or #'o as&e)#s o" vis(al &ro)essing are "airl 'ell (nders#ood* 9hese involve #he

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Borod, ,! C* @n#erhemis&heri) and in#rahemis&heri) )on#rol o" emo#ion* %ournal of Consulting and Clinical Psychology .0- 332$43= 1222* Appleton, 2! - Baldwin, 5! Managemen# o" 5rain @nA(red Children* 2nd ;d* FIP-200.* 5romp, 8! - Mulder, 5! Ne(ro&s )hologi)al slo'ness o" in"orma#ion &ro)essing a"#er a #ra(ma#i) head inA(r * %ournal of Clinical and 2xperimental Neuropsychology 13- 321$30= 1221* Bu.an, 5! Ise 8o(r /ead* Jondon0 55C P(bli)a#ions*= 1274* @an 8ssen, 6C! +(n)#ional organiGa#ion o" &rima#e vis(al )or#e%* @n Cerebral cortex- A* Pe#ers and ;* <* :ones- eds* (Ne' 8ork0 Plen(m)- &&*2,2$323= 123,*

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Revised 17 November 2013 de#ermina#ion o" E'ha#! a s#im(l(s is and E'here! i# is lo)a#ed* 2,- 2. 9he E'ha#! &a#h'a involves a &ro)essing s#ream #ha# #ravels "rom #he &rimar vis(al )or#e% #o #he #em&oral lobe and #he E'here! &a#h'a goes "rom #he &rimar )or#e% #o #he &arie#al lobe* 9he &arie#al and "ron#al regions in B(es#ion are ana#omi)all in#er)onne)#ed C #he &arie#al area sends a%ons #o #he &re"ron#al region and #he &re"ron#al region sends a%ons ba)k #o #he &arie#al area* 9hese "indings s(gges# #ha# #he &arie#al lobe vis(al area 'orks 'i#h #he la#eral &re"ron#al )or#e% #o main#ain in"orma#ion abo(# #he s&a#ial lo)a#ion o" vis(al s#im(li in 'orking memor * Similarl - Rober# 6esimone "o(nd eviden)e "or re)i&ro)al in#era)#ions be$ #'een #he vis(al areas o" #he #em&oral lobe (#he E'ha#! &a#h'a ) and #he la#eral &re"ron#al )or#e% in s#(dies involving #he re)ogni#ion o" 'he#her a &ar#i)(lar obAe)# had been seen re)en#l 27* 9he main#enan)e o" vis(al in"orma#ion in 'orking memor #h(s a&&ears #o de&end )r()iall on in#era)#ions be#'een #he la#eral &re"ron#al region and s&e)ialiGed areas o" #he vis(al )or#e%* <oldman$Raki) and )olleag(es re)orded "rom )ells in #he &arie#al lobe E'here! &a#h'a d(ring shor#$#erm memor #es#s reB(iring #he #em&orar remembering o" #he s&a#ial lo)a#ion o" vis(al s#im(li* 9he "o(nd #ha# )ells #here- like )ells in #he la#eral &re"ron#al )or#e%- 'ere a)#ive- s(gges#ing #ha# #he 'ere kee&ing #ra)k o" #he lo)a#ion- d(ring #he dela * (<oldman$Raki)- 1237- idem) S#(dies- es&e)iall b Wilson and asso)ia#es23- have raised B(es#ions abo(# #he role o" #he &re"ron#al )or#e% as a general &(r&ose 'orking memor &ro)essor* +or e%am&le- #he have "o(nd #ha# di""eren# &ar#s o" #he la#eral &re"ron#al )or#e% &ar#i)i&a#e in 'orking memor 'hen animals have #o de#ermine E'ha#! a vis(al s#im(l(s is as o&&osed #o E'here! i# is lo)a#ed- s(gges#ing #ha# di""eren# &ar#s o" #he &re"ron#al )or#e% are s&e)ialiGed "or di""eren# kinds o" 'orking memor #asks* While #hese "indings sho' #ha# &ar#s o" #he &re"ron#al )or#e% &ar#i)i&a#e (niB(el in di""eren# shor#$#erm memor #asks- #he do no# r(le o(# #he e%is#en)e o" a general$&(r&ose 'orks&a)e and a se# o" e%e)(#ive "(n)#ions #ha# )oordina#e #he a)#ivi# o" #he s&e)ialiGed s s#ems- es&e)iall sin)e #he #asks s#(died do no# #a% #he )a&a)i# o" 'orking memor in a 'a #ha# 'o(ld reveal a limi#ed )a&a)i# s s#em* 9he &a#h'a "rom #he s&e)ialiGed vis(al areas #ells #he &re"ron#al )or#e% E'ha#! is o(# #here and E'here! i# is lo)a#ed (bo##om$(& &ro)essing)* 9he &re"ron#al )or#e%- b 'a o" &a#h'a s ba)k #o #he vis(al areas- &rimes #he vis(al s s#em #o a##end #o #hose obAe)#s and s&a#ial lo)a#ions #ha# are being &ro)essed in 'orking memor (#o&$do'n &ro)essing)* 9hese kinds o" #o&$do'n in"l(en)es on sensor &ro)essing are believed #o be im&or#an# as&e)#s o" #he e%e)(#ive )on#rol "(n)#ions o" 'orking memor *

2,

Angerleider, <14 - Mishkin, M! 9'o )or#i)al vis(al s s#ems* @n Analysis of "isual beha"ior- 6* :* @ngle- M* A* <oodale- and R* :* W* Mans"ield- eds* (Cambridge0 M@9 Press)- &&* ,42$3.= 1232* Angerleider, <14 - 'a&by, ,! Wha# and 'here= in #he h(man brain* Current 3pinion in Neurobiology 4- 1,7$.,= 1224* 6esimone, 24 Miller, 894 Chela..i, <4 - <ueschow, A! M(l#i&le memor s s#ems in #he vis(al )or#e%* @n The cogniti"e neurosciences, M* S* <aGGaniga- ed* (Cambridge0 M@9 Press)- &&* 47,$ 3.= 122,* Wilson, /AW4 0calaidhe, 034 - 1oldman)2akic, 30! 6isso)ia#ion o" obAe)# and s&a#ial &ro)essing domains in &rima#e &re"ron#al )or#e%* #cience 2.0- 12,,$,3= 1223*

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Revised 17 November 2013

2elation of the BWhatB and BWhereB @isual 3athways to Working Memory! Visual information, received by the visual cortex, is distributed to cortical areas that perform specialized visual processing functions. Two well-studied specialized functions are those involved in object recognition (mediated by the what pathway) and object location (mediated by the where pathway). These specialized visual pathways provide inputs to the prefrontal cortex (PFC), which plays a crucial role in working memory. The specialized systems also receive inputs back from the PFC, allowing the information content of working memory to influence further processing of incoming information. Leftward-going arrows represent bottom-up processing and rightward-going ones top-down processing.

F#her s#(dies #ha# have #a%ed #he s s#em- like imaging s#(dies in h(mans- s(gges# #ha# ne(rons in #he la#eral &re"ron#al )or#e% are &ar# o" a general$&(r&ose 'orking memor ne#'ork* A# #he same #ime- i# is &ossible- given <oldman$Raki)Ds "indings #ha# #he general$ &(r&ose as&e)#s o" 'orking memor are no# lo)aliGed #o a single &la)e in #he la#eral &re"ron#al )or#e% b(# ins#ead are dis#rib(#ed over #he region* 9ha# #his ma o))(r is s(gges#ed b #he "a)# #ha# some )ells in #he s&e)ialiGed areas o" #he la#eral &re$"ron#al )or#e% &ar#i)i&a#e in m(l#i&le 'orking memor #asks*22 Working$memor a))o(n#s o" &re"ron#al "(n)#ion are "avo(red "or a n(mber o" reasons300 /irst- #he are &arsimonio(s- in #ha# 'orking$memor #heories )on#ain onl #he individ(al &ro)essing )om&onen#s needed #o &er"orm #he #ask 'i#ho(# needing a )en#ral e%e)(#ive (s()h as #he SAS) #o )oordina#e #hese )om&onen#s (and #o serve as #he lo)(s o" damage 'hen e%&laining &a#ien# behavio(r)* 0econd- #he have &roven )a&able o" e%&laining a 'ider range o" seemingl dis&ara#e im&airmen#s #han o#her non$e%e)(#ive #heories* 31 5hird- #he are s(&&or#ed b a 'eal#h o" eviden)e "rom monke ne(ro&h siolog and- in$ )reasingl - "rom ne(ro$imaging s#(dies in h(mans* /ourth- #he s(gges# a 'a o" resolving 'ha# is &erha&s #he )en#ral &roblem o" #he ne(ro$
22 30

3etrides, M! +ron#al lobes and behavio(r* Current 3pinion in Neurobiology 4- 207$11= 1224* 9imberg, 6>4 6 8sposito, M - /arah, M,! +ron#al Jobes0 Cogni#ive and Ne(ro&s )hologi)al As&e)#s* @n +einberg- 9 ; K +arah- M :* )eha"ioural Neurology and Neuropsychology* M)<ra'$/ill- Ne' 8ork- 1227* 9imberg 6> - /arah M,! A (ni"ied a))o(n# o" )ogni#ive im&airmen#s "ollo'ing "ron#al lobe damage0 9he role o" 'orking memor in )om&le%- organiGed behavior* % 2xp Psychol 4en 1220411$423- 1223*

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Revised 17 November 2013 &s )holog o" #he "ron#al lobe "(n)#ion C #he &arado% o" disso)iable im&airmen#s 'i#h an (nin#(i#ive )om&elling E"amil resemblan)e!* @" 'e ass(me #ha# 'orking memor is )om&ar#men#aliGed in #he &re"ron#al )or#e% a))ording #o 'ha# is being re&resen#ed in memor ("or 'hi)h eviden)e e%is#s)- #hen &er"orman)e in di""eren# #asks )an be im&aired or s&ared de&ending on 'hi)h # &es o" 'orking memor have been damaged* Never#heless- a))ording #o 'orking$memor a))o(n#s- #here is an (nderl ing )ommonali# among #he #asks sensi#ive #o &re"ron#al damage namel - #hose #ha# have de&enden)e on 'orking memor * (Limb(rg- et al - 1227- /dem) 9here is also eviden)e #ha# #he general$&(r&ose "(n)#ions o" 'orking memor involve areas o#her #han #he la#eral &re"ron#al )or#e%* +or e%am&le- imaging s#(dies in h(mans have sho'n #ha# ano#her area o" #he "ron#al lobe- #he an#erior )ing(la#e )or#e%- is also a)#iva#ed b 'orking memor and rela#ed )ogni#ive #asks* (6!;s&osi#o- et al- 122,- /dem) K 32- 33 Jike #he la#eral &re$"ron#al )or#e%- #he an#erior )ing(la#e region re)eives in&(#s "rom #he vario(s s&e)ialiGed sensor b(""ers- and #he an#erior )ing(la#e and #he la#eral &re$"ron#al )or#e% are ana#omi)all in#er)onne)#ed* (+(s#er- 1232- /dem) K 34* Moreover- bo#h regions are &ar# o" 'ha# has been )alled #he "ron#al lobe a##en#ional ne#'ork- a )ogni#ive s s#em involved in sele)#ive a##en#ion- men#al reso(r)e allo)a#ion- de)ision$making &ro)esses- and vol(n#ar movemen# )on#rol* 3, @# is #em&#ing #o #hink o" #he general$&(r&ose as&e)#s o" 'orking memor as involving ne(rons in #he la#eral &re"ron#al and an#erior )ing(la#e regions 'orking #oge#her* Fne o#her area o" #he &re"ron#al )or#e%- #he orbi#al region- lo)a#ed on #he (ndernea#h side o" #he "ron#al lobe- has emerged as im&or#an# as 'ell* 6amage #o #his region in animals in#er"eres 'i#h shor#$#erm memor abo(# re'ard in"orma#ion- abo(# 'ha# is good and bad a# #he momen# 3.- and )ells in #his region are sensi#ive #o 'he#her a s#im(l(s has A(s# led #o a re'ard or &(nishmen# 37- 33- 32* /(mans 'i#h orbi#al "ron#al damage be)ome oblivio(s #o so)ial and emo#ional )(es and some e%hibi# so)io&a#hi) behavio(r* 40 9his area re)eives in&(#s "rom sensor &ro)essing s s#ems (in)l(ding #heir #em&orar b(""ers) and is also in#ima#el )onne)#ed 'i#h #he am gdala and #he an#erior )ing(la#e region* 9he orbi#al )or#e% &rovides a link #hro(gh 'hi)h emo#ional &ro)essing b #he am gdala migh# be

