A Place For Nouns and A Place For Verbs

Nouns and verbs in the brain
A PLACE FOR NOUNS AND A PLACE FOR VERBS?

A CRITICAL REVIE OF NEUROCO!NITIVE DATA ON !RA""ATICAL
CLASS EFFECTS
Davide Cre#a$di% "anue$a Ber$in&eri% Era$do Pau$esu% and C$audio Lu''atti
De#art(ent o) Ps*+ho$o&*% Universit* o) "i$ano,Bi+o++a% Ita$*
RUNNIN! -EAD. Nouns and verbs in the brain
To a##ear in Brain and Lan&ua&e
Corres#ondin& author.
Davide Cre#a$di
Di#arti(ento di Psi+o$o&ia% Universit/ di "i$ano,Bi+o++a
#ia''a de$$0Ateneo Nuovo 1
23124 "i$ano
Ita$*
E,(ai$. davide5+re#a$di16uni(ib5it
Phone 7dire+t8. 9:; 7382,4<<=:=<3
Fa> 7de#art(enta$8. 9:; 7382,4<<=:?34
1
Abstract
It is &enera$$* he$d that noun #ro+essin& is s#e+i)i+a$$* sub,served b* te(#ora$ areas% @hi$e the
neura$ under#innin&s o) verb #ro+essin& are $o+ated in the )ronta$ $obe5 -o@ever% this vie@ is no@
+ha$$en&ed b* a si&ni)i+ant bod* o) eviden+e a++u(u$ated over the *ears5 "oreover% the resu$ts
obtained so )ar on the neura$ i(#$e(entation o) noun and verb #ro+essin& a##ear to be Auite
in+onsistent5 The #resent revie@ brie)$* des+ribes and +riti+a$$* re,+onsiders the
anato(o,+orre$ative% neuroi(a&in&% "E!% T"S and +orti+a$ sti(u$ation studies on nouns and verbs
@ith the ai( o) assessin& the +onsisten+* o) their resu$ts% #arti+u$ar$* @ithin te+hniAue5 The #a#er
a$so addresses the Auestion as to @hether the in+onsisten+* o) the data +ou$d be due to the variet* o)
the tasBs used5 -o@ever% it e(er&ed that neither the di))erent investi&ation te+hniAues used nor the
di))erent +o&nitive tasBs e(#$o*ed )u$$* e>#$ain the variabi$it* o) the data5 In the )ina$ se+tion @e
thus su&&est that the (ain reason )or the e(er&en+e o) in+onsistent data in this )ie$d is that the
+erebra$ +ir+uits under$*in& noun and verb #ro+essin& are not s#atia$$* se&re&ated% at $east )or the
s#atia$ reso$ution +urrent$* used in (ost neuroi(a&in& studies5
Ce*@ords. nouns% verbs% &ra((ati+a$ +$ass% )un+tiona$ inde#enden+e% anato(i+a$
inde#enden+e% )"RI% PET% anato(o,)un+tiona$ +orre$ations% neura$ +ir+uit5
2
1. Introduction
Re&ard$ess o) their +o((uni+ative )un+tion 7e5&5% asBin& vs5 a))ir(in&8% a$$ hu(an $in&uisti+
a+ts have a +ore in @hi+h 7i8 a s#e+i)i+ obDe+t is sin&$ed out 7denoted8 and 7ii8 a nu(ber o)
#ro#erties are attributed to this obDe+t 7#redi+atedE e5&5% Cho(sB*% 1;;F85 This basi+
+o((uni+ationa$ distin+tion bet@een denotation and #redi+ation (a* be +on+eived as $e>i+a$$*
re)$e+ted in the noun,verb di+hoto(*. e$e(ents in the )or(er +ate&or* natura$$* sub,serve
denotation% @hi$e @ords be$on&in& to the $atter +$ass natura$$* #redi+ate the #ro#erties and
re$ationshi#s re$ative to @hat is denoted b* nouns5 !iven that virtua$$* a$$ $an&ua&es% @ith ver* )e@
and +ontroversia$ e>+e#tions 7e5&5% Fo$e*% 1;;=E Lan&a+Ber% 1;=?E Robins% 1;F28% (aBe a basi+
di))erentiation bet@een nouns and verbs% it )o$$o@s that these t@o #arts o) s#ee+h have a #rivi$e&ed
status a(on& &ra((ati+a$ +$asses5
A behavioura$ dia&nosti+ o) the #rivi$e&ed status o) nouns and verbs +an be )ound in the
e>tensive neuro#s*+ho$o&i+a$ $iterature re#ortin& on a#hasi+ #atients @ith dis#ro#ortionate
i(#air(ent o) either o) these t@o &ra((ati+a$ +$asses5 In a se(ina$ #a#er% -o$(es% "arsha$$ and
Ne@+o(be 71;?18 des+ribed a d*s$e>i+ #atient @hose #er)or(an+e on nouns in a readin& tasB @as
si&ni)i+ant$* better than on the verbs in the sa(e tasB5 This re#ort @as )o$$o@ed b* (an* others
7e5&5% "i+e$i% Si$veri% Vi$$a% G Cara(a''a% 1;=<E "+Cart* G arrin&ton% 1;=F8 and toda*
noun,verb disso+iation has been do+u(ented throu&h severa$ di))erent tasBs% i5e5% @ritin& to
di+tation 7Ra## G Cara(a''a% 23328% ora$ #i+ture na(in& 7Hin&eser G Berndt% 1;==8% @ritten
#i+ture na(in& 7Cara(a''a G -i$$is% 1;;18% @ord,to,#i+ture (at+hin& 7-i$$is G Cara(a''a% 1;;F8
and s#ontaneous s#ee+h 7Bastiaanse G IonBers% 1;;=E Cre#a$di% In&i&no$i% Ver&a% Contardi%
Se(en'a% G Lu''atti% in #ress85 The ran&e o) a#hasi+ s*ndro(es asso+iated @ith #redo(inant
i(#air(ent o) either nouns or verbs is ver* @ideE so(e verb,i(#aired #atients su))er )ro( )$uent
a#hasia% others )ro( non,)$uent a#hasia 7see Lu''atti% Ra&&i% Hon+a% Pistarini% Contardi% G Pinna%
:
23328E (oreover% @hi$e so(e +ases o) disso+iation e(er&ed on$* in s#e+i)i+ tasBs 7e5&5% Berndt%
S$oan% "it+hu(% G -eandi&es% 1;;?a% 1;;?b8 or even a))e+ted di))erent &ra((ati+a$ +$asses in
di))erent tasBs 7-i$$is G Cara(a''a% 1;;F8% other #atients sho@ed a ver* +onsistent #attern over
di))erent behavioura$ tasBs 7Berndt et a$5% 1;;?85 This @ide variet* o) #atterns (aBes it #$ausib$e to
assu(e that noun,verb disso+iation +an e(er&e as a +onseAuen+e o) di))erent t*#es o) )un+tiona$
da(a&e in di))erent #atients. indeed% it has been su&&ested that noun, and verb,s#e+i)i+ i(#air(ent
arises at $e>i+a$,#hono$o&i+a$ 7e5&5% Ra## G Cara(a''a% 23328% $e>i+a$,s*nta+ti+ 7e5&5% Cre#a$di%
A&&uDaro% Arduino et a$5% 23348% se(anti+ 7e5&5% Bird% -o@ard and FranB$in% 23338 and s*nta+ti+
$eve$s 7e5&5% Fried(ann% 233385 hatever )un+tiona$ inter#retation o) the #heno(enon is ado#ted 7as
su&&ested above% this (i&ht var* )ro( #atient to #atient8% the e>isten+e o) a doub$e disso+iation in
brain,da(a&ed individua$s uneAuivo+a$$* ar&ues )or se#arate and )un+tiona$$* inde#endent
re#resentation )or nouns and verbs5 The Auestion then arises as to @hether )un+tiona$ inde#enden+e
is a$so anato(i+a$% i5e5% @hether noun and verb #ro+essin& is +arried out b* neura$ +ir+uits invo$vin&
s#atia$$* se&re&ated brain areas5 This issue @i$$ be addressed in the #resent #a#er b* ana$*sin& the
eviden+e #rovided so )ar b* anato(o,+orre$ative% neuroi(a&in&% T"S and +orti+a$ sti(u$ation
studies5
O) +ourse% noun and verb #ro+essin& deve$o#s throu&h a series o) di))erent +o(#utationa$
sta&es 7e5&5% se(anti+ #ro+essin&% $e>i+a$ retrieva$% arti+u$ation or even $e>i+a$ ortho&ra#hi+
identi)i+ation85 As +$ear$* sho@n b* neuro#s*+ho$o&i+a$ i(#air(ents and% (ore +ontroversia$$*% b*
neuroi(a&in& resear+h% these $eve$s o) #ro+essin& are so(e@hat )un+tiona$$* inde#endent 7i5e5% one
or (ore $eve$s +an +ease to )un+tion a)ter brain da(a&e or be s#ared% as the +ase (a* be8 and (i&ht
be based on di))erent neura$ +ir+uits 7e5&5% )ronta$ da(a&es t*#i+a$$* +ause s*nta+ti+ and arti+u$ator*
de)i+its% @hereas $e>i+a$ de)i+its nor(a$$* arise a)ter te(#ora$ da(a&e85 There)ore% the Auestion o)
@hether nouns and verbs are #ro+essed in brain areas that are s#atia$$* se&re&ated (ust be
<
addressed se#arate$* )or ea+h #ro+essin& $eve$ 7ortho&ra#hi+% #hono$o&i+a$% $e>i+a$% s*nta+ti+%
(or#ho$o&i+a$% and se(anti+85 Co&nitive neuro#s*+ho$o&* has sho@n )un+tiona$ inde#enden+e
bet@een nouns and verbs at virtua$$* a$$ +o&nitive $eve$s 7due to the )a+t that #atients have been
sho@n to disso+iate in severa$ tasBs8E ho@ever% be+ause )un+tiona$ inde#enden+e does not a$@a*s
i(#$* anato(i+a$ inde#enden+e% di))erent &ra((ati+a$ +$asses (a* re+ruit di))erent brain areas at
s#e+i)i+ +o&nitive $eve$s 7e5&5% $e>i+a$,se(anti+ #ro+essin&8% but not at others 7e5&5% #hono$o&i+a$
en+odin&85 In overt +ontrast @ith these +onsiderations% noun and verb #ro+essin& have been o)ten
treated as t@o (ono$ithi+ entities in the $iterature% and this (i&ht be the reason @h* a ver*
in+onsistent #i+ture e(er&es )ro( #revious studies 7as @e @i$$ do+u(ent be$o@85 One the (ain ai(
o) this revie@ is to assess @hether a (ore a##ro#riate +onsideration o) the +o&nitive $eve$7s8
addressed in ea+h stud* +ontributes to +$ari)* the resu$ts obtained so )ar% #erha#s sho@in& that
nouns and verbs are s#atia$$* se&re&ated at +ertain $eve$s o) #ro+essin&% but over$a# anato(i+a$$* at
so(e others5
2. Anatomo-correlative studies on patients suffering from focal brain damage
In a revie@ +ondu+ted b* Ca##a and Perani 7233:8% the authors +on+$uded that $e>i+a$ retrieva$
de)i+its #redo(inant$* invo$vin& a+tion na(in& (ain$* arise in #atients @ith )ronta$ 7and% to a $esser
e>tent% #arieta$8 brain $esions% @hi$e noun,s#e+i)i+ $e>i+a$ i(#air(ents e(er&e a)ter te(#ora$
$esions5 -o@ever% as these authors )ree$* a+Bno@$ed&e% this &enera$i'ation a##ears to be )$a@ed b*
severa$ e>+e#tions5
The )ronto,te(#ora$ di+hoto(* h*#othesis 7FTD-8 ori&inated in a stud* b* Da(asio and
Trane$ 71;;:8% @here three brain,da(a&ed individua$s @ith an a$(ost #ure na(in& de)i+it @ere
des+ribed. t@o #atients had se$e+tive di))i+u$ties in retrievin& nouns as o##osed to verbs in a #i+ture
na(in& tasB% @hi$e one #atient @as i(#aired in retrievin& verbs5 The $esions +hara+teri'in& the
F
noun,i(#aired #atients @ere $i(ited to the te(#ora$ $obe and over$a##ed in the anterior (id, and
in)erior te(#ora$ re&ions% Dust behind the te(#ora$ #o$e5 Juite si&ni)i+ant$*% one o) the t@o
noun,i(#aired #atients +ou$d na(e too$s (u+h better than ani(a$s and )ruits 7Auite as @e$$ as
a+tions% in )a+t8% so that his i(#air(ent a##eared to be s#e+i)i+ to natura$ obDe+ts rather than to
nouns #er seE a si(i$ar K even i) $ess +$ear,+ut K #attern a$so e(er&ed in the se+ond noun,i(#aired
#atient5 The $esion +hara+teri'in& the verb,i(#aired #atient @as $o+ated in the #osterior se&(ent o)
the $e)t in)erior )ronta$ &*rus and the anterior se&(ent o) the $e)t #re+entra$ &*rus5
A$thou&h severa$ studies 7e5&5% Danie$e% !iusto$isi% Si$veri% Co$osi(o% G !ainotti% 1;;<8 have
sho@n resu$ts +onsistent @ith those re#orted in Da(asio and Trane$ 71;;:8% a nu(ber o) other
#a#ers have re#orted eviden+e a&ainst the FTD-5 A#hasi+ #atients have been des+ribed as su))erin&
)ro( verb i(#air(ent a)ter $esions $*in& outside the $e)t )ronta$ re&ions 7Si$veri G Di Betta% 1;;?E
Si$veri% Perri% G Ca##a% 233:8% @hereas verb retrieva$ @as s#ared in other individua$s% in s#ite o)
vast $e)t #re)ronta$ $esions5 In #arti+u$ar% the #atient des+ribed b* De Ren'i and Di Pe$$e&rino 71;;F8
had a ver* $ar&e $e)t )ronta$ $esion invo$vin& the )ronta$ o#er+u$u( and the #re(otor +orti+es in
Brod(ann area 4% and #artia$$* e>tendin& into the #re)ronta$ re&ionE neverthe$ess% he @as sti$$ ab$e
to retrieve verbs both in #i+ture na(in& and senten+e +o(#$etion tasBs 7he +onsistent$* na(ed over
=FL o) the verbs +orre+t$*85
"ore re+ent$*% A&&uDaro% Cre#a$di% Pistarini% Tari++o and Lu''atti 723348 )ound a (ore
+o(#$e> anato(i+a$ #attern in a stud* based on a $ar&e sa(#$e o) verb,i(#aired a#hasi+ #atients5 In
their stud*% usin& a +$assi+a$ #i+ture na(in& tasB% verb i(#air(ent e(er&ed in )our #atients as a
+onseAuen+e o) e>tensive $e)t #eris*$vian da(a&e invo$vin& virtua$$* a$$ the $e)t he(is#here
$an&ua&e areas and in other )our #atients su))erin& )ro( a $esion $i(ited to the #osterior #art o) the
$e)t su#erior te(#ora$ &*rus at its Dun+tion @ith the $e)t in)erior #arieta$ &*rusE )ina$$*% it @as a$so
#resent in another )our a#hasi+ #atients a)ter #ure$* sub,+orti+a$ and insu$ar da(a&e5 No #atient
4
@ith verb i(#air(ent @as re#orted @ith a +erebra$ $esion that @as $i(ited to either the )ronta$ or the
#arieta$ $obe5
Another @ide sa(#$e o) verb,i(#aired a#hasi+ #atients has been des+ribed b* Trane$% "an'e$%
As#% and Ce((erer 7233=8% @ho asBed their #atients to na(e short video +$i#s re#resentin& a+tions5
As in A&&uDaro et a$5 723348% di))erent anato(i+a$ #atterns e(er&ed. nine #atients had $esions
over$a##in& in the $e)t )ronta$ o#er+u$u( and the under$*in& @hite (atter% @hi$e t@o other #atients
had $esions over$a##in& in the (edio,te(#ora$ re&ion and the o++i#ito,te(#ora$ Dun+tion o) the $e)t
he(is#here5 Interestin&$*% a(on& the ; #atients @hose $esions (a>i(a$$* over$a##ed in the $e)t
)ronta$ o#er+u$u(% < a$so had non,ne&$i&ib$e da(a&e to the in)erior #arieta$ $obu$e5
A$to&ether% the FTD- see(s to be over$* si(#$isti+% at $east as )ar as verbs are +on+erned5 No
sin&$e brain area a##ears to be s*ste(ati+a$$* asso+iated @ith verb retrieva$ i(#air(ent% as @ou$d
be e>#e+ted i) the neura$ +ir+uitr* res#onsib$e )or verb #ro+essin& @as s#atia$$* +on+entrated in a
s#e+i)i+ +erebra$ re&ionE si(i$ar +on+$usions @ere dra@n in a re+ent revie@ o) anato(o,+orre$ative
studies on &ra((ati+a$ +$ass e))e+ts 7"Mt'i&% DruBs% "asterson% G Vi&$io++o% 233;85 Parti+u$ar$* in
the vie@ o) the (ost re+ent studies on a $ar&e nu(ber o) #atients 7i5e5% A&&uDaro et a$5% 2334 and
Trane$ et a$5% 233=8% @e +an )air$* assu(e that verb retrieva$ is sub,served b* a
)ronto,te(#oro,#arieta$ net@orB% @hi+h is (ost$* ,, but not e>+$usive$* ,, based in the )ronta$
o#er+u$u( and in the in)erior #arieta$ $obu$eE this +ir+uitr* (ost $iBe$* invo$ves a$so a set o) sub,
+orti+a$ stru+tures brid&in& the +o((uni+ation bet@een the )ronta$ and #arieta$ areas5 "oreover% in
order to a++ount )or those @ho @ere s#ared at retrievin& verbs in s#ite o) havin& )ronta$ or #arieta$
$esions% it (ust be he$d that this net@orB has a hi&h de&ree o) interna$ vi+ariousness% i5e5% it +an
easi$* to$erate the $oss o) #art o) its stru+ture% ar&uab$* be+ause its sub,+o(#onents +an taBe over
the ro$e o) other 7da(a&ed8 sub,+o(#onentsE a$ternative$*% one (a* assu(e a $i(ited de&ree o)
sub,s#e+ia$i'ation @ithin this net@orB5
?