32

Corbetta, M4 Mie.in, /M4 6obmeyer, 04 0hulman, 1<4 - 3etersen, 08* Sele)#ive and divided a##en#ion d(ring vis(al dis)rimina#ions o" sha&e- )olor- and s&eed0 +(n)#ional ana#om b &osi#ron emission #omogra&h * %ournal of Neuroscience 11- 2333$2402= 1221* 3osner, M4 - 3etersen, 0! 9he a##en#ion s s#em o" #he h(man brain* Annual +e"ie5 of Neuroscience 13- 2,$42= 1220* 1oldman)2akic, 30! 9o&ogra&h o" )ogni#ion0 Parallel dis#rib(#ed ne#'orks in &rima#e asso)ia#ion )or#e%* Annual +e"ie5 of Neuroscience 11- 137$,.= 1233* 3osner, M! A##en#ion as a )ogni#ive and ne(ral s s#em* Current 6irections in Psychological #cience 1- 11$14= 1222* 1affan, 64 Murray, 8A4 - /abre)5horpe, M! @n#era)#ion o" #he am gdala 'i#h #he "ron#al lobe in re'ard memor * 2uropean %ournal of Neuroscience ,- 2.3$7,= 1223* 5horpe, 0,4 2olls, 854 - Maddison, 0! 9he orbi#o"ron#al )or#e%0 Ne(ronal a)#ivi# in #he behaving monke * 2xperimental )rain +esearch 42- 23$11,= 1233 * 2olls, 85! Ne(ro&h siolog and "(n)#ions o" #he &rima#e am gdala* @n The amygdala: Neurobiological aspects of emotion, memory, and mental dysfunction, :* P* Aggle#on- ed* (Ne' 8ork0 Wile $Jiss)- &&* 143$.,= 1222* (no, 5 - ?ishijo, '! Ne(ro&h siologi)al basis o" #he Lliiver5() s ndrome0 Res&onses o" monke am gdaloid ne(rons #o biologi)all signi"i)an# obAe)#s* In The amygdala: Neurobiological aspects of emotion, memo,78 and mental dysfunction, :* P* Aggle#on- ed* (Ne' 8ork0 Wile $Jiss)- &&* 1.7$20= 1222* 6amasio, A! 6escartes error: 2motion, reason, and the human brain. Ne' 8ork0 <rosse#1P(#nam= 1224*

33

34

3,

3.

37

33

32

40

Page 7 of 22

Revised 17 November 2013 rela#ed in 'orking memor #o in"orma#ion being &ro)essed in sensor or o#her regions o" #he neo)or#e%* Je6o(% 'ri#es #ha# #he involvemen# o" s&e)ialiGed shor#$#erm b(""ers in #he sensor s s#ems and a general$&(r&ose 'orking memor me)hanism in #he &re"ron#al )or#e% is a ver )om&li)a#ed s s#em* 41 9he &re"ron#al )or#e% i#sel" seems #o have regions #ha# are s&e)ialiGed- a# leas# #o some degree- "or s&e)i"i) kinds o" 'orking memor "(n)#ions* S()h "indings- ho'ever- do no# dis)redi# #he no#ion #ha# #he &re"ron#al )or#e% is involved in #he general$&(r&ose or e%e)(#ive as&e)#s o" 'orking memor sin)e onl some )ells in #hese areas &la s&e)ialiGed roles* @n#era)#ions be#'een #he general$&(r&ose )ells in di""eren# areas ma )oordina#e #he overall a)#ivi# o" 'orking memor * @# is #h(s &ossible #ha# #he e%e)(#ive "(n)#ions o" #he &re"ron#al )or#e% migh# be media#ed b )ells #ha# are dis#rib(#ed a)ross #he di""eren# &re"ron#al s(bs s#ems ra#her #han b )ells #ha# are )olle)#ed #oge#her in one region* (Je6o(%- 1223- idem)

What are BuffersC

A )ri#i)al )om&onen# o" 5addele !s 'orking memor model is #he e%is#en)e o" verbal mid$s&a#ial s#orage b(""ers42* 9he )ogni#ive )on)e&# o" a b(""er #ransla#ed in#o ne(ral #erms 'o(ld &ro&ose #ha# #em&orar re#en#ion o" #ask$relevan# in"orma#ion reB(ires #rans"er o" #ha# in"orma#ion #o a &ar# o" #he brain #ha# is dedi)a#ed #o #he s#orage o" in"orma#ion* Pres(mabl - s()h b(""ers are analogo(s #o a )om&(#er!s RAM'hi)h serves as a )a)he "or in"orma#ion #rans"erred "rom #he hard drive #ha# is &ro)essed b a CPI* Consis#en# 'i#h #his in#er&re#a#ion o" a 'orking memor Eb(""er!- man des)ri&#ions o" )ogni#ive models o" 'orking memor re"er #o #he in"orma#ion being Ein! or Eo(#! o" 'orking memor * 2elation of 0peciali.ed 0hort)5erm Buffers, <ong)5erm 8&plicit Memory and Working Memory!

Stimuli processed in different specialized systems (such as sensory, spatial, or language systems) can be held simultaneously in short-term buffers. The various short-term buffers provide potential inputs to working memory, which can deal most effectively with only one of the buffers at a time. Working memory integrates information received from short-term buffers with long-term memories that are also activated.

Je6o(% s(gges#s #ha# #his involvemen# o" s&e)ialiGed shor#$#erm b(""ers in #he sensor s s#ems and a general$&(r&ose 'orking memor me)hanism in #he &re"ron#al )or#e% is a ver )om&li)a#ed s s#em* 9he &re"ron#al )or#e% i#sel" seems #o have regions #ha# are
41 42

<e6ou&, ,! 9he ;mo#ional 5rain* Weiden"eld K Ni)holson- 1223 @S5N 0227341034* Baddeley, A! 9or!ing *emory. Ne' 8ork- N80 F%"ord Iniversi# Press= 123.*

Page 8 of 22

Revised 17 November 2013 s&e)ialiGed- a# leas# #o some degree- "or s&e)i"i) kinds o" 'orking memor "(n)#ions* S()h "indings- ho'ever- do no# dis)redi# #he no#ion #ha# #he &re"ron#al )or#e% is involved in #he general$&(r&ose or e%e)(#ive as&e)#s o" 'orking memor sin)e onl some )ells in #hese areas &la s&e)ialiGed roles* @n#era)#ions be#'een #he general$&(r&ose )ells in di""eren# areas ma )oordina#e #he overall a)#ivi# o" 'orking memor * @# is #h(s &ossible #ha# #he e%e)(#ive "(n)#ions o" #he &re"ron#al )or#e% migh# be media#ed b )ells #ha# are dis#rib(#ed a)ross #he di""eren# &re"ron#al s(bs s#ems ra#her #han b )ells #ha# are )olle)#ed #oge#her in one region* (Je6o(%- 1223- idem) @n a revie' o" 'orking memor - Re&ovs K 5addele 43 s#a#e #ha#- >#he "(n)#ion o" #he ar#i)(la#or rehearsal &ro)ess is #o re#rieve and rear#i)(la#e #he )on#en#s held in #his &honologi)al s#ore and in #his 'a #o re"resh #he memor #ra)e?* 44 +(r#her- 'hile s&ee)h in&(# enters #he &honologi)al s#ore a(#oma#i)all - in"orma#ion "rom o#her modali#ies en#ers #he &honologi)al s#ore onl #hro(gh re)oding in#o &honologi)al "orm- a &ro)ess &er"ormed b >ar#i)(la#or rehearsal?* Ja#er- #he a(#hors re"er #o- >"o)al shi"#s o" a##en#ion #o memoriGed lo)a#ions #ha# &rovide a rehearsal$like "(n)#ion o" main#aining in"orma#ion a)#ive in s&a#ial 'orking memor ?* 9h(s- one B(es#ion #ha# ne(ro$s)ien#i"i) da#a )an address regarding ho' #he brain im&lemen#s 'orking memor &ro)esses is 'he#her s()h b(""ers or s#orage si#es e%is# in dis#in)# &ar#s o" #he brain #o s(&&or# #he a)#ive main#enan)e o" #ask$relevan# in"orma#ion* A )ogni#ive model o" 'orking memor - has been &(# "or#h b Co'an- 4,- 4. i# &ro&oses #ha# #he E)on#en#s o" 'orking memor ! are no# main#ained 'i#hin dedi)a#ed s#orage b(""ers- b(# ra#her are sim&l #he s(bse# o" in"orma#ion #ha# is 'i#hin #he "o)(s o" a##en#ion a# a given #ime* /e des)ribes an embedded$&ro)esses model 'here 'orking memor )omes "rom hierar)hi)all arranged "a)(l#ies )om&rising long$#erm memor - #he s(bse# o" 'orking long$ #erm memor #ha# is )(rren#l a)#iva#ed and #he s(bse# o" a)#iva#ed memor #ha# is #he "o)(s o" a##en#ion* 9hese ideas are similar #o #ha# &(# "or#h b Anderson 47 'ho re"erred #o 'orking memor as #hose re&resen#a#ions )(rren#l a# a high level o" a)#iva#ion* 9h(s- #ask$ relevan# re&resen#a#ions are no# in 'orking memor - b(# #he do have levels o" a)#iva#ion #ha# )an be higher or lo'er* A"#er (se- "or e%am&le- re&resen#a#ions ma be #em&oraril more a)#ive or E&rimed!* @n #his "orm(la#ion- 'orking memor does no# have a siGe- or ma%im(m n(mber o" i#ems- as a s#r()#(ral "ea#(re* @ns#ead- &er"orman)e on 'orking memor #asks is de#ermined b #he level o" a)#iva#ion o" relevan# re&resen#a#ions- and #he dis)riminabili# o" a)#iva#ion levels be#'een relevan# and irrelevan# re&resen#a#ions* 43 Again- in ne(ral #erms- Co'an!s or Anderson!s )ogni#ive model o" 'orking memor 'o(ld &redi)# #ha# in"orma#ion #ha# is re&resen#ed #hro(gho(# #he brain is no# #rans"erred #o an inde&enden# b(""er or s#orage si#e- b(# ra#her #ha# #em&orar re#en#ion o" #ask$relevan# in"orma#ion is media#ed b #he a)#iva#ion o" #he ne(ral s#r()#(res #ha# re&resen# #he in"orma#ion being main#ained or s#ored ("or a "(r#her dis)(ssion o" #hese and rela#ed ideas-