ith re&ard to the neura$ +ir+uitr* under$*in& noun #rodu+tion% the +orre$ation bet@een
se$e+tive i(#air(ent )or na(in& obDe+ts and $esions +entred on the te(#ora$ $obe has re+eived
inde#endent su##ort 7!$osser and Dono)rio% 2331E -i$$is% Tu))iash% it*B% G BarBer% 233285
-o@ever% the #i+ture is sti$$ )ar )ro( bein& +$ear 7e5&5% "Mt'i& et a$5% 233;85 In )a+t% in !$osser and
Dono)rio0s stud* 723318 #atients that under@ent $e)t anterior te(#ora$ $obe+to(* 7LATL8 @ere
(ore i(#aired in na(in& obDe+ts than #atients @hose ri&ht anterior te(#ora$ $obe 7RATL8 @as
re(ovedE ho@ever% LATL #atients @ere (ore i(#aired than RATL on na(in& a+tions a$so% even
thou&h the di))eren+e @as s$i&ht$* $ess than in the obDe+t na(in& tasB5 This &ives rise to the
#ossibi$it* that the $e)t anterior te(#ora$ $obe #arti+i#ates in verb #rodu+tion a$so% even thou&h its
ro$e in noun retrieva$ (a* be (ore )unda(enta$5 "oreover% the te(#ora$ $esion in these #atients
see(s to be (ore anterior than in Da(asio and Trane$0s 71;;:8 #atients5 A$$ noun,i(#aired #atients
+onsidered in A&&uDaro et a$50s stud* 723348 su))ered )ro( a te(#ora$ $esionE ho@ever% @hi$e the
da(a&e a))e+ted the +entra$ #art o) the (id and in)erior te(#ora$ &*ri in three #atients% in other t@o
#atients it @as $i(ited to the te(#oro,o++i#ita$ Dun+tion5 It is @orth notin& that no noun,i(#aired
#atient in+$uded in this stud* su))ered )ro( an anterior te(#ora$ $esion ana$o&ous to that des+ribed
b* !$osser and Dono)rio 7233185
A$to&ether% the anato(o,+orre$ative studies on noun retrieva$ +onsistent$* re#ort a ro$e )or
te(#ora$ areas in obDe+t na(in&% but there is +ontrastin& eviden+e as to @hi+h #art o) the te(#ora$
$obe under$ies noun #ro+essin&5
The )un+tiona$ $eve$ at @hi+h the noun,verb disso+iation ori&inated in the di))erent #atients is a
)a+tor that shou$d be taBen into +onsideration @hen inter#retin& the resu$ts o) the
anato(o,+orre$ative $iterature5 In )a+t% noun and verb #ro+essin& (a* be se$e+tive$* da(a&ed at a
#hono$o&i+a$% ortho&ra#hi+% $e>i+a$% se(anti+ or $e>i+a$,s*nta+ti+ $eve$% and thus brain areas that
have been identi)ied as res#onsib$e )or noun and verb #ro+essin& (i&ht di))er a+ross studies si(#$*
=
be+ause the* under$ie di))erent $eve$s o) $in&uisti+ #ro+essin&5 A$$ the anato(o,+orre$ative studies
des+ribed above (ade use o) #i+ture na(in& tasBsE the brain areas des+ribed in these studies are
thus to be +onsidered as housin& noun, or verb,s#e+i)i+ $e>i+a$,se(anti+ #ro+essin&% $e((a
se$e+tion and $e>e(e se$e+tion 7see Leve$t% Roe$o)s% and "e*er% 1;;;85 The issue o) the +o&nitive
$eve$ at @hi+h noun and verb anato(i+a$ +orre$ates are studied @i$$ be addressed (ore s#e+i)i+a$$*
@hi$e +onsiderin& the neuroi(a&in& $iteratureE in these studies% in )a+t% the +o&nitive $eve$ o)
#ro+essin& under investi&ation +an be e>#eri(enta$$* (ani#u$ated throu&h the +hoi+e o) a s#e+i)i+
e>#eri(enta$ and base$ine tasB5
3. Functional neuroimaging studies
This se+tion e>a(ines a$$ the studies that 7i8 investi&ated the neuro)un+tiona$ net@orB
under$*in& noun and verb #ro+essin& in nor(a$ *oun& adu$ts and 7ii8 in+$uded both a noun and a
verb +ondition5
The behavioura$ and anato(o,+$ini+a$ eviden+e dis+ussed above sets the rationa$e )or )un+tiona$
neuroi(a&in& 7PET% )"RI and "E!8 and trans+rania$ (a&neti+ sti(u$ation 7T"S8 investi&ations
o) noun and verb #ro+essin& in nor(a$ subDe+ts. to date there are (ore or $ess )ort* #a#ers re#ortin&
@ith these investi&ations in the #eer revie@ed $iterature5
As (entioned above% no +$ear #i+ture e(er&es )ro( this vast database and there is no +onsensus
on an anato(i+a$ disso+iation that (a* under#in the nounNverb di+hoto(*E a +$ear e>a(#$e o) this
is &iven in Ber$in&eri% Cre#a$di% Roberti% S+ia$)a% Lu''atti% and Pau$esu 7233=85 In Fi&ure 1 the
authors re#orted the noun,re$ated and verb,re$ated a+tivations )ound in #revious )un+tiona$
neuroi(a&in& studies on a brain te(#$ate5 The resu$t @as a #attern o) noun,re$ated and verb,re$ated
a+tivations distributed a+ross +erebra$ re&ions and he(is#heres% @ith no area e>+$usive$* asso+iated
@ith either nouns or verbs5
;
There is a nu(ber o) +auses @hi+h (a* have +ontributed to &enerate su+h in+onsistent resu$tsE
so(e are @e$$ Bno@n )a+tors that need to be taBen into a++ount @hen assessin& i(a&in& data )ro(
di))erent e>#eri(ents.
7a8 the nature o) the #ri(ar* tasB. noun and verb #ro+essin& (a* be e$i+ited throu&h #rinted or
auditor* sti(u$i and this inevitab$* deter(ines the a+tivation o) +ertain re&ions rather than others
due si(#$* to the di))erent in#ut +hanne$s5
7b8 The nature o) the Oe>#eri(enta$P +o(#onent @ithin the #ri(ar* tasB. subDe+ts (a* be
reAuired to (aBe e>#$i+it Dud&e(ents on s#e+i)i+ Auestions K is OsAuareP a noun or a verb? K or (a*
be asBed to &enerate one +orre+t res#onse 7as in #i+ture na(in&8 or severa$ di))erent res#onses 7as
@hen the* are reAuired to thinB o) as (an* verbs K or nouns K as #ossib$e in res#onse to a &iven
+ue85
7+8 The te+hniAue used 7PET vs5 )"RI vs5 "E!8% the sa(#$e si'e% the nu(ber o) s+ans #er
subDe+t 7in PET and )"RI studies8% the statisti+a$ a##roa+h ado#ted in the ana$*ses o) the data% and
the thresho$d used to se#arate si&ni)i+ant )ro( non,si&ni)i+ant vo>e$s% a$$ #$a* a dra(ati+ ro$e in
deter(inin& @hi+h areas are a+tivated5 Ne&$e+tin& these )a+tors (a* $ead to erroneous +on+$usions5
7d8 Fina$$*% the +hoi+e o) the base$ine +ondition +$ear$* #$a*s an i(#ortant ro$e in deter(inin&
see(in&$* di))erent out+o(es )or the sa(e e>#eri(enta$ tasB 7see Friston% 233F% and Ne@(an%
T@ie&% G Car#enter% 2331% )or an in,de#th +onsideration o) this issue
1
85
1
Ne@(an and +o$$ea&ues 723318 e>#$i+it$* tested the ro$e o) di))erent base$ines 7i5e5% rest state%
#assive $istenin& and tasB,re$ated base$ine8 in deter(inin& di))erent )un+tiona$ neuroi(a&in& resu$ts
@ith the sa(e e>#eri(enta$ tasB% the sa(e (ateria$% and the sa(e sa(#$e o) #arti+i#ants5 A +ertain
de&ree o) base$ine,re$ated variabi$it* in the resu$ts is intrinsi+ in the e>#eri(enta$ desi&ns based on
subtra+tive $o&i+ 7#arti+u$ar$* those @hi+h in+$uded a base$ine +ondition8 and +an on$* be
13
A$$ the +onsiderations e>a(ined above +$ear$* (i$itate a&ainst a s*ste(ati+ and #edanti+
dis+ussion o) ea+h and ever* #eaB o) a+tivation re#orted in the i(a&in& studies on noun and verb
#ro+essin&% and a$so su&&est that it (aBes $itt$e sense to +o(#are resu$ts obtained in studies @hi+h
use non,+o(#arab$e e>#eri(enta$ tasBs and re$ative$* re(ote base$ines5 In $ine @ith this ar&u(ent%
7i8 @e @i$$ re#ort the resu$ts obtained in si(#$e e))e+ts 7tasB vs5 base$ine8 in the tab$es% @ithout
dis+ussin& the( in de#th into the te>t bod*E these a+tivation #eaBs are in )a+t the (ost $iBe$* to be
do(inated b* e))e+ts that are e>#eri(ent,s#e+i)i+ 7i5e5% the e))e+t o) the #ri(ar* tasB8, rather than b*
&enera$ &ra((ati+a$ +$ass e))e+ts
2
5
7ii8 @e @i$$ +onsider the resu$ts o) neuroi(a&in& studies on nouns and verbs +arried out usin&
di))erent e>#eri(enta$ tasBs se#arate$*. this shou$d &uarantee that the brain areas bein& +o(#ared
under$ie $eve$s o) +o&nitive #ro+essin& 7e5&5% #hono$o&i+a$% $e>i+a$% se(anti+8 that are +o(#arab$e% i)
not identi+a$5
This #ro+edure shou$d (aBe +$ear @here @e e>#e+t to observe +onsistent and re#$i+ab$e resu$ts
a+ross di))erent studies% na(e$*% 7i8 in dire+t +o(#arisons bet@een noun and verb s+ans a+ross
studies usin& the sa(e 7or ver* si(i$ar8 e>#eri(enta$ tasBs% and 7ii8 in the intera+tion e))e+ts K
e$i(inated 7or% at $east% +onsiderab$* redu+ed8 b* usin& (ethodo$o&ies su+h as #ara(etri+ or
)a+toria$ desi&ns 7e5&5% Friston 233F85 -o@ever% in the noun,verb $iterature% a$$ )"RI and PET
studies ado#ted a subtra+tive $o&i+ and on$* : out o) :1 did not in+$ude a base$ine +ondition5
2
Be+ause si(#$e e))e+ts o) nouns,(inus,base$ine and verbs,(inus,base$ine +an be obtained a$so in
e>#eri(ents that investi&ate on$* one o) the t@o &ra((ati+a$ +$asses% the tab$es a$so des+ribe those
studies that on$* in+$uded either a noun or a verb +ondition5 e a$so i(#osed as a )urther in+$usion
+riterion that the si(#$e e))e+t ana$*sis @as (ade a&ainst a base$ine that +$ear$* did not reAuire
either noun or verb #ro+essin&5
11
(ain$* bet@een tasB and &ra((ati+a$ +$ass K e(er&in& in studies based on a )a+toria$ desi&n5
A$thou&h it +annot be e>+$uded that so(e s#urious +ross,e>#eri(enta$ variabi$it* @i$$ a))e+t the
resu$ts obtained @ith this #ro+edure% it sti$$ see(s to +onstitute the best too$ to address the issue o)
the anato(i+a$ inde#enden+e o) noun and verb #ro+essin&% as it a$$o@s better +ontro$ over the
+on)ounds due to the t*#e o) e>#eri(enta$ tasB% the base$ine and the statisti+a$ a##roa+h ado#ted5
So% i) a $in&uisti+ $eve$ o) ana$*sis 7e5&5% #hono$o&i+a$% ortho&ra#hi+% $e>i+a$8 e>ists @here the neura$
+ir+uits res#onsib$e )or noun and verb #ro+essin& are s#atia$$* se&re&ated in the brain% @e +an
e>#e+t to )ind +onsistent eviden+e o) this se&re&ation in the resu$ts o) noun,verb dire+t +o(#arison
ana$*ses and in the e(er&en+e o) &ra((ati+a$ +$ass,b*,tasB intera+tions5
3.1 Written lexical decision
e @i$$ +onsider )irst the studies addressin& the (ost #eri#hera$ as#e+ts o) $e>i+a$ #ro+essin&%
i5e5% those re$ated to @ord re+o&nition as investi&ated throu&h $e>i+a$ de+ision5 The studies des+ribed
in this #ara&ra#h ta# on #eri#hera$ ortho&ra#hi+ #ro+essin& 7$etter identi)i+ation and
(or#ho,ortho&ra#hi+ se&(entation8 and $e>i+a$ identi)i+ation5 Lin&uisti+a$$* s#eaBin&% there is $itt$e
&round )or anato(i+a$ se&re&ation o) nouns and verbs at this $eve$ o) ana$*sis% be+ause the
distin+tion bet@een these +$asses is based on s*nta+ti+ and se(anti+s K rather than ortho&ra#hi+ K
as#e+ts5 -o@ever% (u+h #s*+ho$in&uisti+ eviden+e indi+ates that $e>i+a$ de+ision auto(ati+a$$*
re+ruits +on+e#tua$ 7e5&5% Perea G Rosa% 23328 and #hono$o&i+a$ #ro+essin& 7Rast$e G Br*sbaert%
23348% thus risin& e>#e+tations o) di))erent brain a+tivations )or nouns and verbs5
Perani% Ca##a% S+hnur et a$5 71;;;8 +ondu+ted a PET stud* @here a @ritten $e>i+a$ de+ision tasB
@as +o(#ared to an ortho&ra#hi+ dete+tion tasB% in @hi+h the subDe+ts had to de+ide @hether a
strin& +ontained an OQP5 The dire+t +o(#arisons bet@een noun and verb s+ans revea$ed that nouns
did not a+tivate an* brain area (ore than verbs did% @hi$e verbs a+tivated a +o(#$e> #attern o) areas
12
s#annin& a$$ )our $obes in the $e)t he(is#here 7see Tab$e 185
Juite di))erent resu$ts @ere obtained b* T*$er% Russe$$% Fadi$i and "oss 723318 usin& a ver*
si(i$ar (ethodo$o&* 7sa(e e>#eri(enta$ and +ontro$ tasBs and sa(e neuroi(a&in& te+hniAue% i5e5%
PET85 A$thou&h Perani et a$5 71;;;8 and T*$er et a$5 723318 both re#ort that nouns did not a+tivate
an* brain area to a &reater e>tent than verbs 7as revea$ed b* the dire+t +o(#arison bet@een the t@o
&ra((ati+a$ +$asses8% on$* the ri&ht substantia ni&ra 7and no sin&$e +orti+a$ area8 e(er&ed in the
o##osite dire+t +ontrast in T*$er et a$50s stud* 7Tab$e 185
Le>i+a$ de+ision @as a$so +o(#ared @ith an ortho&ra#hi+ base$ine in the e>#eri(ents +arried
out in Ia#anese CataBana s+ri#t b* FuDi(aBi% "i*au+hi% Put' et a$5 71;;;8 and RoBo*a(a%
"i*a(oto% Riera et a$5 7233485 In their ver* arti+u$ated e>#eri(ent 7see Tab$e 1b )or (ore detai$s
about the severa$ areas @hi+h e(er&ed in the si(#$e e))e+t ana$*sis8% FuDi(aBi and +o$$ea&ues )ound
no eviden+e o) di))erentia$ resu$ts )or nouns and verbs% @ith the e>+e#tion o) a hi&her a+tivation )or
nouns bi$atera$$* in the in)erior #arieta$ su$+us and in the #re+entra$ su$+us 7Tab$e 1b85 A++ordin& to
the dire+t +o(#arison ana$*ses +arried out b* RoBo*a(a et a$5 723348% no brain area @as
s#e+i)i+a$$* asso+iated @ith either nouns or verbs% a$thou&h in a re&ion,o),interest ana$*sis the $e)t
(id te(#ora$ &*rus turned out to be (ore a+tivated b* both a+tive and #assive verbs than b* nouns5
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
TABLE 1 ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
In t@o other studies% $e>i+a$ de+ision @as +o(#ared @ith #assive vie@in& o) a )i>ation #oint5 In
a stud* +arried out in Chinese% Li% Iin and Tan 7233<8 )ound noun,s#e+i)i+ a+tivation in the +audate
nu+$eus @hi$e ana$*sin& the dire+t +o(#arison bet@een nouns and verbsE on the +ontrar*% no area
e(er&ed as bein& s#e+i)i+a$$* a+tivated b* verb #ro+essin& 7Tab$e 185 Tho(#son% BonaBdar#our%
1:
Fi> et a$5 7233?8 )o+used their attention on the neura$ +orre$ates o) verb ar&u(ent stru+ture
#ro+essin& and )ound no area to be s#e+i)i+a$$* asso+iated @ith either nouns or verbs in their dire+t
+o(#arison ana$*ses 7Tab$e 185
3.1.1 Summary of the lexical decision literature on written stimuli
A$to&ether% these resu$ts a##ear to be $ar&e$* in+onsistent5 Verbs,(inus,nouns dire+t
+o(#arisons var* striBin&$* a+ross studies. so(e authors re#orted no a+tivation at a$$ 7RoBo*a(a et
a$5% 23348% others sho@ed on$* sub,+orti+a$ a+tivation 7T*$er et a$5% 23318 and so(e others )ound a
+o(#$e> +orti+a$ +ir+uit% (ain$* )ronto,#arieta$ 7Perani et a$5% 1;;;85 The on$* resu$t that a##ears to
be +onsistent is the $a+B o) noun,s#e+i)i+ areas e(er&in& )ro( dire+t +o(#arisons 7but see FuDi(aBi
et a$5% 1;;;8% #ossib$* re)$e+tin& the )a+t that the neura$ +ir+uit under$*in& $e>i+a$ a++ess to noun
re#resentations is the sa(e as that de#uted to verb re+o&nition 7see arburton% ise% Pri+e et a$5%
1;;485
3.2 Semantic judgement task
Se(anti+ Dud&e(ent tasBs on #rinted @ords reAuire the sa(e #eri#hera$ ste#s as $e>i+a$
de+ision tasBs 7i5e5% ortho&ra#hi+N#hono$o&i+a$ #ro+essin& o) $ettersN#hone(es and $e>i+a$
identi)i+ation8% but +$ear$* reAuire #arti+i#ants to #ro+ess the (eanin& o) the sti(u$i (ore
thorou&h$*% renderin& the e(er&en+e o) &ra((ati+a$ +$ass e))e+ts (ore $iBe$*5 -o@ever% the $e>i+a$
de+ision e>#eri(ents i$$ustrated in the #revious #ara&ra#h indi+ate that even i) the e>#eri(enta$ tasB
e$i+its se(anti+ #ro+essin&% nouns and verbs do not ne+essari$* &enerate +onsistent a+tivations
a+ross studies5
Lon&e% Randa$$% Sta(ataBis and T*$er 7233?8 +ondu+ted a )a+toria$,desi&n )"RI e>#eri(ent
)o+used on +ontrastin& (or#ho$o&i+a$$* si(#$e and +o(#$e> sti(u$i in a se(anti+ Dud&e(ent tasB
1<
7i5e5% reAuirin& the #arti+i#ants to indi+ate @hether the sti(u$us is #$easant or not85 The authors did
not obtain a+tivation in the dire+t noun,verb +o(#arisons @hen bare ste(s @ere +onsideredE on the
+ontrar*% @hen in)$e+ted sti(u$i @ere ana$*sed% the #osterior #ortion o) the $e)t (id te(#ora$ &*rus
turned out to be asso+iated @ith verbs% @hi$e no area e(er&ed in the nouns,(inus,verbs +ontrast
7see Tab$e 285
In a b$o+B,desi&n )"RI e>#eri(ent% Cab$e% Lease,S#e$$(e*er and ChatterDee 723328 used t@o
#ara$$e$ versions o) a )or+ed,+hoi+e se(anti+ asso+iation tasB 7si(i$ar to the P*ra(ids and Pa$(
Tree testE -o@ard and Patterson% 1;;28% one @ith #i+tures o) obDe+tsNa+tions and one @ith #rinted
nounsNverbsE this e>#eri(ent a$so in+$uded a visua$ (at+hin& tasB on s+ra(b$ed #i+tures as a
base$ine +ondition5 The dire+t +o(#arison bet@een the obDe+t and the a+tion +onditions revea$ed
that the $atera$ o++i#ito,te(#ora$ re&ion @as a+tivated (ore b* verbs than b* nouns5 Interestin&$*%
this @as not the +ase in the #rinted @ord +onditions 7a$thou&h a #osterior $atera$ te(#ora$ area o)
a+tivation )or verbs @as )ound @hen a re&ion,o),interest a##roa+h @as usedE see Tab$e 285
In three studies +ondu+ted b* T*$er and +o$$ea&ues 7T*$er et a$5% 2331E T*$er% Sta(ataBis% Di+B%
Bri&ht% F$et+her% and "oss% 233:E T*$er% Bri&ht% F$et+her and Sta(ataBis% 233<8 a se(anti+
Dud&e(ent tasB on three +onse+utive sti(u$i 7Auite +ha$$en&in& )or the @orBin& (e(or*8 @as
+o(#ared @ith a $etter identit* tasB 7a$so on three +onse+utive sti(u$i85 SubDe+ts @ere sho@n three
@ords in seAuen+e 7t@o $o@er+ase +ue @ords and an u##er+ase tar&et @ord8 and had to de+ide
@hether the tar&et @ord @as se(anti+a$$* re$ated to both +ue @ords5 ithin this series o) #a#ers% the
+o(#arison bet@een the 2331 PET and the 233< )"RI studies is #arti+u$ar$* interestin&% as the* are
si(i$ar in ever*thin& e>+e#t the sti(u$i used% @ith bare ste(s bein& ado#ted in one stud* and
in)$e+ted @ords in the other5 As )ar as the dire+t +o(#arison bet@een the t@o &ra((ati+a$ +$asses is
+on+erned% neither stud* )ound an* area that @as (ore a+tive )or nouns than verbs% but resu$ts are
in+onsistent @ith res#e+t to the areas that are (ore a+tive )or verbs than @ith nouns. the ri&ht
1F
in)erior te(#ora$ &*rus e(er&ed in the 2331 PET stud*% @hi$e the LIF! turned out to be si&ni)i+ant
in the 233< )"RI e>#eri(ent5 In the 233: stud*% the authors )o+used on the +o(#arison bet@een
ani(a$ nouns and too$ nouns% and bet@een bio$o&i+a$ verbs and too$,re$ated verbsE ho@ever% the
resu$ts do not di))er ver* (u+h a$on& these di(ensions5 As re)$e+ted b* the )a+t that the noun and
verb a+tivation #atterns $ar&e$* over$a# in the si(#$e e))e+t ana$*ses 7see Tab$e 2b )or (ore detai$s8%
ver* )e@ #eaBs o) a+tivation e(er&ed as &ra((ati+a$,+$ass,s#e+i)i+ in the noun,verb dire+t
+o(#arisons 7see Tab$e 2a )or detai$s85
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
TABLE 2 ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
A )urther stud* usin& $etter identit* de+ision as the base$ine tasB @as +ondu+ted b* Davis%
"eunier and "ars$en,i$son 7233<85 Usin& a one,ba+B s*non*(,(onitorin& tasB in @hi+h
#arti+i#ants @ere asBed to de+ide @hether ea+h sti(u$us @as a s*non*( o) the i((ediate$*
#re+edin& ite(% the authors )ound a+tivation )or verbs in dire+t +o(#arison @ith nouns and
adDe+tives in the $e)t #osterior (idd$e te(#ora$ &*rus5 No s#e+i)i+ area o) a+tivation e(er&ed )or
nouns 7see Tab$e 285
T*$er% Randa$$ and Sta(ataBis 7233=8 used a #$easantness Dud&e(ent tasB @ith visua$
#resentation o) nouns and verbs% either in iso$ation or e(bedded in short #hrase 7e5&5% a s(e$$% I
+ou&h85 Durin& the base$ine +ondition% #arti+i#ants @ere #resented @ith the sa(e sti(u$i as in the
e>#eri(enta$ +ondition% but @ere asBed to re)rain )ro( res#ondin&5 The* @ere instru+ted to $ooB at
the( @ithout &ivin& an* Bind o) res#onse 7#assive vie@in&85 The resu$ts sho@ed an in+reased verb,
re$ated a+tivation in the $e)t #osterior (idd$e te(#ora$ &*rus% @hi$e no brain re&ion resu$ted (ore
a+tive )or noun #ro+essin& as o##osed to verb,#ro+essin& 7Tab$e 285
14
ith identit* de+ision on non@ords as their base$ine% Bedn* and Tho(#son,S+hi$$ 723348
investi&ated the noun and verb #ro+essin& neura$ substrate in an i(a&eabi$it*,(at+hed desi&n5 In
this stud*% #arti+i#ants @ere #resented @ith a tar&et @ord )o$$o@ed b* a #air o) @ords5 The* @ere
reAuired to indi+ate @hi+h o) the #air @as (ost si(i$ar in (eanin& to the tar&et @ord5 Usin& an
event,re$ated )"RI stud* and a re&ion,o),interest a##roa+h% the authors dis+overed noun,re$ated
a+tivation in the LIF! and the $e)t (idd$e )ronta$ &*rus% @hi$e the $e)t su#erior te(#ora$ &*rus @as
asso+iated @ith verb #ro+essin& 7see Tab$e 285 The sa(e data% ana$*sed usin& a @ho$e,brain vo$u(e
a##roa+h% sho@ed &reater verb,re$ated a+tivation in the $e)t #osterior +in&u$ate &*rus and the $e)t
#osterior,su#erior te(#ora$ &*rus 7thou&h usin& a $o@er thresho$d8% @hi$e nouns a+tivated the $e)t
in)erior te(#ora$ &*rus 7a&ain at a $o@er thresho$d8 (ore than verbs5
Pa$ti% Sha+har% -end$er and -adar 7233?8 a$so (ade use o) a se(anti+ tasB in their noun,verb
)"RI e>#eri(entE ho@ever% as this stud* @as based on a (ore +o(#$e> )a+toria$ desi&n that a$so
in+$uded a (or#ho$o&i+a$ tasB% its resu$ts @i$$ be re#orted and +o((ented in a se#arate #ara&ra#h5
3.2.1 Summary of the semantic judgement literature
The resu$ts are not ver* +$ear as )ar as verb #ro+essin& is +on+erned. @hi$e so(e authors
re#orted a+tivation in the LIF! and the $e)t insu$a% )air$* stron& eviden+e su&&ests verb #ro+essin&
is a$so based in the #osterior #art o) the te(#ora$ $obe5 Interestin&$*% none o) the studies re#ort
eviden+e )or verb,s#e+i)i+ a+tivation in both re&ions5 As e(er&ed in the $e>i+a$ de+ision
e>#eri(ents% resu$ts are so(e@hat +$earer @hen noun #ro+essin& is +onsidered5 The (aDorit* o) the
studies did not re#ort a+tivation in noun,(inus,verbs dire+t +o(#arisonsE the on$* e>+e#tion is
Bedn* and Tho(#son,S+hi$$ 723348% @ho des+ribed noun,re$ated a+tivation in the LIF! and in the
$e)t in)erior te(#ora$ &*rus5
1?