43

2epovs, 1!- Baddeley, A! 9he m(l#i$)om&onen# model o" 'orking memor 0 e%&lora#ions in e%&erimen#al )ogni#ive &s )holog * Neuroscience 132- ,$2= 200.* 9he phonological loop (or Mar#i)(la#or loo&M) as a 'hole deals 'i#h so(nd or &honologi)al in"orma#ion* @# )onsis#s o" #'o &ar#s0 a shor#$#erm phonological store 'i#h a(di#or memor #ra)es #ha# are s(bAe)# #o ra&id de)a and an articulatory rehearsal component (some#imes )alled #he articulatory loop) #ha# )an revive #he memor #ra)es* Cowan, ?! ;volving )on)e&#ions o" memor s#orage- sele)#ive a##en#ion- and #heir m(#(al )ons#rain#s 'i#hin #he h(man in"orma#ion &ro)essing s s#em* Psycho /. )ull. 104- 1.3$171= 1233* Cowan, ?! An embedded$&ro)ess model o" 'orking memor * @n *odels of 5or!ing memory: mechanisms of acti"e maintenance and executi"e control (eds A* Mi ake K P* Shah)- &&* .2$ 101* Cambridge- IL0 Cambridge Iniversi# Press= 1222* Anderson, ,! 2! The architecture of cognition. Cambridge- MA0 /arvard Iniversi# Press= 1233* Limberg, 6! >!, 678sposito, M! - /arah, M! ,! Cogni#ive "(n)#ions in #he &re"ron#al )or#e%$ 'orking memor and e%e)(#ive )on#rol* Curro 6ir. Psychol. S)i* ;, 13,$122= 1227*

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4,

4.

47 43

Page 9 of 22

Revised 17 November 2013 see R()hkin- et al 42* @n o#her 'ords- #he #em&orar re#en#ion o" a "a)e- "or e%am&le- 'o(ld reB(ire a)#iva#ion o" )or#i)al areas #ha# are involved in #he &er)e&#(al &ro)essing o" "a)es* +rom a ne(ros)ien)e &ers&e)#ive- i# is )o(n#er$in#(i#ive #ha# all #em&oraril s#ored in"orma#ion d(ring goal$dire)#ed behavio(r reB(ires s&e)ialiGed dedi)a#ed b(""ers* Clearl #here )o(ld no# be a s(""i)ien# n(mber o" inde&enden# b(""ers #o a))ommoda#e #he in"ini#e # &es o" in"orma#ion #ha# need #o be a)#ivel main#ained #o a))ommoda#e all &o#en#ial or in#ended a)#ions* +or ins#an)e- in a s s#em 'i#h onl #'o b(""ers- s()h as verbal and vis(o$s&a#ial- ho' 'o(ld #he re#en#ion o" odo(rs or #a)#ile sensa#ions- 'hi)h )anno# al'a s be re)oded in#o verbal or vis(o$s&a#ial re&resen#a#ions- be a))om&lishedN An addi#ional e&isodi) b(""er has been &ro&osed #o be a s#ore )a&able o" m(l#i$dimensional )oding #ha# allo's #he binding o" in"orma#ion #o )rea#e an in#egra#ed e&isode,0 * /o'ever- even 'i#h #he addi#ion o" #his b(""er- 5addele !s 'orking memor model )anno# a))ommoda#e s#orage o" all &ossible # &es o" in"orma#ion &ro)essed b #he h(man brain (i# is im&or#an# #o no#e- ho'ever- #ha# #his 'as no# &robabl #he original in#en# o" his model)* Al#erna#ivel - in Co'an!s &ro&osal- 'hi)h does no# rel on #he )on)e&# o" s&e)ialiGed dedi)a#ed s#orage b(""ers- a)#ive main#enan)e or s#orage o" #ask$relevan# re&resen#a#ions )o(ld be im&lemen#ed 'i#h a ne(ral s s#em 'here memor s#orage o))(rs in #he ver same brain )ir)(i#r #ha# s(&&or#s #he &er)e&#(al re&resen#a#ion o" in"orma#ion* S()h a ne(ral s s#em &res(mabl 'o(ld be more "le%ible and e""i)ien# #han one #ha# #rans"ers in"orma#ion ba)k and "or#h be#'een dedi)a#ed s#orage b(""ers* Fbvio(sl - #here is s#ill 'ork #o be done #o #es# #hese )om&e#ing h &o#heses*

Cogni#ion and Working Memor

9he in"l(en)e o" memor on &er)e&#ion is an e%am&le o" 'ha# )ogni#ive s)ien#is#s some#imes )all #o&$do'n &ro)essing- 'hi)h )on#ras#s 'i#h #he b(ild$(& o" &er)e&#ions "rom sensor &ro)essing- kno'n as bo##om$(& &ro)essing* Working memor - in shor#- si#s a# #he )rossroads o" bo##om$(& and #o&$do'n &ro)essing s s#ems and makes high$level #hinking and reasoning &ossible* S#e&hen Lossl n- a leading )ogni#ive s)ien#is#- &(#s i# #his 'a 0
Working memor *** )orres&onds #o #he a)#iva#ed in"orma#ion in long$#erm memories- #he in"orma#ion in shor#$#erm memories- and #he de)ision &ro)esses #ha# manage 'hi)h in"orma#ion is a)#iva#ed in #he long$#erm memories and re#ained in #he shor#$#erm memories **** 9his kind o" 'orking memor s s#em is ne)essar "or a 'ide range o" #asks- s()h as &er"orming men#al ari#hme#i)- reading- &roblem solving and * * * reasoning in general* All o" #hese #asks reB(ire no# onl some "orm o" #em&orar s#orage- b(# also in#er&la be#'een in"orma#ion #ha# is s#ored #em&oraril and a larger bod o" s#ored kno'ledge,1*

S#(dies )ond()#ed in #he 1230s b C*+* :a)obsen &rovide #he "o(nda#ion "or o(r (nders#anding o" #his &roblem,2* /e #rained monke s (sing some#hing )alled #he dela ed res&onse #ask* 9he monke sa# in a )hair and 'a#)hed #he e%&erimen#er &(# a raisin (nder one o" #'o obAe)#s #ha# 'ere side b side* A )(r#ain 'as #hen lo'ered "or a )er#ain amo(n#
42

2uchkin, 6! 0!, 1rafman, ,!, Cameron, 9! - Berndt, 2! 0! Working memor re#en#ion s s#ems0 a s#a#e o" a)#iva#ed long$#erm memor * )eha". )rain #ci. 2.- 702$723- dis)(ssion 723$777= 2003* Baddeley, A! 9he e&isodi) b(""er0 a ne' )om&onen# o" 'orking memor N Trends Cogn. S)i* 4417C423= 2000 9osslyn, 0M!, and 9oenig, (* 9et mind: The ne5 cogniti"e neuroscience8 Ne' 8ork0 Ma)millan= 1222* ,acobsen, C84 - ?issen, 'W! S#(dies o" )erebral "(n)#ion in &rima#es0 @7* 9he e""e)#s o" "ron#al lobe lesions on #he dela ed al#erna#ion habi# in monke s* %ournal of Comparati"e and Physiological Psychology 23- 101$12= 1237*

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Revised 17 November 2013 o" #ime (#he dela ) and #hen #he monke 'as allo'ed #o )hoose* @n order #o ge# #he raisin#he monke had #o remember no# 'hi)h obAe)# #he raisin 'as (nder b(# 'he#her #he raisin 'as (nder #he le"# or #he righ# obAe)#* Corre)# &er"orman)e- in o#her 'ords- reB(ired #ha# #he monke holds in i#s mind #he s&a#ial lo)a#ion o" #he raisin d(ring #he dela &eriod (d(ring 'hi)h #he &la ing "ield 'as hidden "rom vie')* A# ver shor# dela s (a "e' se)onds)- normal monke s did B(i#e 'ell- and &er"orman)e go# &redi)#abl 'orse as #he dela in)reased ("rom se)onds #o min(#es)* /o'ever- monke s 'i#h damage #o #he &re"ron#al )or#e% &er"ormed &oorl - even a# #he shor# dela s* Fn #he basis o" #his and resear)h #ha# "ollo'ed- #he &re"ron#al )or#e% has )ome #o be #ho(gh# o" as &la ing a role in #em&orar memor &ro)esses- &ro)esses #ha# 'e no' re"er #o as 'orking memor * Previo(sl a##en#ion has been dra'n #o #he role o" #he medial &re"ron#al )or#e% in #he e%#in)#ion o" emo#ional memor * @n )on#ras#- i# is #he lateral &re"ron#al )or#e% #ha# has mos# o"#en been im&li)a#ed in 'orking memor * 9he la#eral &re"ron#al )or#e% is believed #o e%is# onl in &rima#es and is )onsiderabl larger in h(mans #han in o#her &rima#es,3* @# is no# s(r&rising #ha# one o" #he mos# so&his#i)a#ed )ogni#ive "(n)#ions o" #he brain sho(ld involve #his region* @n re)en# ears- #he role o" #he la#eral &re"ron#al )or#e% in 'orking memor has been s#(died e%#ensivel b #he labora#ories o" :oaB(in +(s#er a# ICJA and Pa# <oldman$Raki) a# 8ale,4-,,-,.-,7 * 5o#h resear)hers have re)orded #he ele)#ri)al a)#ivi# o" la#eral &re"ron#al ne(rons 'hile monke s &er"ormed dela ed res&onse #asks and o#her #es#s reB(iring shor#$ #erm s#orage* 9he have sho'n #ha# )ells in #his region be)ome &ar#i)(larl a)#ive d(ring #he dela &eriods* @# is likel #ha# #hese )ells are a)#ivel involved in holding on #o #he in"orma#ion d(ring #he dela *

9he Pre"ron#al Cor#e% and Working Memor

9here is no' a )ri#i)al mass o" s#(dies #ha# "ind la#eral Pre"ron#al Cor#e% (P+C) a)#ivi# in h(mans d(ring dela #asks* ,3 +or e%am&le- in a "(n)#ional magne#i) resonan)e imaging ("MR@) s#(d (sing an o)(lomo#or dela #ask iden#i)al #o #ha# (sed in monke s#(dies- i# 'as observed #ha# no# onl #he "ron#al )or#e% a)#ivi# d(ring #he re#en#ion in#erval b(# also #he magni#(de o" #he a)#ivi# - )orrela#ed &osi#ivel 'i#h #he a))(ra) o" #he memor $g(ided sa))ade #ha# "ollo'ed la#er* 9his rela#ionshi& s(gges#s #ha# #he "ideli# o" #he a)#ivel main#ained lo)a#ion is re"le)#ed in #he dela $&eriod a)#ivi# ,2* 9h(s- #he e%is#en)e o" &ersis#en# ne(ral a)#ivi# d(ring blank
,3

3reuss, 5M! 6o ra#s have &re"ron#al )or#e%N 9he Rose$Woolse $Aker# &rogram re)onsidered* %ournal of Cogniti"e Neuroscience 7- 1$24= 122,* /uster, ,M! The prefrontal cortex. Ne' 8ork0 Raven= 1232)* 1oldman)2akic, 30* Cir)(i#r o" &rima#e &re"ron#al )or#e% and reg(la#ion o" behavior b re&resen#a#ional memor * @n andboo! of physiology. #ection 10 The ner"ous system. $ol. ,0 igher Functions of the )rain, +* Pl(m- ed* (5e#hesda- M60 Ameri)an Ph siologi)al So)ie# - &&* 373$417= 1237* 1oldman)2akic, 30* Working memor and #he mind* @n *ind and brain: +eadings from #cientific American maga:ine, W* /* +reeman- ed* (Ne' 8ork0 +reeman)- &&* ..$77= 1223* Wilson, /AW4 0calaidhe, 034 - 1oldman)2akic, 30! 6isso)ia#ion o" obAe)# and s&a#ial &ro)essing domains in &rima#e &re"ron#al )or#e%* #cience 2.0- 12,,$,3= 1223* Curtis, C! 8! - 678sposito, M! Persis#en# a)#ivi# in #he &re"ron#al )or#e% d(ring 'orking memor * Trends. Cogn. #ci. 7-41,C423= 2003* Curtis, C! 8!, 2ao, @! >! - 678sposito, M! Main#enan)e o" s&a#ial and mo#or )odes d(ring o)(lomo#or dela ed res&onse #asks! %. Neurosci. 24- 3244$32,2= 2004.