3.3 Picture naming tasks
"ost )un+tiona$ disso+iations bet@een nouns and verbs in brain,da(a&ed #atients e(er&ed in
#i+ture na(in& e>#eri(entsE there)ore% this tasB is ver* $iBe$* to invo$ve +o&nitive $eve$s @here
nouns and verbs are )un+tiona$$* se&re&ated% and thus shou$d be idea$ )or e$i+itin& +onsistent
&ra((ati+a$ +$ass e))e+ts in )"RI5 -o@ever% #i+ture na(in& en+o(#asses a visua$ ana$*sis o) the
sti(u$us% obDe+t identi)i+ation% $e>i+a$ se$e+tion% #hono$o&i+a$ se$e+tion and seAuen+in&% and
arti+u$ation 7even i) res#onses are &enerated sub,vo+a$$* in (ost )"RI studies and thus arti+u$ation
is not reAuired85 Su+h a broad )o+us #$a*s a&ainst +onsisten+* in the resu$ts% be+ause di))erent
studies (i&ht hi&h$i&ht di))erent areas res#onsib$e )or di))erent +o&nitive $eve$s due to di))eren+es in
the base$ine or in the sti(u$i +hara+teristi+s5
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
TABLE : ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
In Trane$% "artin% Da(asio% !rabo@sBi and -i+h@a0s PET stud* 7233F8% #arti+i#ants @ere
invited to si$ent$* na(e #i+tures o) too$s and a+tionsE this e>#eri(enta$ +ondition @as +o(#ared to
an orientation Dud&e(ent tasB on un)a(i$iar )a+es5 The dire+t +o(#arison ana$*ses revea$ed that
verbs deter(ined (ore a+tivation than nouns in the $e)t )ronta$ o#er+u$u(% the $e)t su#erior
te(#ora$ su$+us and the $e)t (idd$e te(#ora$ &*rus 7see Tab$e :85 The +hoi+e o) a +o(#$ete$*
non,$in&uisti+ tasB as a base$ine (aBes it ver* di))i+u$t to understand @hi+h +o&nitive #ro+esses
under$ie these brain areasE these resu$ts are thus di))i+u$t to +o(#are @ith other data obtained in
#i+ture na(in& e>#eri(ents5
The sa(e +onsideration a##$ies to Da(asio% !rabo@sBi% Trane$% Ponto% -i+h@a% and Da(asio
7233185 In a #i+ture,na(in& PET e>#eri(ent these authors +o(#ared $e>i+a$ retrieva$ o) verbs and
1=
nouns @ith an orientation Dud&e(ent on un)a(i$iar )a+es5 In the dire+t +o(#arison ana$*ses% the*
)ound bi$atera$ verb,s#e+i)i+ a+tivation in the (idd$e te(#ora$ &*rus 7Tab$e :85 As )or Trane$%
"artin et a$50s stud* 7233F8% the +hoi+e o) a +o(#$ete$* non,$in&uisti+ base$ine (aBes this stud*
di))i+u$t to inter#ret in +o&nitive ter(s. in )a+t% the verb,s#e+i)i+ a+tivation )ound in the (idd$e
te(#ora$ &*rus +ou$d re)$e+t $e>i+a$ retrieva$% as @e$$ as se(anti+ or #hono$o&i+a$ #ro+essin&5
Base$ine tasBs @ere not even in+$uded in the )a+toria$ e>#eri(enta$ desi&n e(#$o*ed b*
Sa++u(an% Ca##a% Bates et a$5 723348% and Siri% Tetta(anti% Ca##a et a$5 7233=85 In the )or(er stud*%
no brain areas #assed the FDR,+orre+ted 53F thresho$d in the verbs,(inus,nouns +o(#arisonE @hen
a (ore $enient thresho$d @as ado#ted% the $e)t in)erior #arieta$ $obu$e% the ri&ht )usi)or( &*rus and
the $e)t +erebe$$u( a##eared to be asso+iated @ith verbs5 Nouns a+tivated the ri&ht +uneus% the
#osterior +in&u$ate +orte> and the ri&ht +audate nu+$eus (ore than verbs 7see Tab$e :85 Siri et a$5
7233=8 e(#$o*ed the sa(e #i+tures to e$i+it either a+tion verbs or a+tion nouns in se#arate
e>#eri(enta$ b$o+BsE the noun, and verb,re$ated #atterns o) a+tivation @ere )ound to be Auite si(i$ar
in this stud* 7see Tab$e :a )or detai$s85
A si(i$ar e>#eri(enta$ #aradi&( @as a$so used b* Li$DestrS(% TarBiainen% Parviainen et a$5
7233=85 In this stud* the authors e(#$o*ed a #i+ture na(in& tasB @ith a+tion and obDe+t i(a&es
@here the a+tion i(a&es in+$uded the use o) an obDe+t5 The #arti+i#ants @ere instru+ted to na(e the
obDe+t re#resented in iso$ation 7ObD +ondition8% or the obDe+t re#resented in the a+tion i(a&e
7A+tObD +ondition8% or the a+tion itse$) 7A+t +ondition85 The dire+t +o(#arison bet@een the ObDA+t
and the A+t +ondition sho@ed &reater a+tivation in the $e)t in)erior #arieta$ $obu$e% in the $e)t
#re+uneus% in the ri&ht (idd$e )ronta$ &*rus% in the ri&ht su#erior #arieta$ $obu$e% and in the ri&ht
in)erior #arieta$ $obu$e5 The reverse +ontrast 7A+t,(inus,ObDA+t8 did not sho@ an* si&ni)i+ant
resu$t5 On the +ontrar*% @hen +o(#arin& the A+t +ondition @ith the ObD +ondition% &reater
verb,re$ated a+tivation @as )ound in a nu(ber o) areas $o+ated bi$atera$$* in the )ronta$% te(#ora$%
1;
and #arieta$ $obes 7see Tab$e :a )or detai$s85
Ber$in&eri and +o$$ea&ues 7233=8 used a #i+ture na(in& tasB in their )a+toria$,desi&n
e>#eri(ent and re#orted bi$atera$ verb,s#e+i)i+ a+tivation in a @ide #attern o) areas in+$udin&
)ronta$% te(#ora$% and #arieta$ re&ions5 None o) the brain areas @ere a+tivated (ore b* nouns than
b* verbs5
The resu$ts o) #i+ture,na(in& neuroi(a&in& e>#eri(ents are even $ess +$ear than those o) the
PET and )"RI studies usin& $e>i+a$ de+ision and se(anti+ tasBs5 Verb #ro+essin& has in )a+t been
re#orted to be asso+iated @ith brain areas as diverse as the $e)t )ronta$ o#er+u$u(% the (idd$e
te(#ora$ &*rus% the )usi)or( &*rus% the intra,#arieta$ su$+us% and the +erebe$$u(5 The a+tivation
#eaBs @hi+h survived in nouns,(inus,verbs dire+t +o(#arisons @ere $o+ated either in the LIF!
7)reAuent$* re#orted to be asso+iated @ith verbs in i(a&in& studies usin& other tasBsE e5&5% Perani et
a$5% 1;;;8 or in the ri&ht +uneus and #osterior +in&u$ate +orte>5
"a&netoen+e)a$o&ra#h* 7"E!8 over+o(es the te(#ora$ reso$ution $i(itations t*#i+a$ o) )"RI
studies% so that the ti(in& o) the neura$ a+tivit* a+ross brain re&ions +an be e>#$ored as the sti(u$i
e$aboration un)o$ds5 This (ethodo$o&* there)ore #er(its a +$earer #er+e#tion o) @hen noun
#ro+essin& diver&es )ro( verb #ro+essin&% a$thou&h @ith a $ess a++urate s#atia$ reso$ution than
)"RI5 So(e re+ent data K obtained @ith Intra,+rania$ e$e+tro#h*sio$o&*% or ICE% a ne@ te+hniAue
that% $iBe "E!% a$$o@s a (i$$ise+ond a++ura+* K have indeed sho@n that the neura$ #ro+essin& o)
di))erent t*#es o) sti(u$i 7or o) di))erent as#e+ts o) sti(u$i8 (i&ht di))er (ore in ter(s o) ti(in&
than in ter(s o) their s#atia$ $o+ation 7e5&5% Sahin% PinBer% Cash% S+ho(er% and -a$&ren% 233;85 It is
thus i(#ortant to inte&rate resu$ts obtained @ith te+hniAues o))erin& &ood s#atia$ reso$utions K )"RI
K @ith those obtained @ith te+hniAues o))erin& &ood te(#ora$ reso$utions K "E!5
SSrSs% Corne$issen% Laine% and Sa$(e$in 7233:8 used "E! to +hara+teri'e the brain a+tivation
23
#attern asso+iated @ith noun and verb $e>i+a$ #ro+essin&5 A #i+ture na(in& tasB e$i+ited si(i$ar
s#atio,te(#ora$ a+tivation #atterns under$*in& noun and verb retrieva$E brain a+tivit* started in the
o++i#ita$ areas bi$atera$$* and s#read to the #osterior te(#oro,#arieta$ +orte>% #ro&ressive$*
invo$vin& the (ore anterior re&ionsE in the )ina$ ste#s% a+tivation @as re#orted in the $e)t )ronta$
+orte>5
In a re+ent "E! stud* Li$DestrS(% -u$tTn% ParBBonen% and Sa$(e$in 7233;8 used the
e>#eri(enta$ #aradi&( and (ateria$s o) their #revious )"RI stud* 7Li$DestrS(% TarBiainen%
Parviainen et a$5% 233=8 to eva$uate the ti(in& and sour+e o) a+tivation asso+iated @ith noun or verb
#ro+essin&5 The resu$ts indi+ate stron&er ear$* noun,re$ated res#onses 7133,233 (s8 in the #osterior
+hanne$sE ho@ever% as Li$DestrS( and +o$$ea&ues e(#hasise% the e))e+t o) &ra((ati+a$ +$ass
disa##eared outside this ti(e @indo@% and no other verbNnoun e))e+t @as )ound e$se@here5
3.3.1 Summary of the picture naming literature
TaBen to&ether% the resu$ts )ro( #i+ture,na(in& i(a&in& studies su&&est that noun and verb
#ro+essin& is sub,served b* the sa(e neura$ net@orB5 -o@ever% an over$a# bet@een the neura$
net@orBs under$*in& noun and verb #ro+essin& in #i+ture na(in& (i&ht de#end% at $east in #art% on
the )a+t that a+tion #i+tures unavoidab$* in+$ude the re#resentation o) obDe+ts and a&ents invo$ved in
the a+tion 7e5&5% the #i+toria$ re#resentation o) the verb Oto +utP reAuires the i(a&e o) an a&ent and
an instru(ent% but intransitive a+tions $iBe Oto runP (ust a$so in+$ude the a&ent #er)or(in& the
a+tion8E to date there is no eviden+e a&ainst the #ossibi$it* that obDe+ts and a&ents are i(#$i+it$*
#ro+essed% thus a+tivatin& the neura$ +ir+uits asso+iated @ith obDe+t na(in&5
3. !orphological tasks
"or#ho$o&i+a$ tasBs have been @ide$* ado#ted in neuroi(a&in& e>#eri(ents on noun and verb
#ro+essin&% #arti+u$ar$* a)ter the +ase o) the a#hasi+ #atient @ho @as ab$e to #rodu+e the #$ura$
21
)or(s o) nouns% but not to in)$e+t verbs to the third #erson sin&u$ar% not@ithstandin& the )a+t that
these (or#ho$o&i+a$ #ro+esses are #hono$o&i+a$$* identi+a$ 7Sha#iro% She$ton% G Cara(a''a% 233385
hen +ontrasted @ith an a##ro#riate base$ine% these tasBs +an iso$ate (or#ho$o&i+a$ #ro+essin&
)ro( other +o&nitive $eve$s o) ana$*sisE )air$* +onsistent resu$ts +an thus be e>#e+ted a+ross studies
i) (or#ho$o&i+a$ #ro+essin& o) nouns and verbs re+ruit di))erent brain areas5
In a PET stud*% Sha#iro% "otta&h*% S+hi$$er et a$5 7233F8 asBed !er(an,s#eaBin& subDe+ts to
in)$e+t nouns or verbs a++ordin& to a s*(bo$i+ +ue #resented i((ediate$* be)ore ea+h tria$ 7U
#ro(#ted the #arti+i#ants to #rodu+e the sin&u$ar )or( in noun tria$s and the )irst,#erson,sin&u$ar
)or( in verb tria$s% @hi$e UUU reAuired the subDe+ts to #rodu+e the #$ura$ )or( in noun tria$s and the
)irst,#erson,#$ura$ )or( in verb tria$s85 Pseudo,@ord #assive vie@in& and readin& +onstituted the
+ontro$ +onditions5 In the dire+t nouns,(inus,verbs +ontrast 7(asBed @ith the base$ine8% nouns
*ie$ded si&ni)i+ant a+tivation in the (idd$e #art o) the su#erior te(#ora$ &*rus% o) the (idd$e
#ortions o) the $e)t )usi)or( &*rus and o) the ri&ht insu$a and +erebe$$u(E on the +ontrar*% verbs
a+tivated #arti+u$ar$* the anterior #ortion o) the $e)t su#erior )ronta$ &*rus% the in+$udin& Bro+a0s
area and the ri&ht +erebe$$u( 7see Tab$e <85
In a $ater event,re$ated )"RI stud* +ondu+ted in En&$ish% Sha#iro% "oo and Cara(a''a 723348
asBed the subDe+ts to #er)or( the sa(e in)$e+tiona$ tasB throu&h +o(#$etin& a #hrase that )or+ed the
#rodu+tion o) a sin&u$arN#$ura$ tar&etE #arti+i#ants @ere #resented @ith a sti(u$us #hrase 7e5&5% one
idea8 )o$$o@ed b* a +ue 7e5&5% (an* ... 8 that the* had to +o(#$ete5 In so(e tria$s subDe+ts had to
+han&e the (or#ho$o&i+a$ )or( o) the sti(u$us 7the* @ere #resented @ith one idea and had to
#rodu+e (an* ideas8% @hi$e in others the* had si(#$* to re#rodu+e the ori&ina$ sti(u$us 7the* @ere
#resented @ith one idea and had to #rodu+e one idea8. As in their #revious stud*% in the dire+t
+ontrast bet@een nouns and verbs the (idd$eNanterior #ortion o) the $e)t )usi)or( &*rus @as (ore
a+tivated b* nouns% irres#e+tive o) the (or#ho$o&i+a$ status o) the tar&et @ord 7+itation vs5 in)$e+ted
22
)or(85 B* +ontrast% the $e)t #re)ronta$ +orte>% the $e)t su#erior #arieta$ $obu$e and the $e)t su#erior
te(#ora$ +orte> @ere (ore a+tivated b* verbs than b* nouns% the $atter area a$so bein& sensitive to
+on+reteness% (or#ho$o&i+a$ re&u$arit* and (or#ho$o&i+a$ status 7+itation vs5 in)$e+ted )or(85
As e(er&ed )ro( the +o(#arison bet@een the t@o e>#eri(ents des+ribed above% resu$ts on$*
#artia$$* over$a# even in ver* si(i$ar tasBs5 The (idd$e #ortion o) the $e)t )usi)or( &*rus
+onsistent$* +orre$ates @ith noun in)$e+tion% but the a+tivation in the su#erior te(#ora$ &*rus% the
ri&ht insu$a and the ri&ht +erebe$$u( @ere not re#$i+ated5 Ana$o&ous$*% @hi$e the $e)t in)erior )ronta$
+orte> is asso+iated @ith verbs in both studies% #arieta$ and te(#ora$ areas e(er&ed as asso+iated
@ith verb in)$e+tion in the se+ond on$*5
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
TABLE < ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
Sahin% PinBer and -a$&ren 723348 asBed #arti+i#ants to +o(#$ete a s(a$$ senten+e 7e5&5%
Resterda* the*555 or Those are the5558 @ith a +orre+t$* in)$e+ted re&u$ar or irre&u$ar verbNnounE a
)i>ation +ondition @as used as a base$ine5 The si(#$e +ontrasts a&ainst the base$ine sho@ed ver*
si(i$ar #atterns o) a+tivation )or verbs and nouns 7see Tab$e <b )or (ore detai$s8E the on$*
di))eren+e @as a stron&er a+tivation o) BA <<N<FN<? and the se$e+tive a+tivation o) the intra#arieta$
su$+us )or nouns% as revea$ed b* the dire+t +o(#arison ana$*ses 7Tab$e <a85 This stud* does not
su##ort the )indin&s o) Sha#iro et a$5 in either 233F or 2334% i5e5% the asso+iation bet@een verb
in)$e+tion and #re)ronta$ a+tivation5 On the +ontrar*% #re(otorN#re)ronta$ #eris*$vian re&ions are
si&ni)i+ant$* (ore a+tivated b* nounsE this is a$so the +ase )or the intra#arieta$ su$+us% +ontrar* to
@hat is su&&ested b* other resu$ts @hi+h hi&h$i&ht a ro$e )or #arieta$ areas in verb #ro+essin& 7see
Ber$in&eri et a$5% 233=E "aran&o$o% Piras% !a$ati% and Burani% 2334E Sha#iro et a$5% 233485
2:
Re+ent$*% Burton% Crebs,Nob$e% !u$$a#a$$i and Berndt 7233;8 investi&ated the neura$ net@orB
under$*in& noun and verb #ro+essin& in a )"RI stud*5 Parti+i#ants @ere instru+ted to e>#ress a
(or#ho$o&i+a$ Dud&e(ent in a si(#$e senten+e +onte>tE in #arti+u$ar the* @ere #resented @ith verbs
and nouns #re+eded b* either OtheP or OtoP and @ere asBed to Dud&e @hether the senten+e @as
&ra((ati+a$$* +orre+t5 The resu$ts sho@ed noun,s#e+i)i+ a+tivation in the $e)t in)erior )ronta$ &*rus%
@hi$e no area @as s#e+i)i+a$$* a+tivated b* verbs5
Fina$$*% Ber$in&eri et a$5 7233=8 and Pa$ti et a$5 7233?8 in+$uded a (or#ho$o&i+a$ tasB in their
)a+toria$,desi&n e>#eri(ents5 In the dire+t +o(#arison ana$*ses% the )or(er authors re#orted
verb,s#e+i)i+ a+tivation in a )ronto,#arieta$ bi$atera$ +ir+uit 7not in+$udin& the LIF! or the $e)t
insu$aE see Tab$e Fa8 and noun,s#e+i)i+ bi$atera$ a+tivation in the LIF!% the $e)t in)erior #arieta$
$obu$e% the $e)t #re+uneus and the su#erior and (idd$e o++i#ita$ &*rus5 Pa$ti and +o$$ea&ues% on the
other hand% re#orted verb,s#e+i)i+ a+tivation in the $e)t #re(otor area @hi+h (a* be in $ine @ith the
resu$ts des+ribed b* Sha#iro and +o$$ea&ues 7233FE 233485
3..1 Summary of the morphological task literature
A$to&ether% as @as the +ase )or the studies usin& $e>i+a$ de+ision% se(anti+ Dud&e(ent and
#i+ture na(in& tasBs% the resu$ts re#orted above are )ar )ro( +onstitutin& a +$ear #i+ture. no sin&$e
brain area e(er&es as +onsistent$* asso+iated @ith either noun or verb #ro+essin& in an* o) the
i(a&in& studies usin& (or#ho$o&i+a$ tasBs5
3." #luency tasks
Fro( a +o&nitive #oint o) vie@% )$uen+* tasBs see( to en+o(#ass the @ho$e @ord &eneration
#ro+ess% )ro( sear+h and se$e+tion in the $e>i+a$,se(anti+ s#a+e to #hono$o&i+a$ en+odin& and
arti+u$ation5 Due to their broad )o+us% these tasBs are ver* sensitive in the +$ini+a$ eva$uation o)
a#hasi+ #atients% but @hen the* are used in neuroi(a&in& studies the* #ose +$ear $i(itations on the
2<
+o&nitive inter#retation o) the resu$tsE un$ess the base$ine has been ver* +are)u$$* desi&ned% it is hard
to Bno@ @hether an a+tivated area re)$e+ts se(anti+% $e>i+a$% #hono$o&i+a$% or arti+u$ator* #ro+esses5
arburton et a$5 71;;48 +ondu+ted a PET stud* in @hi+h subDe+ts @ere asBed to #rodu+e as
(an* nouns or verbs as #ossib$e% re$ated to a s#oBen noun used as a +ue 7E>#eri(ent :8E as a
+ontro$ +ondition% subDe+ts @ere asBed to Bee# their e*es +$osed and to Oe(#t* their (indP5
arburton et a$5Vs stud* sho@ed the sa(e a+tivation #attern )or verb and noun #ro+essin&. this
#attern s#reads over the @ho$e brain @ith the e>+$usion o) the o++i#ita$ $obe 7see Tab$e Fb )or (ore
detai$s85 hen nouns and verbs @ere dire+t$* +o(#ared% verb &eneration #rodu+ed &reater
a+tivation in the in)erior )ronta$ &*rus% the S"A% the te(#ora$ su$+us% the te(#oro,#arieta$ Dun+tion
and the in)erior #arieta$ $obe% @hi$e noun &eneration a+tivated the ri&ht su#erior )ronta$ su$+us and
the $e)t anterior +in&u$ate &*rus5 It is virtua$$* i(#ossib$e to Bno@ @hi+h +o&nitive $eve$s are
re)$e+ted in these a+tivationsE there)ore% these resu$ts +ontribute $itt$e to our understandin& o) the
neura$ i(#$e(entation o) noun and verb #ro+essin&5 -o@ever% the stud* is theoreti+a$$* re$evant
both be+ause it @as the )irst neuroi(a&in& e>#eri(ent (ani#u$atin& &ra((ati+a$ +$ass as an
inde#endent )a+tor and be+ause it @as the )irst to re#ort that verbs,(inus,nouns +ontrasts &enerate
ver* @ide net@orBs o) a+tivation% @hi$e nouns,(inus,verbs +o(#arisons revea$ )e@ brain areas% i)
an*5
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
TABLE F ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
3.$ %ask&'y&grammatical&class factorial studies
As i$$ustrated above% noun and verb a+tivations obtained in studies usin& di))erent tasBs do not
+onver&e on the sa(e brain areas5 This is a$so true )or tasBs ta##in& si(i$ar +o&nitive $eve$s o)
2F
ana$*sisE this (a* indi+ate that brain a+tivations o)ten re)$e+t tasB,s#e+i)i+ e))e+ts rather than
&ra((ati+a$ +$ass di))eren+es5 In this res#e+t% an interestin& a##roa+h has been introdu+ed re+ent$*
re&ardin& the e>#eri(enta$ and base$ine tasBs used @hen investi&atin& noun and verb #ro+essin&5
Pa$ti et a$5 7233?8 and Ber$in&eri et a$5 7233=8 e(#$o*ed )a+toria$ e>#eri(ents in @hi+h t@o di))erent
tasBs @ere used% (aBin& it #ossib$e to address tasB,de#endent as @e$$ as tasB,inde#endent
&ra((ati+a$,+$ass e))e+ts5 These e>#eri(enta$ desi&ns a$$o@ a better assess(ent o) @hether #atterns
o) brain a+tivation are asso+iated to &enuine &ra((ati+a$ +$ass e))e+ts rather than to tasB de(and%
#arti+u$ar$* i) the tasBs e(#$o*ed share on$* one +o&nitive #ro+ess. brain areas that +onsistent$*
e(er&e as asso+iated to a &ra((ati+a$ +$ass in both tasBs re)$e+t the #ro+essin& at that +o&nitive
$eve$ and are un$iBe$* to re)$e+t tasB,s#e+i)i+ #ro+essin&% thus &eneratin& resu$ts that shou$d be (ore
eas* to re#$i+ate 7)or an ear$* su&&estion o) the advanta&e o) )a+toria$ desi&ns in )"RI see Friston%
Pri+e% F$et+her% "oon% Fra+Bo@iaB% G Do$an% 1;;485
Pa$ti and +o$$ea&ues used a (or#ho$o&i+a$ and a se(anti+ tasB% in @hi+h #arti+i#ants @ere
asBed to Dud&e @hether an auditori$* #resented nounNverb @as in)$e+ted in the #$ura$ )or( or @as
re$ated to a &iven se(anti+ +ate&or*5 The authors )ound &reater a+tivation )or verbs than )or nouns
a+ross tasBs in the LIF!% in the #re(otor area and in the $e)t su#erior te(#ora$ su$+us% @hi$e none o)
the brain areas @ere a+tivated (ore b* nouns than b* verbs5 In a si(#$e e))e+t ana$*sis% the authors
a$so )ound that the VWN e))e+t e(er&ed in the LIF! in the se(anti+ tasB on$*% in the #re(otor area
in the (or#ho$o&i+a$ tasB on$*% and in the $e)t su#erior te(#ora$ su$+us in both tasBs5 These $atter
resu$ts are ver* re$evant as the* sho@ ho@ the e>#eri(enta$ tasB deter(ines the e(er&en+e o)
di))erent &ra((ati+a$,+$ass,s#e+i)i+ areas even @hen identi+a$ sti(u$i are e(#$o*ed5
Ber$in&eri et a$5 7233=8 (ade an e>#$i+it use o) the #otentia$ o) )a+toria$ desi&ns b* usin& t@o
e>#eri(enta$ tasBs that share on$* one s#e+i)i+ +o&nitive #ro+ess 7na(e$*% $e>i+a$ #ro+essin&85 To
this ai(% the* asBed their #arti+i#ants to +arr* out a #i+ture na(in& and a &ra((ati+a$ +$ass
24
s@it+hin& tasBE in this $atter% #arti+i#ants @ere invited to #rodu+e the nounNverb +orres#ondin& to a
verbNnoun +ue 7e5&5% a##$auso% a##$ause% )ro( a##$audire% to a##$aud% or vi+e versa85 This tasB
reAuires $e>i+a$ #ro+essin& in Ita$ian as the (or#he(e used to #rodu+e a deverba$ noun is
idios*n+rati+ and +annot be derived b* a##$*in& a (or#ho$o&i+a$ ru$e% i5e5% there is no +ue in the
verb that sa)e$* indi+ates @hi+h deverba$ (or#he(e (ust be atta+hed to &enerate a noun 7e5&5%
+a$+o$ & are % to +a$+u$ate% &enerates +a$+o$ & o % +a$+u$ation% but so$$eva & are % to $i)t% &enerates
so$$eva & a(ento % $i)tin&
N
85 In this e>#eri(ent% verbs @ere +onsistent$* asso+iated @ith bi$atera$
a+tivation in the #re+entra$ and #ost+entra$ &*ri% the #ara+entra$ $obu$e% the su#erior #arieta$ $obu$e
and the #re+uneus% and @ith a+tivation in the S"A and the in)erior #arieta$ $obu$e in the ri&ht
he(is#here5 No brain area sho@ed &reater a+tivation )or nouns than )or verbs +onsistent$* a+ross
tasBs5 ThanBs to the )a+toria$ desi&n e(#$o*ed% Ber$in&eri et a$5 7233=8 @ere a$so ab$e to identi)*
brain areas @here &ra((ati+a$ +$ass intera+ted @ith the e>#eri(enta$ tasBE in #arti+u$ar% the $e)t
in)erior )ronta$ &*rus @as )ound to be #redo(inant$* a+tivated b* verbs in #i+ture na(in&% but b*
nouns in the &ra((ati+a$,+$ass s@it+hin& tasB des+ribed above5 On the basis o) this )indin&% the
authors su&&ested that the $eve$ o) a+tivation in the LIF! (a* re)$e+t the overa$$ +o&nitive de(and
i(#osed b* a &iven tasB% rather than a )ir( asso+iation @ith a &iven &ra((ati+a$ +$ass 7see a$so
Bedn*% -u$bert% G Tho(#son,S+hi$$% 2334E Fie' G Petersen% 1;;=E Sabsevit'% "ed$er% Seidenber&%
G Binder% 233F8
4. Noun and verb processing in the brain as assessed through T!
On$* T"S studies investi&atin& the neura$ basis o) nouns and verbs #ro+essin& in *oun&
hea$th* adu$ts are in+$uded in this se+tion5
Studies usin& re#etitive trans+rania$ (a&neti+ sti(u$ation 7rT"S8 a$so revea$ed a re$evant
i(#a+t o) the tasB used to investi&ate noun and verb #ro+essin& in the brain5 Sha#iro%
2?
Pas+ua$,Leone% "otta&h*% !an&itano and Cara(a''a 723318 )ound that #re)ronta$ sti(u$ation 7:33
(s% 1 -'8 +aused in+reased res#onse $aten+ies )or verbs but not )or nounsE on the other hand% Ca##a%
Sandrini% Rossini% Sosta and "iniussi 723328 obtained the o##osite e))e+t 7verb,s#e+i)i+ )a+i$itation
re$ative to base$ine8 in a #i+ture na(in& tasB a)ter sti(u$atin& the sa(e re&ion 7F33 (s% 23 -'85
These $atter )indin&s are #arti+u$ar$* striBin& +onsiderin& that a sti(u$ation )reAuen+* o) 23 -' is
+o((on$* he$d to +ause inhibition in the +erebra$ +orte>. i) the dorso$atera$ #re)ronta$ +orte> is
dire+t$* res#onsib$e )or verb #ro+essin& K as su&&ested b* Sha#iro et a$5 723318 K @e @ou$d have
e>#e+ted to observe se$e+tive inhibition o) verb #rodu+tion in Ca##a et a$5Vs stud* 723328% @hi+h
@as not the +ase5 The diver&in& resu$ts re#orted b* Sha#iro et a$5 723318 and b* Ca##a et a$5 723328
+ou$d be e>#$ained b* the )a+t that in the )or(er stud* subDe+ts @ere asBed to +arr* out a
(or#ho$o&i+a$ tasB% @hi$e in the $atter the* had to na(e #i+turesE ho@ever% this @ou$d indi+ate that
)ronta$ re&ions are +ru+ia$ )or in)$e+tin& verbs% but not )or retrievin& the(% in +on)$i+t @ith so(e
)"RI and PET eviden+e 7e5&5% Sa++u(an et a$5% 2334E Sahin et a$5% 233485 O$iveri% Fino++hiaro%
Sha#iro% !an&itano% Cara(a''a% and Pas+ua$ Leone 7233<8 +ondu+ted a )urther rT"S stud*% in
@hi+h the $e)t #ri(ar* (otor +orte> @as )ound to #$a* a +riti+a$ ro$e in the retrieva$ o) a+tion @ords
irres#e+tive o) their &ra((ati+a$ +$ass 7i5e5 both nouns and verbs8E on+e a&ain the tasB @as to in)$e+t
a visua$$* #resented @ord )o$$o@in& a +ue in the )or( o) a s*(bo$% indi+atin& @hether the verbNnoun
had to be #rodu+ed in the sin&u$ar or in the #$ura$ )or(5
On the basis o) these resu$ts% Lo !er)o% O$ivieri% Torriero% Sa$erno% Co+h% and Ca$ta&irone
7233=8 e>#$ored the ro$e o) the #re)ronta$ +orte> and o) the #ri(ar* (otor +orte> in both verb and
a+tion #ro+essin&5 The* too used a (or#ho$o&i+a$ tasB% si(i$ar to that used b* Sha#iro and
+o$$ea&ues 723318 and b* O$ivieri et a$5 7233<85 Resu$ts sho@ed that @hen rT"S @as a##$ied to the
$e)t #re)ronta$ +orte>% there @as a se$e+tive de$a* in a+tion verb retrieva$5 On the +ontrar*% an rT"S
i(#u$se a##$ied to the $e)t (otor +orte> +aused a si&ni)i+ant de$a* in #ro+essin& a+tion @ords%
2=
irres#e+tive o) &ra((ati+a$ +$ass5 A$thou&h these resu$ts see( to better +$ari)* the ro$e o) the )ronta$
area in verbNa+tion #ro+essin&% the +hoi+e o) a #ure (or#ho$o&i+a$ tasB rather than another t*#e o)
tasB does not in itse$) +ontribute to de+idin& @hether the ro$e o) the #re)ronta$ +orte> is s#e+i)i+ )or
verb in)$e+tion or (ore &enera$$* asso+iated to verb $e>i+a$,se(anti+ #ro+essin&5
In order to address this issue% Ca##e$$etti% Fre&ni% Sha#iro% Pas+ua$,Leone and Cara(a''a
7233=8 re+ent$* e>#$ored the ro$e #$a*ed b* distin+t #arts o) the #re)ronta$ +orte> in noun and verb
#rodu+tion @ithin a #hrasa$ +onte>t5 The* obtained a si&ni)i+ant in+rease in res#onse $aten+ies )or
verb #rodu+tion @hen sti(u$atin& the $e)t anterior (id)ronta$ &*rus5 No si&ni)i+ant verb,s#e+i)i+
inter)eren+e e))e+t @as )ound a)ter the sti(u$ation o) the #osterior (idd$e )ronta$ &*rus and the
Bro+aVs area and% in addition% none o) these re&ions @as #re)erentia$$* invo$ved in the #rodu+tion o)