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Revised 17 November 2013 memor in#ervals o" dela #asks is a &o'er"(l em&iri)al "inding- 'hi)h lends s#rong s(&&or# "or #he h &o#hesis #ha# s()h a)#ivi# re&resen#s a ne(ral me)hanism "or #he a)#ive main#enan)e or s#orage o" #ask$relevan# re&resen#a#ions* 9he ne)essi# o" #he P+C "or #he a)#ive main#enan)e o" #ask$relevan# re&resen#a#ions has been demons#ra#ed b s#(dies #ha# have "o(nd im&aired &er"orman)e on dela #asks in monke s 'i#h sele)#ive lesions o" #he la#eral P+C .0-.1 . /o'ever- monke &h siolog s#(dies re)ording "rom o#her brain areas and h(man "MR@ s#(dies o" 'orking memor have also "o(nd #ha# #he P+C is no# #he onl region #ha# is a)#ive d(ring #he #em&orar re#en#ion o" #ask$relevan# in"orma#ion* +or e%am&le di""eren# brain regions are involved d(ring #he &er"orman)e o" #he o)(lomo#or dela ed res&onse #ask* S&e)i"i)all - di""eren# brain regions 'ere a)#ive de&ending on 'he#her #he #ask reB(ired #he #em&orar main#enan)e o" re#ros&e)#ive (e*g* &as# sensor even#s) or &ros&e)#ive (e*g* re&resen#a#ions o" an#i)i&a#ed a)#ion and &re&ara#or se#) )odes* 9his s#(d demons#ra#ed no# onl #ha# man di""eren# brain regions e%hibi# &ersis#en# ne(ral a)#ivi# d(ring a)#ive main#enan)e o" #ask$relevan# in"orma#ion- b(# also #ha# a (niB(e ne#'ork o" brain regions are re)r(i#ed de&ending on #he # &e o" in"orma#ion being a)#ivel main#ained* 9he "MR@ da#a also s(&&or# #he no#ion #ha# even 'i#hin #he domain o" s&a#ial in"orma#ion- se&arable ne(ral me)hanisms are engaged "or #he a)#ive main#enan)e o" Emo#or! &lans vers(s Es&a#ial! )odes* Moreover- given #ha# #he #ask onl reB(ired #he o)(lomo#or s s#em- i# is &robable #ha# dis#in)# ne(ral )ir)(i#r 'ill be re)r(i#ed 'hen #he mo#or a)# involves o#her modali#ies- s()h as s&ee)h or limb o(#&(#* .2 9h(s- #his is #he "irs# &ie)e o" eviden)e #ha# #he )on)e&# o" s&e)ialiGed b(""ers ("or- sa verbal vers(s s&a#ial in"orma#ion) ma no# ma& adeB(a#el on#o ne(ral ar)hi#e)#(re* Ra#her- #he "indings a&&ear more )onsis#en# 'i#h a s s#em in 'hi)h a)#ive main#enan)e involves #he re)r(i#men# o" #he same )ir)(i#r #ha# re&resen#s #he in"orma#ion i#sel"- 'i#h di""eren# )ir)(i#s "or di""eren# # &es o" s&a#ial in"orma#ion (e*g* vis(al vers(s o)(lomo#or)* Similar "indings e%is# 'hen #he Evis(al! )om&onen# o" 'orking memor is inves#iga#ed 'i#h ne(ro$s)ien#i"i) me#hods* +or e%am&le- in ano#her "MR@ s#(d .3 - #he &ar#i)i&an#s 'ere asked #o learn a series o" "a)es- ho(ses and "a)e$ho(se asso)ia#ions and #he 'ere s)anned 'hile &er"orming a dela ed ma#)h$#o$sam&le (6MS) and dela ed &aired $ asso)ia#e (6PA) #ask 'i#h #hese s#im(li* Res(l#s sho'ed #ha# dela $&eriod a)#ivi# 'i#hin )a#egor $sele)#ive in"erior #em&oral s(b$ regions re"le)#ed #he # &e o" in"orma#ion #ha# 'as being a)#ivel main#ained C #he "(si"orm g r(s sho'ed enhan)ed a)#ivi# 'hen &ar#i)i&an#s main#ained &revio(sl sho'n "a)es on 6MS #rials- and 'hen s(bAe)#s re)alled "a)es in res&onse #o a ho(se )(e on 6PA #rials* Jike'ise- #he &ara$hi&&o)am&al g r(s sho'ed enhan)ed a)#ivi# 'hen &ar#i)i&an#s main#ained &revio(sl sho'n ho(ses on 6MS #rials and 'hen #he re)alled ho(ses in res&onse #o a "a)e )(e on 6PA #rials*

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Bauer, 2! '! - /uster, ,! M! 6ela ed$ma#)hing and dela ed$res&onse de"i)i# "rom )ooling dorsola#eral &re"ron#al )or#e% in monke s* ;. %. 2xp. Psychol. 5 20- 223$302= 127.* /unahashi, 0!, Bruce, C! ,! - 1oldman)2akic, 3! 0! 6orsola#eral &re"ron#al lesions and o)(lomo#or dela ed$res&onse &er"orman)e0 eviden)e "or mnemoni) Ms)o#omasM* %. Neurosci. 13- 1472$1427= 1223* 'ickok, 1!, Buchsbaum, B!, 'umphries, C! - Muftuler- 9* A(di#or $mo#or in#era)#ion revealed b "MR@0 s&ee)h- m(si)- and 'orking memor in area S&#* %. Cogn. Neurosci. #E, .73$.32= 2003* 2anganath, C!, Cohen, M! F!, 6am, C! - 678sposito, M* @n"erior #em&oral- &re"ron#al- and hi&&o)am&al )on#rib(#ions #o vis(al 'orking memor main#enan)e and asso)ia#ive memor re#rieval* %. Neurosci. 24- 3217$322,= 2004*

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Revised 17 November 2013 9hese "MR@ "indings are )onsis#en# 'i#h several monke ne(ro$&h siologi)al s#(dies 'hi)h have also sho'n #ha# #em&oral lobe ne(rons e%hibi# &ersis#en# s#im(l(s$sele)#ive a)#ivi# in #asks reB(iring #he a)#ive main#enan)e o" vis(al obAe)# in"orma#ion a)ross shor# dela s .4- .,..* Again- like s&a#ial and mo#or )odes- a)#ive main#enan)e o" vis(al s#im(li is media#ed b #he a)#iva#ion o" )or#i)al regions #ha# also s(&&or# &ro)essing o" #ha# in"orma#ion&er)e&#(al in #his )ase* Work on ne(ro$s)ien#i"i) s#(dies o" verbal 'orking memor - 'hi)h has been mos# e%#ensivel s#(died b behavio(ral me#hods= (7allar K Shalli)e- &rovide a similar vie' regarding #he ne(ral me)hanisms (nderl ing 'orking memor .7 )* Consis#en#l &er"orman)e on #asks #ha# #a& #he E&honologi)al loo&!- as )on)e&#(aliGed b 5addele engage a se# o" brain regions #ha# are #ho(gh# #o be involved in &honologi)al &ro)essing* +or e%am&le- (sing "(n)#ional ne(ro$imaging #e)hniB(es d(ring verbal 'orking memor #asks- #he le"# in"erior &arie#al lobe- &os#erior in"erior "ron#al g r(s (5ro)a!s area)- &remo#or )or#e% and #he )erebell(m are # &i)all a)#iva#ed* .3- .2

0chematic of Baddeley7s Model

.4

Miyashita, >! - Chang, '! 0* Ne(ronal )orrela#e o" &i)#orial shor#$#erm memor in #he &rima#e #em&oral )or#e%* Nature 331-.3$70= 1233* Miller, 8! 9!, <i, <! - 6esimone, 2* A)#ivi# o" ne(rons in an#erior in"erior #em&oral )or#e% d(ring a shor#$#erm memor #ask* %. Neurosci. #%, 14.0$1473= 1223* ?akamura, 9! - 9ubota, 9! Mnemoni) "iring o" ne(rons in #he monke #em&oral &ole d(ring a vis(al re)ogni#ion memor #ask* %. Neurophysiol. 74- 1.2$173= 122,* @allar, 1! - 0hallice, 5! Neuropsychological im&airmen#s of short<term memory. CambridgeIL0 Cambridge Iniversi# Press= 1220* 3aulesu, 8!, /rith, C! 6! - /rackowiak, 2! 0! 9he ne(ral )orrela#es o" #he verbal )om&onen# o" 'orking memor * Nature 3.2-342$34,= 1223 Awh, 8!, ,onides, ,!, 0mith, 8! 8!, 0chumacher, 8! '!, 9oeppe, 2! A! - 9at., 0! 6isso)ia#ion o" s#orage and rehearsal in verbal 'orking memor 0 eviden)e "rom P;9* Psycho /. #ci. 7- 2,$3= 122.*