either re&u$ar or irre&u$ar in)$e+tion5
". #ortical stimulation in pre-surgical patients
Corina% !ibson% "artin% Po$iaBov% BrinB$e*% and ODe(ann 7233F8 investi&ated the neura$
re#resentation o) verbs and nouns in 1: neurosur&i+a$ subDe+ts under&oin& +orti+a$ (a##in& @hi$e
a@aBe5 In this stud* nouns *ie$ded a Auite +oherent )ra(e. inabi$it* to na(e obDe+t #i+tures @as
+onsistent$* +aused b* +orti+a$ sti(u$ation o) the $e)t anterior su#erior and o) the $e)t (idd$e
te(#ora$ &*rus5 On the +ontrar*% the areas +ausin& inabi$it* to na(e verbs varied re(arBab$* a+ross
#arti+i#ants% in+$udin& the $e)t (idd$e su#erior te(#ora$ &*rus% the $e)t su#ra(ar&ina$ &*rus and the
$e)t #osterior (idd$e te(#ora$ &*rus5 Un)ortunate$*% due to te+hni+a$N+$ini+a$ +onstraints it @as not
#ossib$e to sti(u$ate an* )ronta$ areasE ho@ever% the stud* +$ear$* hi&h$i&hts an i(#ortant ro$e o) the
te(#ora$ re&ions in retrievin& verbs5 ODe(ann% ODe(ann and Letti+h 723328 had +ondu+ted a si(i$ar
e>#eri(ent so(e *ears be)ore% but @ith t@o i(#ortant di))eren+es. )ronta$ re&ions @ere a$so
sti(u$ated and a verb &eneration tasB instead o) a #i+ture na(in& tasB @as used to e$i+it verbs5 This
2;
stud* +on)ir(ed that verb,s#e+i)i+ areas (a* var* notab$* a+ross subDe+ts. in : out o) the 1:
#arti+i#ants tested% both )ronta$ and te(#ora$ sti(u$ation +aused verb,s#e+i)i+ de)i+itsE in 2
#arti+i#ants% a verb de)i+it a##eared a)ter )ronta$ sti(u$ation% @hereas it e(er&ed a)ter te(#ora$
sti(u$ation in another = #arti+i#ants5 A$to&ether% te(#ora$ verb,re$ated areas see(ed to be (ore
s#e+i)i+ than )ronta$ verb,re$ated areas. in )a+t% a(on& the 1: )ronta$ sites in @hi+h sti(u$ation
ha(#ered verb &eneration% ? a$so +aused an obDe+t,na(in& de)i+it5 On the +ontrar*% ho@ever% 12 out
o) 23 te(#ora$ verb,s#e+i)i+ sites @ere $o+ated at $east 1 +( a@a* )ro( an* obDe+t,na(in& re$ated
site 7(ost$* $o+ated in the #osterior #art o) the (edia$ and u##er te(#ora$ &*rus and in the
su#ra(ar&ina$ &*rus85 A$to&ether% dire+t +orti+a$ sti(u$ation studies e(#hasi'e the ro$e o) the
#osterior te(#ora$ $obe in retrievin& verbs% as @e$$ as &reat a+ross,subDe+t variabi$it* re&ardin& the
e>a+t $o+a$i'ation o) the neura$ sites% @hi+h% @hen sti(u$ated% hindered either noun or verb $e>i+a$
retrieva$5
$. %iscussion
There is +$ear$* a &reat dea$ o) in+onsisten+* in the eviden+e des+ribed in this #a#er5 It (i&ht
have been e>#e+ted that the di))erent e>#eri(enta$ te+hniAues ado#ted 7e5&5% anato(o,+$ini+a$
+orre$ations% )"RI% "E!8 and various behavioura$ tasBs used to investi&ate the anato(i+a$
under#innin&s o) noun and verb #ro+essin& @ou$d a++ount )or the dis+re#an+ies% but +ontrastin&
)indin&s @ere re#orted even in studies usin& the sa(e (ethodo$o&* and% in severa$ +ases% a$so the
sa(e e>#eri(enta$ tasBs 7e5&5% Perani et a$5% 1;;; and T*$er et a$5% 2331E Siri et a$5% 233= and
Sa++u(an et a$5% 2334E Sha#iro et a$5% 2331 and Ca##a et a$5% 233285 The #resen+e o) in+onsistent
resu$ts @ithin tasB and (ethodo$o&* (aBes it even $ess a##ro#riate to +o(#are resu$ts a+ross tasBs
and (ethodo$o&ies5
The dis+re#an+ies do not see( to be due to te+hni+a$ detai$s 7e5&5% statisti+a$ thresho$d% sa(#$e
:3
si'e8 K @hi+h have been sho@n to in)$uen+e the out+o(e o) i(a&in& studies 7e5&5% Friston% 233F8 K
or to noise si&na$s K @hi+h unavoidab$* a))e+t the de#endent variab$e in neuroi(a&in& e>#eri(ents5
I) this @ere the +ase% @e @ou$d e>#e+t so(e rando( noise over a set o) rather +onsistent resu$tsE but
no su+h set o) resu$ts e(er&ed in our revie@5 Indeed% the resu$ts are ver* in+onsistent overa$$5 This
(a* be due si(#$* to (ethodo$o&i+a$ as#e+ts and sour+es o) noise intrinsi+ to the (ethodo$o&*E
ho@ever% this @ou$d basi+a$$* i(#$* that neuroi(a&in& data are so #oor$* re#$i+ab$e to be o) $itt$e
s+ienti)i+ uti$it* overa$$5
The )ronto,te(#ora$ di+hoto(* h*#othesis 7FTD-8 #ut )or@ard in severa$ anato(o,+orre$ative
and neuroi(a&in& studies% and su##orted b* Ca##a and Perani in a #revious revie@ 7233:8% is )ar
)ro( bein& even #artia$$* +on)ir(ed5 A++ordin& to the FTD-% 7$e)t8 )ronta$ areas under$ie verb
#ro+essin&% @hereas the 7$e)t8 te(#ora$ re&ion is the +erebra$ +ounter#art o) noun #ro+essin&5
-o@ever% o) 2= neuroi(a&in& studies% on$* 1< have re#orted a )ronta$ area s#e+i)i+a$$* a+tivated b*
verbs in either si(#$e +ontrasts or dire+t +o(#arisons% @hi$e 1< have )ai$ed to do so and 4 o) these
1< studies have even )ound noun,s#e+i)i+ )ronta$ area a+tivation 7"artin% i&&s% Un&er$eider% G
-a>b*% 1;;4E Chao G "artin% 2333E T*$er et a$5% 233:E Sahin et a$5% 2334E Bedn* G
Tho(#son,S+hi$$% 2334E Siri et a$5% 233=85 Noun,re$ated a+tivation in the )ronta$ areas @as &enera$$*
obtained @ith too$ nouns 7e5&5% "artin et a$5% 1;;4E T*$er et a$5% 233:8% but this +ate&or*,s#e+i)i+it*
@as not +on)ir(ed in other si(i$ar studiesE )or e>a(#$e% T*$er et a$5 7233:8 and Sahin et a$5 723348
)ound )ronta$ a+tivation asso+iated @ith nouns that did not re)er to (ani#u$ab$e obDe+ts5
Further(ore% the FTD- obvious$* #redi+ts the e(er&en+e o) )ronta$ a+tivation asso+iated @ith
verbs and te(#ora$ a+tivation asso+iated @ith nouns in dire+t +o(#arison ana$*sesE ho@ever% o) the
1F i(a&in& studies re#ortin& verb,noun dire+t +ontrasts% on$* F )ound verb,re$ated a+tivation in
)ronta$ areas 7Perani et a$5% 1;;;E Sha#iro et a$5% 233FE Sha#iro et a$5% 2334E T*$er et a$5% 233:E T*$er
et a$5% 233<8 and on$* 2 sho@ed noun,re$ated a+tivation in te(#ora$ areas 7Sha#iro et a$5% 233FE
:1
Sha#iro et a$5% 233485 A @ord o) +aution shou$d be s#ent here on the )a+t that% in this $ine o)
reasonin&% #ositive and ne&ative eviden+e )or the FTD- are &iven the sa(e @ei&ht% @hereas the
+o((on #ro+ess o) statisti+a$ in)eren+e ado#ted in neuroi(a&in& studies 7and in the @ho$e )ie$d o)
#s*+ho$o&*% in )a+t8 is so(e@hat biased to@ards #ositive )indin&s% @hi$e ne&ative resu$ts (a*
re(ain not )u$$* assessed5 Fi&ure 1 i$$ustrates a (ore detai$ed ana$*ses o) ho@ (an* studies
re#orted noun or verb,s#e+i)i+ a+tivations in either si(#$e e))e+ts or dire+t +o(#arisonsE even at a
)iner s#atia$ reso$ution 7i5e5% +onsiderin& se#arate$* di))erent #ortions o) the in)erior )ronta$ &*ri and
o) the te(#ora$ $obes8% no )ronta$ area is s#e+i)i+a$$* asso+iated @ith verbs% nor an* te(#ora$ area is
s#e+i)i+a$$* asso+iated @ith nouns5 It is +$ear that the FTD- does not survive +on)rontation @ith the
eviden+e avai$ab$e to date5
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
FI!URE 1 ABOUT -ERE
,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,,
The #rob$e( see(s to be that no a$ternative +oherent )ra(e has e(er&ed )ro( this revie@5
A$ternative$* to the FTD-% the neuro$in&uisti+ eviden+e dis+ussed at the be&innin& o) the #a#er
su&&ested that verb #ro+essin& @as (ost $iBe$* +arried out b* a @ide neura$ +ir+uit invo$vin& not
on$* )ronta$% but a$so #arieta$ areas 7e5&5% Si$veri et a$5% 233:8 and sub,+orti+a$ stru+tures brid&in& the
anterior,#osterior +o((uni+ation 7e5&5% A&&uDaro et a$5% 2334E see a$so "Mt'i& et a$5% 233;85
-o@ever% this h*#othesis @ou$d #redi+t +onsistent Doint )ronto,#arieta$ a+tivation in neuroi(a&in&
studies% @hi+h is +$ear$* not the +ase5 This (i&ht be e>#$ained b* the e>isten+e o) various
sub,+ir+uits @ithin the @ider net@orB% ea+h o) @hi+h (i&ht be s#e+ia$i'ed in s#e+i)i+ $in&uisti+
#ro+essin& +o(#onents 7e5&5% $e>i+a$ vs5 se(anti+ vs5 (or#ho$o&i+a$85 I) this @ere the +ase% there
@ou$d be no reason to e>#e+t )ronto,te(#ora$ a+tivation in a$$ neuroi(a&in& studiesE @e shou$d
:2
instead be ab$e to identi)* +onsistent asso+iations bet@een s#e+i)i+ tasBs 7i5e5% s#e+i)i+ +o&nitive
#ro+esses8 and s(a$$er net@orBs o) )ronta$ and #arieta$ areas5 -o@ever% i(a&in& data turned out to
be in+onsistent even a(on& studies usin& the sa(e tasBE on+e a&ain% this $a+B o) +onsisten+* @ithin
tasBs (aBes +o(#arin& resu$ts a+ross tasBs o) $itt$e use5
The (ost re+ent )"RI studies ado#tin& a )a+toria$ a##roa+h 7Ber$in&eri et a$5% 233=E Pa$ti et a$5%
233?8 have +ertain$* taBen a ste# )or@ard b* sear+hin& )or brain areas that sho@ &ra((ati+a$ +$ass
e))e+ts +onsistent$* a+ross di))erent tasBs ta##in& on si(i$ar +o&nitive #ro+esses5 These studies have
#rovided +onvin+in& eviden+e that retro,ro$andi+ areas 7$iBe the $e)t su#erior te(#ora$ su$+us and
the #osterior #arieta$ re&ion8 #$a* an i(#ortant ro$e in the $e>i+a$,se(anti+ #ro+essin& o) verbsE the
Auestion ho@ever re(ains as to @h* these areas have not been )ound (ore +onsistent$* in #revious
studies5 One #ossib$e reason is that #revious studies @ere not so s#e+i)i+ in their )o+us on $e>i+a$
#ro+essin& as Ber$in&eri et a$5 7233=8 and thus )ound a+tivations that a$so re)$e+ted other +o&nitive
$eve$sE (oreover% tasB,s#e+i)i+ a+tivations (i&ht have b$urred the resu$ts in sin&$e,tasB #aradi&(s%
even @hen $e>i+a$ #ro+essin& @as @e$$ tar&eted throu&h +are)u$ desi&n o) the e>#eri(enta$ and the
base$ine tasBs5 In addition to these desi&n,re$ated e>#$anations% (ore &enera$ )a+tors su+h as
di))eren+es in the $e>i+a$,se(anti+ +hara+teristi+s 7e5&5% @ord )reAuen+*% a+tiona$it*8 o) the verbs
used in the di))erent studies (i&ht have +ontributed to the underesti(ation o) the ro$e o) the #arieta$
and #osterior te(#ora$ areas in verb #ro+essin&5
Su+h an inho(o&eneous #attern o) resu$ts (i&ht de#end on the )a+t that nouns nor(a$$* re)er to
obDe+ts @hi$e verbs (ost$* re)er to a+tions% thus (aBin& the $e>i+a$,s*nta+ti+ di+hoto(* hard to
distin&uish )ro( the +on+e#tua$ one5 Indeed% so(e authors have dis+ussed their resu$ts in ter(s o)
obDe+t,vs5,a+tion e))e+ts rather than in ter(s o) &ra((ati+a$ +$ass 7e5&5% Cab$e et a$5% 233285
"oreover% se(anti+ +ate&or* e))e+ts have been de(onstrated @ithin &ra((ati+a$ +$ass in severa$
studies5 This h*#othesis is @eaBened b* the ar&u(ent that re#$i+abi$it* o) resu$ts shou$d be
::
inde#endent )ro( their theoreti+a$ inter#retation. )or e>a(#$e% @hen +o(#arin& the brain a+tivations
e$i+ited b* O+aBeP and O@ritin&P% the sa(e resu$ts shou$d e(er&e in t@o identi+a$ e>#eri(ents
irres#e+tive o) @hether the di))eren+e is due to &ra((ati+a$ +$ass% or to +aBe bein& an obDe+t and
eatin& an a+tion5
Another i(#ortant $e>i+a$,se(anti+ distin+tion +ontrasts (ove(ent verbs vs5 non,(ove(ent
verbs5 A++ordin& to re+ent )indin&s 7e5&5% BaB% RanBo#ou$ou% Nestor% et a$5% 23348% the se(anti+
re#resentations in the )or(er +ate&or* are heavi$* based on (otor in)or(ation% thus re$*in& on
+o(#$ete$* di))erent neura$ +ir+uits in+$udin& the #osterior #arieta$ 7Ber$in&eri et a$5% 233=8 and the
#ri(ar* (otor +orte> 7-auB% Iohnsrude% G Pu$ver(u$$er% 233<85 A$thou&h so(e data are in $ine
@ith this su&&estion 7e5&5% Ce((erer% !on'a$e' Casti$$o% Ta$ava&e% Patterson% G i$e*% 233=8% the
resu$ts o) t@o studies that investi&ated too$,re$ated (otor verbs in the +onte>t o) the sa(e
e>#eri(enta$ tasB 7i5e5% #i+ture na(in&8 are ver* in)or(ative in this res#e+tE @hi$e Sa++u(an et a$5
723348 re#ort that verbs a+tivate the $e)t in)erior #arieta$ $obu$e% the ri&ht )usi)or( &*rus% and the $e)t
+erebe$$u( (ore than nouns% Da(asio et a$5 723318 des+ribe si&ni)i+ant verb,re$ated a+tivations in
the (id te(#ora$ &*rus bi$atera$$*5 -en+e the neura$ +ir+uits des+ribed in the t@o studies do not
over$a# even thou&h the sa(e +$ass o) verbs @as investi&ated5 So(e re+ent resu$ts a$so su&&est the
e>isten+e o) verb,s#e+i)i+ a+tivation in the #osterior,$atera$,te(#ora$ +orte>% inde#endent$* o)
@hether verbs re)er to (otion or to (enta$ states 7Bedn*% Cara(a''a% !ross(an% Pas+ua$,Leone% G
Sa>e% 233=E see a$so A&&uDaro et a$5% 2334% )or si(i$ar )indin&s in neuro#s*+ho$o&*85 O) +ourse%
these +onsiderations do not den* that di))erent t*#es o) verbs in di))erent studies (i&ht be the +ause
o) these dis+re#an+iesE ho@ever% this )a+tor does not see( to be the #ri(ar* +ause o) su+h vast
in+onsisten+* in the data5
I) not e>#$ained b* $e>i+a$,se(anti+ +o,var*in& )a+tors% this $a+B o) +onsisten+* a+ross studies
investi&atin& the anato(i+a$ under#innin&s o) noun and verb #ro+essin& (i&ht su&&est% (ore
:<
si(#$*% that the neura$ +ir+uits u$ti(ate$* distin&uishin& nouns and verbs on anato(i+a$ &rounds are