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Revised 17 November 2013

Sketch of proposed circuitry supporting phonological loop

(A"#er ;dm(nd 5lair 5olles) /o'ever- is #his ne#'ork o" brain regions also res&onsible "or #he a)#ive main#enan)e o" non$&honologi)al lang(age re&resen#a#ions (e*g* le%i)al$seman#i))*70 +or vis(al 'ord re)ogni#ion- a "(n)#ionall s&e)ialiGed &ro)essing s#ream is #ho(gh# #o e%is# 'i#hin in"erior #em&oral )or#e%- re&resen#ing vis(al 'ords a# in)reasingl higher levels o" abs#ra)#ion along a &os#erior$#o$an#erior a%is 71* @n#ra)ranial ele)#ro&h siologi)al re)ordings- "or e%am&le- (Nobre et al 72 ) sho' #ha# &os#erior in"erior #em&oral )or#e% di""eren#ia#es le##er s#rings "rom non$ling(is#i) )om&le% vis(al obAe)#s* 5rain a)#ivi# in more an#erior in"erior #em&oral )or#i)al regions- in )on#ras#dis#ing(ishes 'ords "rom non$'ords and is a""e)#ed b #he seman#i) )on#e%# o" 'ordsindi)a#ing #ha# an#erior in"erior #em&oral )or#e% holds more elabora#e ling(is#i) re&resen#a#ions (see also Marslen$Wilson K 9 ler 73 and Pa##erson 74)* 9o demons#ra#e #ha# #here is dis#in)# ne(ral )ir)(i#r s(&&or#ing #he a)#ive main#enan)e o" non$&honologi)al lang(age re&resen#a#ions- 6!;s&osi#o 7, e%&lored #he role o" lang(age
70

<e&ical semantics is #he s#(d o" ho' and 'ha# #he 'ords o" a lang(age deno#e* Words ma ei#her be #aken #o deno#e #hings in #he 'orld or )on)e&#s- de&ending on #he &ar#i)(lar a&&roa)h #o le%i)al seman#i)s* Cohen, <! - 6ehaene, 0* S&e)ialiGa#ion 'i#hin #he ven#ral s#ream0 #he )ase "or #he vis(al 'ord "orm area* Neuroimage 22-4..C47.= 2004* ?obre, A! C!, Allison, 5! - McCarthy, 1* Word re)ogni#ion in #he h(man in"erior #em&oral lobe* Nature 372- 2.0$2.3= 1224* Marslen) Wilson, W 6! - 5yler, <! 9! Mor&holog - lang(age and #he brain0 #he de)om&osi#ional s(bs#ra#e "or lang(age )om&rehension* Phil. Trans. +. #oc. ) 3.2- 323$33.= 2007* 3atterson, 9! 9he reign o" # &i)ali# in seman#i) memor * Phil. Trans. +. #oc. ) 3.2- 313$321= 2007* 678sposito, M! +rom )ogni#ive #o ne(ral models o" 'orking memor * Phil. Trans. +. #oc. )* 22 Ma 2007= vol! %;$ no! #*G# 7.1$772

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Revised 17 November 2013 regions 'i#hin #he le"# in"ero$#em&ora1 )or#e% (@9C) #ha# are involved in vis(al 'ord re)ogni#ion and 'ord$rela#ed seman#i)s* Ising #MR@- he "irs# lo)aliGed a vis(al E'ord "orm! area 'i#hin in"erior #em&oral )or#e% area and #hen demons#ra#ed #ha# #his area 'as involved in #he a)#ive main#enan)e o" vis(all &resen#ed 'ords d(ring a dela #ask 7.* S&e)i"i)all - he "o(nd #ha# #his area 'as re)r(i#ed more "or #he a)#ive main#enan)e o" 'ords #han &se(do$'ords (i*e* or#hogra&hi)all legal and &rono(n)eable non$'ords)* Main#enan)e o" &se(do$'ords sho(ld no# eli)i# s#rong s(s#ained a)#iva#ion in s()h brain regions- as no s#ored re&resen#a#ions &re$e%is# "or #hese i#ems* 9hese res(l#s s(gges# #ha# verbal 'orking memor ma be )on)e&#(aliGed as involving s(s#ained a)#iva#ion o" all relevan# &re$e%is#ing )or#i)al lang(age (&honologi)al- le%i)al or seman#i)) re&resen#a#ions* <aGGale et al - have (sed a dela #ask #o dire)#l s#(d #he ne(ral me)hanisms (nderl ing #o&$do'n mod(la#ion b inves#iga#ing #he &ro)esses involved 'hen &ar#i)i&an#s 'ere reB(ired #o enhan)e relevan# and s(&&ress irrelevan# in"orma#ion*77 6(ring ea)h #rial- &ar#i)i&an#s observed seB(en)es o" #'o "a)es and #'o na#(ral s)enes &resen#ed in a randomiGed order* 9he #asks di""ered in #he ins#r()#ions in"orming #he &ar#i)i&an#s ho' #o &ro)ess #he s#im(li0 (i) (ii) (iii) remember "a)es and ignore s)enesremember s)enes and ignore "a)es- or &assivel vie' "a)es and s)enes 'i#ho(# a##em&#ing #o remember #hem*

@n ea)h #ask- #he &eriod in 'hi)h #he )(e s#im(li 'ere &resen#ed 'as balan)ed "or bo##om$(& vis(al in"orma#ion- #h(s allo'ing (s #o &robe #he in"l(en)e o" goal$dire)#ed behavio(r on ne(ral a)#ivi# (#o&$do'n mod(la#ion)* @n #he #'o memor #asks- #he en)oding o" #he #ask$relevan# s#im(li reB(ires sele)#ive a##en#ion and #h(s &ermi#s #he disso)ia#ion o" &h siologi)al meas(res o" enhan)emen# and s(&&ression rela#ive #o #he &assive baseline* 9here a&&ears #o be a# leas# #'o # &es o" #o&$do'n signal- one #ha# serves #o enhan)e #ask$relevan# in"orma#ion and ano#her #ha# serves #o s(&&ress #ask$relevan# in"orma#ion* @# is 'ell do)(men#ed #ha# #he nervo(s s s#em (ses in#erleaved inhibi#or and e%)i#a#or me)hanisms #hro(gho(# #he ne(ro$a%is (e*g* s&inal re"le%es- )erebellar o(#&(#s and basal ganglia movemen# )on#rol ne#'orks)* 9h(s- i# ma no# be s(r&rising #ha# enhan)emen# and s(&&ression me)hanisms ma e%is# #o )on#rol )ogni#ion*73-72 5 genera#ing )on#ras# via bo#h enhan)emen#s and s(&&ressions o" a)#ivi# magni#(de and &ro)essing s&eed- #o&$ do'n signals bias #he likelihood o" s())ess"(l re&resen#a#ion o" relevan# in"orma#ion in a )om&e#i#ive s s#em* 9ho(gh i# has been &ro&osed #ha# #he P+C &rovides a maAor so(r)e o" #he # &es o" #o&$ do'n signals #ha# <aGGale has des)ribed- #his h &o#hesis largel origina#es "rom s(gges#ive

7.

/iebach, C! ,!, 2issman, ,! - 678sposito, M! Mod(la#ion o" in"ero$#em&oral )or#e% a)#iva#ion d(ring verbal 'orking memor main#enan)e* Neuron ,1- 2,1$2.1= 200.* 1a..aley, A, Cooney, #! W, Mc8voy, 9!, 9night, 2! 5! - 678sposito, M* 9o&$do'n enhan)emen# and s(&&ression o" #he magni#(de and s&eed o" ne(ral a)#ivi# * %. Cogn. Neurosci. 17- ,07$ ,17* 200, (doi0 10*11.2103232220,3272,22) 9night, 2! 5!, 0taines, W 2!, 0wick, 6! - Chao, <! <! Pre"ron#al )or#e% reg(la#es inhibi#ion and e&citation in distributed neural networks! Acta. Psycho!. (Amst) #D#, #EH)#=G4 #HHH! 0himamura, A! 3* 9he role o" #he &re"ron#al )or#e% in d nami) "il#ering* Psychobiology =>, 207$ 213= 2000*

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Revised 17 November 2013 "indings ra#her #han dire)# em&iri)al eviden)e* /o'ever- a "e' s#(dies lend dire)# )a(sal s(&&or# #o #his h &o#hesis* See Also 30 * Clearl - #here are o#her areas o" m(l#imodal )or#e% s()h as &os#erior &arie#al )or#e%- and #he hi&&o)am&(s- #ha# )an also be #he so(r)e o" #o&$do'n signals* +or e%am&le- #he hi&&o)am&(s has been &ro&osed #o be s&e)ialiGed "or Era&id learning o" arbi#rar in"orma#ion 'hi)h )an be re)alled in #he servi)e o" )on#rolled &ro)essing!* 31 Moreoverin&(# "rom brains#em ne(ro$mod(la#or s s#ems &robabl &la s a )ri#i)al role in mod(la#ing goal$dire)#ed behavio(r* 32 +or e%am&le- #he do&aminergi) s s#em &robabl &la s a )ri#i)al role in )ogni#ive )on#rol &ro)esses* 33 S&e)i"i)all - i# is &ro&osed #ha# &hasi) b(rs#s o" do&aminergi) ne(rons ma be )ri#i)al "or (&da#ing )(rren#l a)#iva#ed #ask$relevan# re&resen#a#ions 'hereas #oni) do&aminergi) a)#ivi# serves #o s#abiliGe s()h re&resen#a#ions* 34 ;m&iri)al s#(dies in animals and man have &rovided s(&&or# "or a role o" do&amine in 'orking memor * 3,- 3. @" 'orking memor main#enan)e &ro)esses re"le)# #he &rolonged a)#iva#ion o" #he same brain regions #ha# s(&&or# online &ro)essing- eviden)e "or )or#i)al a)#ivi# in #he absen)e o" s#im(li sho(ld be eviden# no# onl in asso)ia#ion )or#e% b(# also 'i#hin &rimar )or#i)al regions* 9his indeed is #he )ase as s()h e""e)#s have been observed in &rimar ol"a)#or 37vis(al 33- 32 and a(di#or )or#e% 20- 21* 9h(s- #he ne(ro$s)ien#i"i) da#a &resen#ed here is )onsis#en# 'i#h mos#- i" no# all- ne(ral &o&(la#ions being able #o re#ain in"orma#ion #ha# )an be a))essed and ke&# a)#ive over several se)onds- via &ersis#en# ne(ral a)#ivi# in #he servi)e o" goal$dire)#ed behavio(r*

30

@uilleumier, 3! - 6river, ,! Mod(la#ion o" vis(al &ro)essing b a##en#ion and emo#ion0 'indo's on )a(sal in#era)#ions be#'een h(man brain regions* Phil. Trans. +. #oc. ) 3.2- 337$3,,= 2007*) (72eilly, 2!C!, Braver, 5! 0! - Cohen, ,! 6! A biologi)all based )om&(#a#ional model o" 'orking memor * @n *odels of 5or!ing memory: mechanisms of acti"e maintenance and executi"e control (eds Mi ak'- A* K Shah- P*)* Cambridge- IL0 Cambridge Iniversi# Press* 1222* 2obbins, 5! W! Shi"#ing and s#o&&ing0 "ron#o$s#ria#al s(bs#ra#es- ne(ro)hemi)al mod(la#ion and)lini)al im&li)a#ions* Phil. Trans. +. #oc* 5 3.2- 217$232= 2007* (doi0 10*10231 rs#b*2007*2027) Cools, 2! - 2obbins, 5! W! Chemis#r o" #he ada&#ive mind* Phil. Trans. +. #oc* A 3.2- 2371$ 2333*(doi01 F* 10231rs#a*2004*14.3) 2004* 6urstewit., 6!, 0eamans, I! 9! - 0ejnowski, 5! I* 6o&amine$media#ed s#abiliGa#ion o" dela $ &eriod a)#ivi# in a ne#'ork model o" &re"ron#al )or#e%* %. Neurophysiol* 33- 1733$17,00 2000* 0awaguchi, 5 * 9he e""e)#s o" do&amine and i#s an#agonis#s on dire)#ional dela $ &erioda)#ivi# o" &re"ron#al ne(rons in monke s d(ring an o)(lomo#or dela ed$res&onse #ask* Neurosci. +es* 41- 11,$123= 2001* (doi0 1F*101.1S01.3$0102(01)00270$O) 1ibbs, 0! 8! - 678sposito, M* A "(n)#ional MR@ s#(d o" #he e""e)#s o" bromo)ri&#ine- ado&amine re)e&#or agonis#- on )om&onen# &ro)esses o" 'orking memor * Psychopharmacology (5erlin) 130- .44$ .,3= 200,* (doi0@F*10071 s00213$00,$0077$,) Ielano, c!, Bensafi, M!, 3orter, ,!, Mainland, ,!, ,ohnson, B!, Bremner, 8!, 5elles, C!, 9han, 2! 0obel, ?! A##en#ional mod(la#ion in h(man &rimar ol"a)#or )or#e%* Nat. Neurosci. G, 114$120= 200,* 9lein, I!, 3aradis, A! <!, 3oline, ,! B!, 9osslyn, 0! M! - <e Bihan, 6* 9ransien# a)#ivi# in #he h(man )al)arine )or#e% d(ring vis(al$men#al imager 0 an even#$rela#ed "MR@ s#(d * %. Cogn. Neurosci. 12(S(&&l* 2)- 1,$23= 2000* 0inger, W! - 1ray, C! M! 7is(al "ea#(re in#egra#ion and #he #em&oral )orrela#ion h &o#hesis* Annu. +e". Neurosci. 13-,,,$,3.= 122,* Calvert, 1! A!, Bullmore, 8! 5!, Brammer, M! ,!, Campbell, 2!, Williams, 0! C!, Mc1uire, J3! 9!, Woodruff, 3! W, Iversen, 0! 6! - 6avid, A! 0! A)#iva#ion o" a(di#or )or#e% d(ring silen# li&reading* #cience 27.- ,23$,2.= 1227* 9raemer, 6! ,!, Macrae, C! ?!, 1reen, A! 8! - 9elley, W M! M(si)al imager 0 so(nd o" silen)e a)#iva#es a(di#or )or#e%* Nature *%*,#EG!= 200,*

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Revised 17 November 2013

Cerebellar +(n)#ion in A(#ism K 5rain @nA(r and #he Role o" #he Cerebell(m in Cogni#ion and A""e)#
9he )oe%is#en)e o" bo#h )erebral and )erebellar maldevelo&men# ma e%&lain 'h #he im&airmen#s in higher )ogni#ive "(n)#ions in a(#ism are &ervasive and &ersis#en# a)ross #he li"e s&an* A &os#na#al &eriod o" e%#reme maldevelo&men# o" #hese #'o maAor brain s#r()#(res 'ill almos# )er#ainl be mani"es# in aberran# behavio(ral e%&ression- and i# is d(ring #his earl &eriod (# &i)all 12 #o 24 mon#hs) #ha# &aren#s mos# o"#en "irs# e%&ress )on)ern abo(# #heir )hild!s develo&men#* 22 6(ring #his develo&men#al &eriod- #he h(man brain (ndergoes ra&id s na&#ogenesis- e%&ansion o" dendri#i) and a%on arbors- and sele)#ion o" 'hi)h ne(ronal elemen#s #o kee& or elimina#e* 23- 24- 2,- 2. @n #he normall develo&ing brain- sha&ing o" ne(ral ar)hi#e)#(re and )onne)#ivi# is #heorised #o be signi"i)an#l in"l(en)ed or dire)#ed b "(n)#ional ne(ral a)#ivi# driven b learning and e%&erien)e* 27- 23 @n #he a(#is#i) brain- ho'ever- resear)hers s&e)(la#e #ha# gro'#h and elabora#ion o" ne(ral ar)hi#e)#(re and )onne)#ivi# o))(rs &rema#(rel and 'i#ho(# being g(ided b "(n)#ional e%&erien)es and ada&#ive learning= #ha# is- in earl li"e #he a(#is#i) brain e%hibi#s &rema#(re gro'#h 'i#ho(# g(idan)e* 22 @n addi#ion- #he )erebell(m and )er#ain )erebral regions ma &er"orm analogo(s- &ossibl )om&lemen#ar - "(n)#ions* +or ins#an)e- in #he normal brain- )erebellar )or#e% is a)#iva#ed b #asks #ha# )ommonl a)#iva#e "ron#al )or#e%- s()h as #asks involving 'orking memor a##en#ion- or seman#i) asso)ia#ion* 100- 101- 102 Ad(l#s 'i#h )erebellar lesions sho' im&aired &er"orman)e on similar "ron#al lobe #asks- in)l(ding #es#s o" so(r)e memor and e%e)(#ive "(n)#ions (e*g*- shi"#ing a##en#ion- )ogni#ive &lanning- and 'orking memor )* 103- 104 So- #he

22

Co& A, Charman 5, Baron)Cohen 0, et al! A(#ism s&e)#r(m disorders a# 20 and 42 mon#hs o" age0 s#abili# o" )lini)al and A6@$R diagnosis* % Child Psychol Psychiatry 1222=,0712H732* 0chade ,3, van 1roenigen WB* S#r()#(ral organiGa#ion o" #he h(man )erebral )or#e%* @*Ma#(ra#ion o" #he middle "ron#al g r(s* Acta Anat 12.1=47072H111* 'uttenlocher 32, 6abholkar A0* Regional di""eren)es in s na&#ogenesis in h(man )erebral)or#e%* % Comp Neurol 1227= 33701.7H173* Kuart. 02, 0ejnowski 5,* 9he ne(ral basis o" )ogni#ive develo&men#0 a )ons#r()#ivis# mani"es#o* )eha" )rain #ci 1223=200 ,37H,2.* Courchesne 8, Chisum ', 5ownsend ,! Ne(ral a)#ivi# $de&enden# brain )hanges indevelo&men#0 im&li)a#ions "or &s )ho&a#holog * 6e" Psychopathol 1224=.0.27H722* Kuart. 02, 0ejnowski 5,* 9he ne(ral basis o" )ogni#ive develo&men#0 a )ons#r()#ivis# mani"es#o* )eha" )rain #ci 1223=200 ,37H,2.* Courchesne 8, Chisum ', 5ownsend ,! Ne(ral a)#ivi# de&enden# brain )hanges indevelo&men#0 im&li)a#ions "or &s )ho&a#holog * 6e" Psychopathol 1224=.0.27H722* Cohen, <! - 6ehaene, 0* S&e)ialiGa#ion 'i#hin #he ven#ral s#ream0 #he )ase "or #he vis(al 'ord "orm area* Neuroimage 22-4..C47.= 2004* Allen 1, Bu&ton 2B, Wong 8C, Courchesne 8* A##en#ional a)#iva#ion o" #he )erebell(m inde&enden# o" mo#or involvemen#* #cience 1227=27,01240H1243* 6esmond ,8, 1abrieli ,6, Wagner A6, 1inier B<, 1lover 1'! Job(lar &a##erns o" )erebellar a)#iva#ion in verbal 'orking memor and "inger$#a&&ing #asks as revealed b "(n)#ional MR@* % Neurosci 1227= 1702.7,H2.3,* 2aichle M8, /ie. ,A, @ideen 5( et al * Pra)#i)e$rela#ed )hanges in h(man brain "(n)#ionalana#om d(ring nonmo#or learning* Cereb Cortex 1224= 403H2.* Akshoomoff ?A! @n#ramodali# shi"#ing a##en#ion in )hildren 'i#h damage #o #he )erebell(m* %Cogn Neurosci 1224= .0333H 322* 0chmahmann ,6, 0herman ,C* 9he )erebellar )ogni#ive a""e)#ive s ndrome* )rain 1223=1210,.1H,72*

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Revised 17 November 2013 &resen)e o" bo#h )erebral and )erebellar de"e)#s "rom #he earlies# s#ages o" )ogni#ive develo&men# 'o(ld likel res(l# in severe and &ersis#en# "(n)#ional de"i)i#s* A )hild 'i#h im&aired verbal 'orking memor ma have di""i)(l# "ollo'ing ins#r()#ions and be seen as disobedien#* A )hild 'ho has di""i)(l# &la)ing in"orma#ion in#o longer$#erm memor ma "orge# &revio(s )onversa#ions 'i#h &eers and so a&&ear #o be (nin#eres#ed&ossibl res(l#ing in so)ial reAe)#ion or so)ial 'i#hdra'al* A )hild 'i#h &ros&e)#ive memor &roblems ma be seen as #r ing #o avoid si#(a#ions* ChildrenDs memor di""i)(l#ies )an be a signi"i)an# so(r)e o" s#ress "or #heir "amilies* 10, S)hmahmann and Sherman (1223) s#(died 20 ad(l# &a#ien#s 'i#h disease )on"ined #o #he )erebell(m- eval(a#ing #he na#(re and severi# o" #he )hanges in ne(rologi)al and men#al "(n)#ion* 9he "o(nd behavio(ral )hanges #o be )lini)all &rominen# in &a#ien#s 'i#h lesions involving #he &os#erior lobe o" #he )erebell(m and #he vermis* 9he )hanges 'ere )hara)#eriGed b #he "ollo'ing0 im&airmen# o" e%e)(#ive "(n)#ions s()h as &lanningse#$shi"#ingverbal "l(en) abs#ra)# reasoning and 'orking memor = di""i)(l#ies 'i#h s&a#ial )ogni#ion in)l(ding vis(o$s&a#ial organiGa#ion and memor = &ersonali# )hange 'i#h bl(n#ing o" a""e)# or disinhibi#ed and ina&&ro&ria#e behavio(r= and lang(age de"i)i#s in)l(ding agramma#ism and d s&rosodia*

9he inves#iga#ors named #his ne'l de"ined )lini)al en#i# #he >)erebellar )ogni#ive a""e)#ive s ndrome?* 9he s(gges#ed #he )ons#ella#ion o" de"i)i#s is indi)a#ive o" disr(&#ion o" #he )erebellar mod(la#ion o" ne(ral )ir)(i#s #ha# link &re"ron#al- &os#erior &arie#als(&erior #em&oral and limbi) )or#i)es 'i#h #he )erebell(m* 10. S)o## et al* e%amined #he role o" #he )erebell(m in #he develo&ing )ogni#ive &ro"iles o" )hildren 'i#h )erebellar #(mo(rs and observed an asso)ia#ion be#'een grea#er damage #o righ# )erebellar s#r()#(res and a &la#ea(ing in verbal and1or li#era) skills* @n )on#ras#grea#er damage #o le"# )erebellar s#r()#(res 'as asso)ia#ed 'i#h dela ed or im&aired nonverbal1s&a#ial skills* Jong$#erm )ogni#ive develo&men# o" #he )hildren s#(died #en#a#ivel s(&&or#ed a role "or #he )erebell(m in learning1develo&men#* 9he a(#hors s(gges#ed #ha# la#eraliGed )erebellar damage ma sele)#ivel im&air #he develo&men# o" )ogni#ive "(n)#ions s(bserved b #he )on#ra$la#eral )erebral hemis&here* 107 +rom a s#(d b Mo##agh et al*- ana#omi)al da#a s(gges#ed )rossed re)i&ro)al )onne)#ions be#'een #he )erebell(m and higher order )or#i)al asso)ia#ion areas* @n a righ#$handed vol(n#eer- #he inves#iga#ors "o(nd an a)#iva#ion o" #he le"# "ron#o$&arie#al )or#e% and in #he righ# )erebellar hemis&here= 'hile in #he le"#$handed vol(n#eer #he a)#iva#ion 'as seen in #he righ# "ron#o$&arie#o$#em&oral )or#e% and #he le"# )erebellar hemis&here* 9hese ini#ial res(l#s demons#ra#e #ha# )erebellar a)#iva#ion is )on#ra$ la#eral #o #he a)#iva#ion o" #he "ron#al )or#e% even (nder )ondi#ions o" di""eren# lang(age
10,