not s#atia$$* se&re&ated% $eadin& to noun, or verb,s#e+i)i+ brain areas that +an be re$iab$* dete+ted
b* neuroi(a&in&% T"S% or anato(o,+orre$ative studies5 Rather% these neura$ +ir+uits @ou$d be
$o+ated s#atia$$* +$ose to ea+h other in the brain areas that are (ost )reAuent$* re#orted in the
$iterature% na(e$*% the LIF!% the $e)t insu$a% the (idd$e and in)erior $atera$ te(#ora$ re&ions and the
$e)t in)erior and su#erior #arieta$ &*ri% @hi+h (ost $iBe$* a$so host neura$ +ir+uits that are
res#onsib$e )or both noun and verb #ro+essin&5 "oreover% these areas (i&ht a$so under$ie +o&nitive
#ro+esses that are not s#e+i)i+a$$* $in&uisti+ in nature% but @hi+h are +a$$ed u#on b* (ost tasBs used
to e$i+it noun or verb #rodu+tion and +o(#rehension 7e5&5% tasB (onitorin&% attentiona$ #ro+esses%
+o&nitive resour+e assi&n(entE see Bedn* and Tho(#son,S+hi$$% 2334E Ber$in&eri et a$5% 233=E
Cres+entini% Sha$$i+e% G "a+a$uso% 233;85
O) +ourse% this h*#othesis does not i(#$* that the neura$ +ir+uits under$*in& noun and verb
#ro+essin& over$a# +o(#$ete$*5 Certain$*% there are neura$ a&&re&ates that are not +a$$ed u#on b*
both &ra((ati+a$ +$assesE other@ise% neuro#s*+ho$o&i+a$ disso+iations in brain,da(a&ed #atients
@ou$d never be #ossib$e5 hat @e are su&&estin& here is that &ra((ati+a$,+$ass s#e+i)i+ +ir+uits
(a* not +$uster into se#arate brain areasE rather% the* (a* be dis#ersed in di))erent #arts o) the brain
and be inter$eaved @ith neura$ stru+tures that are shared b* nouns and verbs and that a$so sub,serve
other +o&nitive )un+tions5 This s#e+i)i+ation shou$d a$so (aBe +$earer that this #ro#osa$ is indeed
+o(#atib$e @ith those neuroi(a&in& e>#eri(ents that do )ind noun,verb e))e+ts5 These e))e+ts%
ho@ever% (a* not be easi$* re#$i+ated be+ause &ra((ati+a$,+$ass,s#e+i)i+ stru+tures do not rea+h
the +riti+a$ di(ension to be+o(e +onsistent$* visib$e at the s#atia$ reso$ution nor(a$$* a$$o@ed b*
+urrent neuroi(ain& e>#eri(ents5
This su&&estion @ou$d a$so be in $ine @ith eviden+e )ro( the )"RI and "E! studies +arried
out b* Li$Destro( and +o$$ea&ues 7233=% 233;85 These studies su&&est that the neura$ net@orBs
:F
under$*in& noun and verb #ro+essin& over$a# )or the (ost #artE in addition% the a+tivation d*na(i+s
7i5e5% the te(#ora$ distribution o) the neura$ a+tivations8 are a$so ver* si(i$ar )or nouns and verbs5
hat this h*#othesis i(#$ies is that there is no one,to,one (a##in& bet@een anato(i+a$ and
+o&nitive units at the $eve$ o) ana$*sis that is +urrent$* ado#ted b* (ost anato(o,)un+tiona$ studies
on the noun,verb di+hoto(*5 On the one hand% the anato(i+a$ re&ions 7&*ri or Brod(ann areas8
usua$$* hi&h$i&hted in i(a&in& studies do not rea$$* +orres#ond to sin&$e )un+tiona$ units% but o)ten
+orres#ond to severa$ inde#endent )un+tiona$ units 7)or e>a(#$e% the LIF! has been re#orted to be
res#onsib$e )or the (or#ho$o&i+a$ #ro+essin& o) verbs% the se(anti+ #ro+essin& o) verbs% the
se(anti+ #ro+essin& o) nouns% the $e>i+a$ retrieva$ o) nouns% and even (ore &enera$% non,$in&uisti+
abi$ities $iBe the re+o$$e+tion o) autobio&ra#hi+a$ (e(ories% e5&5% "eu$enbroeB% RiD#Be(a% Cesse$s%
RiBBert G Fernande'% 231385 On the other hand% an* sin&$e +o&nitive #ro+ess +an be +arried out in
+ir+uits that are s#read over severa$ brain areas5 Thus% the in+onsisten+* a+ross studies (a* arise
)ro( the )a+t that our te+hni+a$% statisti+a$ and theoreti+a$ too$s do not a$$o@ to +a#ture univo+a$
+orre$ations bet@een the anato(i+a$ and the )un+tiona$ side5 To i$$ustrate @ith an e>a(#$e% su##ose
that the LIF! hosts #art o) the neura$ +ir+uit res#onsib$e )or the $e>i+a$ retrieva$ o) nouns and #art o)
the neura$ +ir+uit res#onsib$e )or the $e>i+a$ retrieva$ o) verbs% and that these +ir+uits are (odu$ated
b* the +o&nitive de(and i(#osed b* the tasB5 In su+h a situation% it is eas* to see that a+tivation in
the LIF! (a* e(er&e in asso+iation @ith nouns% or @ith verbs% or @ith both% or @ith neither
&ra((ati+a$ +$ass% de#endin& on the s#e+i)i+ e>#eri(enta$ settin&s o) the stud* 7e5&5% tasB% sti(u$i%
statisti+a$ ana$*sis85
Sahin et a$5 7233;8 have re+ent$* #ub$ished a stud* that is i$$u(inatin& in this res#e+t% even i) it
on$* (ar&ina$$* addresses the issue o) &ra((ati+a$ +$ass5 Usin& Intra,+rania$ E$e+tro#h*sio$o&*
7ICE% a te+hniAue that has the advanta&e o) bein& Auite a++urate both in ti(in& and in s#atia$
reso$ution8% the* )ound eviden+e )or $e>i+a$% s*nta+ti+% and #hono$o&i+a$ #ro+essin& in Bro+a0s area
:4
7BA <F8 at di))erent ti(e @indo@s a)ter the #resentation o) the sti(u$us 7X233 (s% X :23 (s% and X
<F3 (s% res#e+tive$*8E these authors a$so re#ort that the neuro)un+tiona$ si&na$ re)$e+tin& these
+o&nitive ste#s is si(i$ar )or nouns and verbs5 There are severa$ reasons @h* this stud* he$#s to
+$ari)* this h*#othesis5 First$*% the resu$ts sho@ that a uniAue brain area 7Bro+a0s area8 sub,serves
di))erent $eve$s o) #ro+essin& 7$e>i+a$% s*nta+ti+% and #hono$o&i+a$8% i5e5% anato(o,)un+tiona$
(a##in& is not one,to,one% but one,to,(an*5 Se+ond$*% the di))erent ti(e @indo@s identi)ied b*
Sahin et a$5 are @e$$ be$o@ the te(#ora$ reso$ution o) )"RIE it @ou$d there)ore a##ear that this
te+hniAue @ou$d never be ab$e to disentan&$e $e>i+a$% s*nta+ti+% and #hono$o&i+a$ #ro+essin& in
Bro+a0s area% un$ess the desi&n o) the e>#eri(enta$ and base$ine tasBs @ere to iso$ate one o) these
$eve$s o) #ro+essin& 5 Third$*% even at su+h a )ine te(#ora$ and s#atia$
:
reso$ution% +ir+uits )or noun
and verb #ro+essin& @ere not distin&uishab$e at $e>i+a$% &ra((ati+a$ or #hono$o&i+a$ $eve$E this
@ou$d i(#$* that either #ro+essin& o) nouns and verbs re$* on the sa(e +ir+uitr* in this area% or that
brain +e$$s de#uted to noun and verb #ro+essin& are indeed se#arate but inter(in&$ed in this sa(e
area5
On the basis o) these +onsiderations% it see(s that a +$ear understandin& o) the +erebra$ +ir+uits
res#onsib$e )or noun and verb #ro+essin& reAuires neuroi(a&in& studies to.
7i8 Condu+t a )ine,&rained +o&nitive ana$*sis o) the tasBs usedE
7ii8 Desi&n e>#eri(enta$ and base$ine tasBs to s#e+i)i+a$$* iso$ate one sin&$e +o&nitive #ro+essE
7iii8 "aBe use o) )a+toria$ desi&ns that a$$o@ better +ontro$ over tasB,s#e+i)i+ e))e+ts and a$$o@
to identi)* areas that are s#e+i)i+ )or &ra((ati+a$ +$ass a+ross tasBs 7throu&h +onDun+tion ana$*sis8E
3
It is di))i+u$t to deter(ine #re+ise$* the s#atia$ reso$ution o) Sahin et a$50s 7233;8 intra+rania$
re+ordin&sE ho@ever% it @as +ertain$* in the ran&e o) 1,2 ((% +onsiderin& that at $east 1< bi#o$ar
+hanne$s @ere re+orded in the Bro+a0s area o) ea+h #atient5
:?
7iv8 "aBe )u$$ use o) the s#atia$ reso$ution a+hieved b* i(a&in& te+hniAues% i5e5% do not $i(it
their ana$*ses to the Brod(ann areas or anato(i+a$ &*ri% but re#ort and +o((ent their resu$ts on the
basis o) s(a$$er +orti+a$ areasE
7v8 Sear+h )or +onver&in& eviden+e )ro( +o(#$e(entar* te+hniAues $iBe "E! and EE!%
@hi+h o))er better te(#ora$ reso$ution than )"RI5
These &uide$ines +ou$d he$# resear+hers to address the issue o) nounNverb #ro+essin& in the
brain at a $eve$ o) ana$*sis @here the re$ationshi# bet@een )un+tiona$ and anato(i+a$ units is $iBe$*
to be +$earer% thus )a+i$itatin& (ore +onsistent and re#$i+ab$e resu$ts5
As a +$osin& re(arB% @e note that the data revie@ed in this #a#er do not +onstitute eviden+e
a&ainst )un+tiona$ (odu$arit* in the hu(an (ind% and do not e>+$ude anato(i+a$ (odu$arit* in
other +o&nitive do(ains 7e5&5% obDe+t identi)i+ation% Boro@sB*% Loher% Ce$$and et a$5% 233F8%
#rovided that an adeAuate s#atia$ reso$ution is +onsidered5
:=
Ac&no'ledgements
The #resent stud* @as +ondu+ted )or the (ost #art at the Universit* o) "i$ano,Bi+o++a%
a$thou&h the (anus+ri#t @as #artia$$* #re#ared @hi$e the )irst author @as at the Ro*a$ -o$$o@a*%
Universit* o) London% )unded b* a #ost,do+tora$ )e$$o@shi# &ranted b* the E+ono(i+ and So+ia$
Resear+h Coun+i$% United Cin&do(E he &rate)u$$* a+Bno@$ed&es this +ontribution5 This @orB has
been #art$* )unded b* the &rant PRIN 233?<ECLSQY332 a@arded to C$audio Lu''atti b* the Ita$ian
"inistr* o) Universit* and Resear+h% and b* the &rant PRIN 233?ILCBL;Y332 a@arded to Era$do
Pau$esu b* the sa(e institution5
:;
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Leve$t% 5I5"5% Roe$o)s% A5P5A5% G "e*er% A5S5 71;;;85 A theor* o) $e>i+a$ a++ess in s#ee+h
#rodu+tion5 Behaviora$ and Brain S+ien+es, 22% 1,:?5
Li% P5% Iin% H5% G Tan% L5-5 7233<85 Neura$ re#resentation o) nouns and verbs in Chinese. an )"RI
stud*5 NeuroI(a&e% 21% 1F::,1F<15
Li$DestrS(% "5% -u$tTn% A5% ParBBonen% L5% G Sa$(e$in% R5 7233;85 Co(#arin& "E! and )"RI
vie@s to na(in& a+tions and obDe+ts5 -u(an Brain "a##in&, :3% 1=<F,1=F45
Li$DestrS(% "5% TarBiainen% A5% Parviainen% T5% CuDa$a% I5% Nu((inen% I5% -i$tunen% I5% et a$5 7233=85
Per+eivin& and na(in& a+tions and obDe+ts5 Neuroi(a&e, <1% 11:2,11<15
Lo !er)o% E5% O$ivieri% "5% Torriero% S5% Sa$erno% S5% Co+h% !5 G Ca$ta&irone C5 7233=85 The
in)$uen+e o) rT"S over #re)ronta$ and (otor areas in a (or#ho$o&i+a$ tasB. !ra((ati+a$ vs5
se(anti+ e))e+ts. Neuro#s*+ho$o&ia, <4% ?4<,??35
Lon&e% O5% Randa$$% B5% Sta(ataBis% E5A5% G T*$er% L5C5 7233?85 !ra((ati+a$ +ate&ories in the
<<
brain. The ro$e o) (or#ho$o&i+a$ stru+ture5 Cerebra$ Corte>% 1?% 1=12,1=235
Lu''atti% C5% Ra&&i% R5% Hon+a% !5% Pistarini% C5% Contardi% A5% G Pinna% !5D5 7233285 Verb,noun
doub$e disso+iation in a#hasi+ $e>i+a$ i(#air(ents. The ro$e o) @ord )reAuen+* and
i(a&eabi$it*5 Brain and Lan&ua&e, =1, <:2,<<<5
"aran&o$o% P5% Piras% F5% !a$ati% !5% G Burani% C5 7233485 Fun+tiona$ anato(* o) derivationa$
(or#ho$o&*5 Corte>% <2% 13;:,11345
"artin% A5% i&&s% C5 L5% Un&er$eider% L5 !5% G -a>b*% I5 V5 71;;485 Neura$ +orre$ates o)
+ate&or*,s#e+i)i+ Bno@$ed&e5 Nature% :?;% 4<;,4F25
"Mt'i&% S5% DruBs% I5% "asterson% I5% G Vi&$io++o% !5 7233;85 Noun and verb di))eren+es in #i+ture
na(in&. #ast studies and ne@ eviden+e5 Corte>, <F% ?:=,?F=5
"+Carth*% R5% G arrin&ton% E5C5 71;=F85 Cate&or* s#e+i)i+it* in an a&ra((ati+ #atient. the
re$ative i(#air(ent o) verb retrieva$ and +o(#rehension5 Neuro#s*+ho$o&ia% 2:% ?3;,?2?5
"eu$enbroeB% O5% RiD#Be(a% "5% Cesse$s% R5P5C5% RiBBert% "5!5"5O5% G Fernande'% !5 7231385
Autobio&ra#hi+a$ (e(or* retrieva$ in #atients @ith A$'hei(er0s disease5 Neuroi(a&e% F:%
::1,:<35
"i+e$i% !5% Si$veri% "5C5% Vi$$a% !5% G Cara(a''a% A5 71;=<85 On the basis )or the a&ra((ati+Vs
di))i+u$t* in #rodu+in& (ain verbs5 Corte>% 23% 23?,2<35
Ne@(an% S5D5% T@ie&% D5B5% G Car#enter% P5A 7233185 Base$ine +onditions and subtra+tive $o&i+ in
neuroi(a&in&5 -u(an Brain "a##in&% 12% 22=,2:F5
ODe(ann% I5!5% ODe(ann% !5A5% G Letti+h% E5 7233285 Corti+a$ sti(u$ation (a##in& o) $an&ua&e
+orte> b* usin& a verb &eneration tasB. e))e+ts o) $earnin& and +o(#arison to (a##in& based on
obDe+t na(in&5 Iourna$ o) Neurosur&er*% ;?% ::,:=5
O$iveri% "5% Fino++hiaro% C5% Sha#iro% C5% !an&itano% "5% Cara(a''a% A5% G Pas+ua$ Leone% A5
7233<85 A$$ ta$B and no a+tion. A trans+rania$ (a&neti+ sti(u$ation stud* o) (otor +orte>
<F
a+tivation durin& a+tion @ord #rodu+tion5 Iourna$ o) Co&nitive Neuros+ien+e% 14% :?<,:=15
Pa$ti% D5% Sha+har% "5B5% -end$er% T5% G -adar% U5 7233?85 Neura$ +orre$ates o) se(anti+ and
(or#ho$o&i+a$ #ro+essin& o) -ebre@ nouns and verbs5 -u(an Brain "a##in&% 2=% :3:,:1<5
Pau$esu% E5% Sha$$i+e% T5% Dane$$i% L5% Fra+Bo@iaB% R5S5I5% G Frith% C5D5 7sub(itted85 Anato(i+a$
(odu$arit* o) verba$ @orBin& (e(or*. eviden+e )ro( a (u+h studied #atient5
Perani% D5% Ca##a% S5F5% S+hnur% T5% Tetta(anti% "5% Co$$ina% S5% Rosa% "5"5% G Fa'io% F5 71;;;85
The neura$ +orre$ates o) verb and noun #ro+essin&5 A PET stud*5 Brain, 122, 2::?,2:<<5
Perea% "5 G Rosa% E5 7233285 The e))e+ts o) asso+iative and se(anti+ #ri(in& in the $e>i+a$ de+ision
tasB5 Ps*+ho$o&i+a$ Resear+h% 44% 1=3,1;<5
Ra##% B5% G Cara(a''a% A5 7233285 Se$e+tive di))i+u$ties @ith s#oBen nouns and @ritten verbs. a
sin&$e +ase stud*5 Iourna$ o) Neuro$in&uisti+s% 1F% :?:,<325
Rast$e5 C5 G Br*sbaert% "5 7233485 "asBed #hono$o&i+a$ #ri(in& e))e+ts in En&$ish. Are the* rea$?