2ivara, ,! B!, /ay, 1!, ,affe, 9!, 3olissar, ?!, 0hurtleff, '!, - Martin, 9! Predi)#ors o" "amil "(n)#ioning one ear "ollo'ing #ra(ma#i) brain inA(r in )hildren* Archi"es of Physical *edicine and +ehabilitation* 73- 322$210= 1222* 0chmahmann, ,6 - 0herman, ,C! 9he )erebellar )ogni#ive a""e)#ive s ndrome*)rain 1210,.1$ ,72= 1223* 0cott 2B4 0toodley4 C,, Anslow 34 3aul C4 0tein ,/4 0ugden 8M - Mitchell C6! Ja#eraliGed )ogni#ive de"i)i#s in )hildren "ollo'ing )erebellar lesions* 6e"elopmental *edicine and ChildNeurology. 43(10)0.3,$.21= 2001*

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Revised 17 November 2013 dominan)e* 9he &rovide eviden)e "or #he h &o#hesis o" a la#eraliGed organiGa#ion o" #he )erebell(m )rossed #o #he )erebral hemis&heres in s(&&or#ing higher )ogni#ive "(n)#ion* 103 9hese "indings are en#irel )ongr(en# 'i#h #he "indings o" Ch(gani e# al* indi)a#ing im&airmen# o" den#a#o$#halamo$)or#i)al &a#h'a s- &rimaril "rom #he righ# den#a#e #o le"# )erebral hemis&here- in o(ng a(#is#i) bo s* 102 5addele !s original advan)e (/dem- 123.) 'as #o move (s "rom #he )on)e&# o" shor#$#erm memor #ha# a))o(n#ed onl "or #he s#orage o" re&resen#a#ions- #o #he )on)e&# o" 'orking memor as a m(l#i$)om&onen# s s#em #ha# allo's "or bo#h s#orage and &ro)essing o" #em&oraril a)#ive re&resen#a#ions* Jike'ise- Jogie 110 )onsidered 'orking memor as a Emen#al 'orks&a)e! #ha# )anno# onl hold b(# is also able #o mani&(la#e a)#iva#ed re&resen#a#ions* @" 'orking memor is de"ined as sim(l#aneo(s s#orage and &ro)essing- #hen #hese #asks 'o(ld &robabl be )onsidered #o assess shor#$#erm memor ra#her #han 'orking memor * 9h(s- i# is &robabl "air #o sa #ha# #he )on)e&# o" 'orking memor as a non$(ni#ar s s#em #ha# allo's "or bo#h s#orage and &ro)essing has gained &o&(lar a))e&#an)e* /o'everless &rogress has been made regarding #he ne(ral me)hanisms (nderl ing #he E&ro)essing! )om&onen# o" 'orking memor as )om&ared 'i#h #he Es#orage! )om&onen# 111 (al#ho(gh see Pe#rides112) C(r#is et al - (2004- idem) also &ro&ose #ha# an &o&(la#ion o" ne(rons 'i#hin &rimar or (nimodal asso)ia#ion )or#e% )an e%hibi# &ersis#en# ne(ronal a)#ivi# - 'hi)h serves #o a)#ivel main#ain #he re&resen#a#ions )oded b #hose ne(ronal &o&(la#ions* Areas o" m(l#imodal )or#e%- s()h as P+C and &arie#al )or#e%- 'hi)h are in a &osi#ion #o in#egra#e re&resen#a#ions #hro(gh )onne)#ivi# #o (nimodal asso)ia#ion )or#e%- are also )ri#i)all involved in #he a)#ive main#enan)e o" #ask$relevan# in"orma#ion (see also 5(rgess et al 113= S#(ss K Ale%ander* 114 Miller K Cohen 11, have &ro&osed #ha# in addi#ion #o #he re)en# sensor in"orma#ionin#egra#ed re&resen#a#ions o" #ask )on#ingen)ies and even abs#ra)# r(les (e*g* i" #his obAe)# C #hen #his la#er res&onse) are also main#ained in #he P+C* 9his is similar #o 'ha# +(s#er 11. has long em&hasiGed- namel #ha# #he P+C is )ri#i)all res&onsible "or #em&oral in#egra#ion and #he media#ion o" even#s #ha# are se&ara#ed in #ime b(# )on#ingen# on one
103

Mottaghy /M4 0hah ?,4 9rause B,4 0chmidt 64 'alsband A4 ,ancke < - Muller)1artner 'W! Ne(ronal )orrela#es o" en)oding and re#rieval in e&isodi) memor d(ring a &aired$'ord asso)ia#ion learning #ask0 a "(n)#ional magne#i) resonan)e imaging s#(d * 2xperimental )rain +esearch 1230332$342= 1222 Chugani 6C4 Musik (4 2othermal 24 Behen M4 Chakraborty 34 Mangner 54 da 0ilva 8A Chugoni '5! Al#ered sero#onin s n#hesis in #he den#a#$o#halamo$)or#i)al &a#h'a in a(#is#i) bo s* Annals of Neurology* 420...$..2= 1227 <ogie, 2! '* 7is(o$s&a#ial 'orking memor * /ove- IL0 ;rlba(m= 122,* 6 8sposito, M* +rom Cogni#ive #o ne(ral models o" 'orking memor * @n 6river- := /aggard- P= K Shalli)e- 9* *ental Processes in the uman )rain* FIP- Ne' 8ork= 2007* 3etrides, M! +ron#al lobes and 'orking memor 0 eviden)e "rom inves#iga#ions o" #he e""e)#s o" )or#i)al e%)isions in nonh(man &rima#es* @n andboo! of Neuropsychology- vol* 2 (eds +* 5oller K :* <ra"man)- &&* ,2$34* Ams#erdam- 9he Ne#herlands0 ;lsevier S)ien)e 5*8= 1224* Burgess, 3! W, 1ilbert, 0! ,! - 6umontheil, I! +(n)#ion and lo)aliGa#ion 'i#hin ros#ral &re"ron#al )or#e% (area 10)* Phil. Trans. +. #oc* 3 3.2-337$322= 2007* 0tuss, 6! 5! - Ale&ander, M! 3* @s #here a d se%e)(#ive s ndromeN Phil. Trans. +. #oc* 5 3.2- 201$ 21,= 2007* Miller, 8! 9! - Cohen, ,! 6* An in#egra#ive #heor o" &re"ron#al )or#e% "(n)#ion* Annu. +e".-, Neurosci. 1.7$202= 2001* /uster, ,! 9he &re"ron#al )or#e%0 ana#om - &h siolog - and ne(ro&s )holog o" #he "ron#al lobes* Ne' 8ork- N80 Raven Press= 1227*

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Revised 17 November 2013 ano#her* @n #his 'a - #he P+C ma e%er# E)on#rol! in #ha# #he in"orma#ion i# re&resen#s )an bias #he &os#erior (nimodal asso)ia#ion )or#e% in order #o kee& ne(ral re&resen#a#ions o" behavio(rall relevan# sensor in"orma#ion a)#iva#ed 'hen #he are no longer &resen# in #he e%#ernal environmen#* 117- 113- 112- 120 @n a real 'orld e%am&le- 'hen a &erson is looking a# a )ro'd o" &eo&le- #he vis(al s)ene &resen#ed #o #he re#ina ma in)l(de a m riad o" angles- sha&es- &eo&le and obAe)#s* /o'ever- i" #ha# &erson is a &oli)e o""i)er looking "or an armed robber es)a&ing #hro(gh #he )ro'd- some me)hanism o" s(&&ressing irrelevan# vis(al in"orma#ion 'hile enhan)ing #ask$ relevan# in"orma#ion is ne)essar "or an e""i)ien# and e""e)#ive sear)h* 9h(s- ne(ral a)#ivi# #hro(gho(# #he brain #ha# is genera#ed b in&(# "rom #he o(#side 'orld ma be di""eren#iall enhan)ed or s(&&ressed- &res(mabl "rom #o&$do'n signals emana#ing "rom in#egra#ive brain regions s()h as P+C- based on #he )on#e%# o" #he si#(a#ion* So- in #his "orm(la#ion- #he &ro)essing )om&onen# o" 'orking memor is #ha# #he )on#rol o" a)#ivel main#ained re&resen#a#ions 'i#hin &rimar and (nimodal asso)ia#ion )or#e% s#ems "rom #he re&resen#a#ional &o'er o" m(l#imodal asso)ia#ion )or#e%- s()h as #he P+C- &arie#al )or#e% and1or hi&&o)am&(s* @" #he P+C- "or e%am&le- s#ores #he r(les and goals- #hen #he a)#iva#ion o" s()h P+C re&resen#a#ions 'ill be ne)essar 'hen behavio(r m(s# be g(ided b in#ernal s#a#es or in#en#ions* As Miller K Cohen 121 elegan#l s#a#e- &(#a#ive #o&$do'n signals origina#ing in P+C ma &ermi# E#he a)#ive main#enan)e o" &a##erns o" a)#ivi# #ha# re&resen# goals and #he means #o a)hieve #hem* 9he &rovide bias signals #hro(gho(# m()h o" #he res# o" #he brain- a""e)#ing vis(al &ro)esses and o#her sensor modali#ies- as 'ell as s s#ems res&onsible "or res&onse e%e)(#ion- memor re#rieval- emo#ional eval(a#ion- e#)* 9he aggrega#e e""e)# o" #hese bias signals is #o g(ide #he "lo' o" ne(ral a)#ivi# along &a#h'a s #ha# es#ablish #he &ro&er ma&&ings be#'een in&(#s- in#ernal s#a#es and o(#&(#s needed #o &er"orm a given #ask!* Com&(#a#ional models o" #his # &e o" s s#em have )rea#ed a P+C mod(le #ha# )onsis#s o" Er(le! (ni#s 'hose a)#iva#ion leads #o #he &rod()#ion o" a res&onse o#her #han #he one mos# s#rongl asso)ia#ed 'i#h a given in&(#* 122 9h(s- #his mod(le is no# res&onsible "or )arr ing o(# in&(#1o(#&(# ma&&ings needed "or &er"orman)e* Ra#her i# in"l(en)es #he a)#ivi# o" o#her (ni#s 'hose res&onsibili# is making #he needed ma&&ings!* 123 9h(s- #here is no need #o &ro&ose #he e%is#en)e o" a hom(n)(l(s (e*g* )en#ral e%e)(#ive) in #he brain #ha# )an &er"orm a 'ide range o" )ogni#ive o&era#ions 'hi)h are ne)essar "or #he #ask a# hand* 9he overall goal o" )ogni#ive ne(ros)ien)e as a dis)i&line is #o de#ermine #he biologi)al basis o" #he mind* As an in#erdis)i&linar dis)i&line #ha# has evolved "rom bo#h ne(ros)ien)e and &s )holog - )ogni#ive ne(ros)ien#is#s )ons(me da#a derived "rom ea)h o" #hese dis)i&lines in #heir a##em&# #o advan)e )ogni#ive #heor as 'ell as de#ermine ho'
117 113

/uster, ,! M! Cor#i)al d nami)s o" memor * /nt. %. Psychophysiol* 3,- 1,,$1.4= 2000* 2anganath, C!, Cohen, M! F!, 6am, C! - 678sposito, M! @n"erior #em&oral- &re"ron#al- and hi&&o)am&al )on#rib(#ions #o vis(al 'orking memor main#enan)e and asso)ia#ive memor re#rieval* %. Neurosci* 24- 3217$322,= 2004* Miller, B! 5! - 678sposito, M* Sear)hing "or M#he #o&M in #o&$do'n )on#rol* Neuron 43-,3,$ ,33=200,* 3ostle, B! 2! 6ela $&eriod a)#ivi# in #he &re"ron#al )or#e%0 one "(n)#ion is sensor ga#ing* %.Cogn. Neurosci. 17- 1.72$1.20= 200,* Miller, 8! 9! - Cohen, ,! 6* An in#egra#ive #heor o" &re"ron#al )or#e% "(n)#ion* Annu. +e". Neurosci* *-1.7$202= 2001* (72eilly, 2! C!, ?oelle, 6! C!, Braver, 5! 0! - Cohen, ,! 6! Pre"ron#al )or#e% and d nami) )a#egoriGa#ion #asks0 re&resen#a#ional organiGa#ion and ne(ro$mod(la#or )on#rol* Cereb! Corte& 12- 24.$2,7= 2002* Cohen, ,! 6!, 6unbar, 9! - McClelland, ,! <! Fn #he )on#rol o" a(#oma#i) &ro)esses0 a &arallel dis#rib(#ed &ro)essing a))o(n# o" #he S#roo& e""e)#* Psychol. +e"* 27- 332$3.= 1220*

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Revised 17 November 2013 #he brain im&lemen#s )ogni#ive "(n)#ion* Advan)es made in (nders#anding #he )ogni#ive and ne(ral basis o" 'orking memor &rovide e%am&les o" #his s nerg * 6!;s&osi#o s(gges#s #ha# "(#(re s#(dies m(s# )on#in(e #o )onsider bo#h )ogni#ive and ne(ral da#a* Resear)h #h(s "ar s(gges#s #ha# 'orking memor )an be vie'ed as nei#her a (ni#ar nor a dedi)a#ed s s#em* A ne#'ork o" brain regions- in)l(ding #he P+C- is )ri#i)al "or #he a)#ive main#enan)e o" in#ernal re&resen#a#ions #ha# are ne)essar "or goal$dire)#ed behavio(r* 9h(s- 'orking memor is no# lo)aliGed #o a single brain region b(# &robabl is an emergen# &ro&er# o" #he "(n)#ional in#era)#ions be#'een #he P+C and #he res# o" #he brain* 9he )on#rib(#ion o" #he la#eral &re"ron#al )or#e% #o 'orking memor is s#ill being e%&lored* /o'ever- )onsiderable eviden)e s(gges#s #ha# #he la#eral &re"ron#al )or#e% is involved in #he e%e)(#ive or general$&(r&ose as&e)#s o" 'orking memor * +or e%am&le- damage #o #his region in h(mans in#er"eres 'i#h 'orking memor regardless o" #he kind o" s#im(l(s in$ "orma#ion involved* (+(s#er- 1232- @dem) K124 +(r#her- brain imaging s#(dies in h(mans have sho'n #ha# a varie# o" di""eren# kinds o" 'orking memor #asks res(l# in #he a)#iva#ion o" #he la#eral &re"ron#al )or#e%*12,- 12.- 127- 123 @n one s#(d - "or e%am&le- s(bAe)#s 'ere reB(ired #o &er"orm a verbal and a vis(al #ask ei#her one a# a #ime or a# #he same #ime* 122 9he res(l#s sho'ed #ha# #he la#eral &re"ron#al )or#e% 'as a)#iva#ed 'hen #he #'o #asks 'ere &er"ormed #oge#her- #h(s #a%ing #he e%e)(#ive "(n)#ions o" 'orking memor - b(# no# 'hen #he #asks 'ere &er"ormed se&ara#el * 9he la#eral &re"ron#al )or#e% is ideall s(i#ed #o &er"orm #hese general &(r&ose 'orking memor "(n)#ions* @# has )onne)#ions 'i#h #he vario(s sensor s s#ems (like #he vis(al and a(di#or s s#ems) and o#her neo)or#i)al s s#ems #ha# &er"orm s&e)ialiGed #em&orar s#orage "(n)#ions (like s&a#ial and verbal s#orage) and is also )onne)#ed 'i#h #he hi&&o)am&(s and o#her )or#i)al areas involved in long$#erm memor (+(s#er- idem- 1232= <oldman$Raki)- 1237- idem)K130-131* @n addi#ion- i# has )onne)#ions 'i#h areas o" #he )or#e% involved in movemen# )on#rol- allo'ing de)isions made b #he e%e)(#ive #o be #(rned in#o vol(n#aril &er"ormed a)#ions (+(s#er- 1232- idem= <oldman$Raki)- 1273- idem)* Re)en# s#(dies have beg(n #o sho' ho' #he la#eral &re"ron#al )or#e% in#era)#s 'i#h some o" #hese areas* /o'ever- #he bes#$(nders#ood are #he in#era)#ions 'i#h #em&orar s#orage b(""ers in #he vis(al )or#e%*

0AMMA2> - C(?C<A0I(?0 C @n normal individ(als #here is no# a )ons)io(s (nders#anding "rom momen# #o momen# o" ho' o(r brains are dealing 'i#h #he )ons#an# "lo' o" in"orma#ion* We onl kno' s()h #hings as a la)k o" )om&rehension and a "ail(re o"

124 12,

3etrides, M! +ron#al lobes and behavio(r* Current 3pinion in Neurobiology 4- 207$11= 1224* 3etrides, M4 Alivsatos, B4 Meyer, 84 - 8vans, AC* +(n)#ional a)#iva#ion o" #he h(man "ron#al )or#e% d(ring #he &er"orman)e o" verbal 'orking memor #asks* Proceedings of the National Academy of #ciences ?#A 20- 373$32= 1223* ,onides, ,4 0mith, 884 9eoppe, 2A4 Awh, 84 Minosbima, 04 - Mintun, M!A* S&a#ial 'orking memor in h(mans as revealed b P;9* Nature 3.3- .23$2,= 1223* 1rasby, 3M4 /irth, C64 /riston, 9,4 Bench, C4 /rackowiak, 20,4 - 6olan, 2,! +(n)#ional ma&&ing o" brain areas im&li)a#ed in a(di#or $verbal memor "(n)#ion* )rain 11.- 1$20= 1223* 0wart., B84 'algren, 84 /uster, ,M4 0impkins, 84 1ee, M4 - Mandelkern, M! Cor#i)al me#aboli) a)#iva#ion in h(mans d(ring a vis(al memor #ask* Cerebral Cortex ,-20,$14= 122,* 678sposito, M4 6etre, ,4 Alsop, 64 0hin, 24 Atlas, 04 - 1rossman, M* 9he ne(ral basis o" #he )en#ral e%e)(#ive s s#em o" 'orking memor * Nature 373-272$31= 122,* 2eep, 2! Rela#ionshi& be#'een &re"ron#al and limbi) )or#e%0 A )om&ara#ive ana#omi)al revie'* )rain, )eha"ior and 2"olution =., ,$30= 1234* Aylings, 'BM4 - van 8den, C1! P(ali#a#ive and B(an#i#a#ive )om&arison o" #he &re"ron#al )or#e% in ra# and in &rima#es- in)l(ding h(mans* Progress in )rain +esearch 3,- 31$.2= 1220*

12.

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Revised 17 November 2013 memor = #hese even#s o))(r in 'ha# 'e )onsider #o be a brain #ha# is 'orking normall * @" o(r brains do no# 'ork as 'ell #oda as #he did sa a ear ago- #hen 'e have #he abili# #o )om&are #he #'o &hases o" o(r li"e* 9hose 'ho have re)eived earl brain damage have #o )o&e 'i#h a disabled brain 'i#ho(# an kno'ledge o" 'ha# #he si#(a#ion 'o(ld be i" i# hadn!# been damaged* Resear)h #h(s "ar s(gges#s #ha# 'orking memor )an be vie'ed as nei#her a (ni#ar nor a dedi)a#ed s s#em* A ne#'ork o" brain regions- in)l(ding #he P+C- is )ri#i)al "or #he a)#ive main#enan)e o" in#ernal re&resen#a#ions #ha# are ne)essar "or goal$dire)#ed behavio(r* 9h(s- 'orking memor is no# lo)aliGed #o a single brain region b(# &robabl is an emergen# &ro&er# o" #he "(n)#ional in#era)#ions be#'een #he P+C and #he res# o" #he brain* @# is im&or#an# #ha# #his ongoing s#(d o" 'orking memor in)l(des 'ork on #he &roblems #ha# a))r(e "rom earl brain inA(r and a(#ism so #ha# 'e )an be##er (nders#and #he di""i)(l#ies o" #hose a""e)#ed in #hese res&e)#s* 9his- o" ne)essi# - needs #here #o be a be##er a))e&#an)e #ha# a(#ism and a(#is#i) s&e)#r(m disorder are in e""e)# bro(gh# abo(# b &ar#iall disabled brains* More im&or#an#l - #ha# #he disablemen# s#ar#s a# an age #ha# &re)edes m()h ne(rologi)al develo&men#* @# is im&or#an# "or ea)h individ(al so a""e)#ed #ha# #he are re)ognised &rimaril as a brain$inA(red &erson and #ha# an &ossible aberran# behavio(r arises "rom #he abnormal develo&men# o" #heir brain* 9his needs #o be given a&&ro&ria#e )onsidera#ion and s#e&s are #aken #o give &ar#i)(lar hel& and s(&&or# #o bo#h #he )hild and #o i#s &aren#s* A(#ism has been a))e&#ed "or a long #ime no' as a ne(rologi)al )ondi#ion and no# a &s )hia#ri) one* /o'ever- #ha# is no# al'a s a))e&#ed and #he )ommon adA(n)# #o #ha#sho(ld i# ha&&en- is #ha# &s )ho#ro&i) dr(gs are im&osed and #hese )an make #he si#(a#ion m()h 'orse b din# o" #he a""e)# #ha# #he 'ill have on a brain and i#s alread abnormal &a#h'a s* An &s )ho#ro&i) medi)a#ion 'ill mos# likel no# have had an )lini)al #rials on &a#ien#s 'i#h earl brain damage and #here"ore #he 'ill be adminis#ered on a s()k$i#$and$ see basis* :(s# as im&or#an# 'ill be #he "a)# #ha# #here )an be li##le "eedba)k "rom #he &a#ien# on #heir e%&erien)es o" #he medi)a#ion and adverse e""e)#s are #hen more easil mis)ons#r(ed or dis)o(n#ed*

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