Do the* (atter? Co&nitive Ps*+ho$o&*% F:% ;?,1<F5
Robins% R5-5 71;F285 Noun and verb in universa$ &ra((ar5 Lan&ua&e, 2=% 2=;,2;=5
Ro@an% A5% LiT&eois% F5% Var&a,Chade(% F5% !adian% D5% Conne$$*% A5% G Ba$de@e&% T5 7233<85
Corti+a$ $atera$i'ation durin& verb &eneration. a +o(bined ERP and )"RI stud*5 NeuroI(a&e%
22% 44F,4?F5
Sabsevit'% D5 S5% "ed$er% D5 A5% Seidenber&% "5% G Binder% I5 R5 7233F85 "odu$ation o) the se(anti+
s*ste( b* @ord i(a&eabi$it*5 Neuroi(a&e% 2?% 1==,2335
Sa++u(an% "5C5% Ca##a% S5F5% Bates% E5A5% Areva$o% A5% De$$a Rosa% P5% Danna% "5% G Perani% D5
7233485 The i(#a+t o) se(anti+ re)eren+e on @ord +$ass. An )"RI stud* o) a+tion and obDe+t
na(in&5 NeuroI(a&e, :2% 1=4F,1=?=5
Sahin% N5 T5% PinBer% S5% Cash% S5 S5% S+ho(er% D5% G -a$&ren% E5 7233;85 SeAuentia$ #ro+essin& o)
$e>i+a$% &ra((ati+a$% and #hono$o&i+a$ in)or(ation @ithin Bro+a0s area5 S+ien+e% :24% <<F,<<;5
<4
Sahin% N5T5% PinBer% S5% G -a$&ren% E5 7233485 Abstra+t &ra((ati+a$ #ro+essin& o) nouns and verbs
in Bro+aVs area. Eviden+e )ro( )"RI5 Corte>% <2% F<3,F425
Sha#iro% C5% G Cara(a''a% A5 7233:85 Loo(in& a $oo(. Eviden+e )or inde#endent a++ess to
&ra((ati+a$ and #hono$o&i+a$ #ro#erties in verb retrieva$5 Iourna$ o) Neuro$in&uisti+s% 14%
=F,1115
Sha#iro% C5% "oo% L5R5% G Cara(a''a% A5 7233485 Corti+a$ si&natures o) noun and verb #rodu+tion5
Pro+eedin&s o) the Nationa$ A+ade(* o) S+ien+es o) USA% 13:% 14<<,14<;5
Sha#iro% C5% "otta&h*% F5"5% S+hi$$er% N5O5% Poe##e$% T5D5% F$uss% "5O5% "u$$er% -55% Cara(a''a%
A5% G Crause% B5I5 7233F85 Disso+iatin& neura$ +orre$ate )or verbs and nouns5 Neuroi(a&e% 2<%
13F=,134?5
Sha#iro% C5% Pas+ua$,Leone% A5% "otta&h*% F5"5% !an&itano% "5% G Cara(a''a% A5 7233185
!ra((ati+a$ distin+tion in the $e)t )ronta$ +orte>5 Iourna$ o) Co&nitive Neuros+ien+e% 1:% ?1:,
?235
Sha#iro% C5% She$ton% I5% G Cara(a''a% A5 7233385 !ra((ati+a$ +$ass in $e>i+a$ #rodu+tion and
(orh#o$o&i+a$ #ro+essin&. Eviden+e )ro( a +ase o) )$uent a#hasia5 Co&nitive
Neuro#s*+ho$o&*% 1?% 44F,4=25
Si$veri% "5C5% G Di Betta% A5"5 71;;?85 Noun,verb disso+iations in brain,da(a&ed #atients. )urther
eviden+e5 Neuro+ase, :% <??,<==5
Si$veri% "5% Perri% R5% G Ca##a% A5 7233:85 !ra((ati+a$ +$ass e))e+ts in brain,da(a&ed #atients.
Fun+tiona$ $o+us o) noun and verb de)i+it5 Brain and Lan&ua&e, =F, <;,445
Siri% S5% Tetta(anti% "5% Ca##a% S5F5% De$$a Rosa% P5% Sa++u(an% C5% S+i)o% P5% G Vi&$io++o% !5
7233=85 The neura$ substrate o) na(in& events. E))e+ts o) #ro+essin& de(ands but not
&ra((ati+a$ +$ass5 Cerebra$ Corte>% 1=% 1?1,1??5
SSrSs% P5% Corne$issen% C5% Laine% "5% G Sa$(e$in% R5 7233:85 Na(in& a+tions and obDe+ts. +orti+a$
<?
d*na(i+s in hea$th* adu$ts and in an ano(i+ #atient @ith a disso+iation in a+tionNobDe+t
na(in&5 Neuroi(a&e, 1;% 1?=?,1=315
Tho(#son% C5C5% BonaBdar#our% B5% Fi>% S5C5% B$u(en)e$d% -5C5% Parrish% T5B5% !ite$(an% D5R5% G
"arse$,"esu$a(% "5 7233?85 Neura$ +orre$ates o) verb ar&u(ent stru+ture #ro+essin&5 Iourna$
o) Co&nitive Neuros+ien+e% 1;% 1?F:,1?4?5
Trane$% D5% !rabo@sBi% T5 I5% L*on% I5% G Da(asio% -5 7233F85 Na(in& the sa(e entities )ro( visua$
or )ro( auditor* sti(u$ation en&a&es si(i$ar re&ions o) $e)t in)erote(#ora$ +orti+es5 Iourna$ o)
Co&nitive Neuros+ien+e% 1?% 12;:,:3F5
Trane$% D5% "an'e$% C5% As#% E5% G Ce((erer% D5 7233=85 Na(in& d*na(i+ and stati+ a+tions.
Neuro#s*+ho$o&i+a$ eviden+e5 Iourna$ o) Ph*sio$o&* , Paris % 132% =3,;<5
Trane$% D5% "artin% C5% Da(asio% -5% !rabo@sBi% T5 I5% G -i+h@a% R5 7233F85 E))e+ts o) noun,verb
ho(on*(* on the neura$ +orre$ates o) na(in& +on+rete entities and a+tions5 Brain and
Lan&ua&e% ;2% 2==,2;;5
T*$er% L5C5% Bri&ht% P5% F$et+her% P5% G Sta(ataBis% E5A5 7233<85 Neura$ #ro+essin& o) nouns and
verbs. the ro$e o) in)$e+tiona$ (or#ho$o&*5 Neuro#s*+ho$o&ia% <2% F12,F2:5
T*$er% L5% Randa$$% B5% G Sta(ataBis% E5 7233=85 Corti+a$ di))erentiation )or nouns and verbs de#ends
on &ra((ati+a$ (arBers5 Iourna$ o) Co&nitive Neuros+ien+e, 23% 1:=1,1:=;5
T*$er% L5C5% Russe$$% R5% Fadi$i% I5% G "oss% -5E5 7233185 The neura$ re#resentation o) nouns and
verbs. PET studies5 Brain, 12<% 141;,14:<5
T*$er% L5C5% Sta(ataBis% E5A5% Di+B% E5% Bri&ht% P5% F$et+her% P5% G "oss% -5E5 7233:85 ObDe+ts and
their a+tions. eviden+e )or a neura$$* distributed se(anti+ s*ste(5 NeuroI(a&e, 1=% F<2,FF?5
RoBo*a(a% S5% "i*a(oto% T5% Riera% I5% Ci(% I5% ABitsuBi% R5% I@ata% C5% Roshi(oto% C5% -orie% C5%
Sato% S5% G Ca@ashi(a% R5 7233485 Corti+a$ (e+hanis(s invo$ved in the #ro+essin& o) verbs.
An )"RI stud*5 Iourna$ o) Co&nitive Neuros+ien+e% 1=% 1:3<,1:1:5
<=
arburton% E5% ise% R5I5S5% Pri+e% C5I5% ei$$er% C5% -adar% U5% Ra(sa*% S5% G Fra+Bo@iaB% S5I5
71;;485 Noun and verb retrieva$ b* nor(a$ subDe+ts% Studies @ith PET5 Brain% 11;% 1F;,1?;5
Hin&eser% L5B5% G Berndt% R5S5 71;==85 !ra((ati+a$ +$ass and +onte>t e))e+ts in a +ase o) #ure
ano(ia. I(#$i+ations )or (ode$s o) $an&ua&e #ro+essin&5 Co&nitive Neuro#s*+ho$o&*% F%
<?:,F145
<;
Figure captions
Fi&ure 15 Per+enta&e o) studies that re#orted noun, or verb,re$ated a+tivations in an* &iven
brain re&ion5 B$ue nu(bers re)er to si(#$e e))e+ts o) nouns or verbs versus base$ine% @hereas red
nu(bers re)er to noun,(inus,verbs or verbs,(inus,nouns dire+t +o(#arisons5 The tota$ nu(ber o)
studies that +arried out si(#$e e))e+t ana$*sis in our sa(#$e is 23% @hereas the tota$ nu(ber o)
studies that #er)or(ed noun,verb dire+t +o(#arisons is 245 The anterior and #osterior #ortions o)
the te(#ora$ $obe are divided b* a $ine ori&inatin& )ro( the )ronto,ventra$ +orner o) -es+h$0s &*rus
and runnin& ortho&ona$$* to the (ain a>is o) the su#erior te(#ora$ su$+us5
F3
Tables
Table 1: PET and fMRI studies on nouns and/or verbs using lexical decision as experimental tas! "irect comparisons bet#een nouns and verbs are
listed in part $a%& #hereas simple effect anal'ses are illustrated in part $b%!
(uthors Techni)ue Threshold "esign *aseline +ontrast (rea
$a% "irect noun,verb comparison
-u.imai et al!
$1///%
fMRI
+ross,
correlation
coefficient&
e)uivalent to
!001
uncorrected
*loc
2ine orientation
.udgement
Nouns,3erbs
R and 2 inf! Parietal sulcus& R and 2 precentral
sulcus
3erbs,Nouns No area
Perani et al!
$1///%
PET
!001
uncorrected
*loc
2etter detection
tas
Nouns,3erbs No area
3erbs,Nouns
2 mid! and inf! -rontal g'rus4 2 sup Parietal lobule4 2
mid! and inf! Temp g'rus4 2 inf! 5ccipital g'rus!
1
2i et al! $6007% fMRI
!001
uncorrected
*loc -ixation Nouns,3erbs +audate Nucleus
3erbs,Nouns No area
Thompson et al!
$6008%
fMRI !01 corrected
Event,
related
-ixation Nouns,3erbs No area
3erbs,Nouns No area
T'ler et al!
$6001%
PET !01 corrected *loc
2etter detection
tas
Nouns,3erbs No area
3erbs,Nouns R substantia nigra
9oo'ama et al!
$600:%
fMRI !01 corrected *loc
Number of
characters
decision
Nouns,3erbs No area
3erbs,Nouns No area4 2 mid! Temporal g'rus $R5I anal'sis%
$b% ;imple effect anal'ses
6
-u.imai et al!
$1///%
fMRI
+ross,
correlation
coefficient&
e)uivalent to
!001
uncorrected
*loc
2ine orientation
.udgement
Nouns,baseline
2 *roca<s area& 2 Insula& 2 ant! +ingulate sulcus&
Precentral sulcus $bilaterall'%& 2 Postcentral sulcus&
bilateral Intraparietal sulcus& 2 ;upramarginal g'rus&
bilateral 5ccipito,Temporal sulcus& R *( 7:,78
3erbs,baseline
*ilateral *roca<s area& Insula and *( 7:& bilateral
Precentral sulcus& 2 Postcentral sulcus& bilateral
Intraparietal sulcus& 2 ;upramarginal g'rus&
Precuneus& bilateral 5ccipito,temporal sulcus!
=
2i et al! $6007% fMRI
!001
uncorrected
*loc
design
-ixation Nouns,baseline
2 and R inf! -rontal g'rus& 2 and R Postcentral
g'rus& 2 and R Precentral g'rus& R ;'lvian fissure& 2
mid,superior Temporal g'rus& R mid Temporal& 2 inf!
Parietal g'rus& 2 mid 5ccipital g'rus& 2 -usiform
g'rus& 2 +uneus& R and 2 +ingulate& Thalamus&
Putamen& +audate Nucleus& >ippocampus&
2enticular Nucleus& R +erebellum
3erbs,baseline
2 and R inf! -rontal g'rus& 2 Precentral g'rus& 2
Insula& R Precentral g'rus& R ;'lvian fissure& R
medial -rontal g'rus& 2 mid,superior Temporal
g'rus& R inf! Parietal g'rus& 2 mid 5ccipital g'rus& 2
sup 5ccipital g'rus& 2 and R -usiform g'rus& 2
+uneus& Thalamus& 2 and R +erebellum
7
Thompson et al!
$6008%
fMRI !01 corrected
Event,
related
R inf! -rontal g'rus& R +entral ;ulcus& 2 Insula& 2
and R Precentral g'rus& 2 and R mid -rontal gr'us& R
sup! Temporal g'rus& 2 and R Parietal 2obule& 2 sup!
Parietal lobule& 2 +ingulate g'rus& 2 and R inf!
5ccipital g'rus& 2 mid 5ccipital g'rus& 2 and R
-usiform g'rus& 2 ?lobus Pallidus& 2 Putamen& 2
;usbtantia Nigra& R Thalamus $32 nucleus%
3erbs,baseline
2 inf! -rontal g'rus& 2 +entral ;ulcus& 2 Insula& 2
and R Precentral g'rus& 2 and R mid -rontal gr'us& R
sup! Temporal g'rus& 2 mid Temporal g'rus& 2 and R
Parietal 2obule& 2 sup! Parietal lobule& R inf!
5ccipital g'rus& 2 mid 5ccipital g'rus& 2 and R
-usiform g'rus& R +audate Nucleus& 2 Red Nucleus&
R Thalamus $M" nucleus%
1
Table 6: PET and fMRI studies on nouns and/or verbs using a semantic experimental tas. "irect comparisons bet#een nouns and verbs are listed in part $a%&
#hereas simple effect anal'ses are illustrated in part $b%!
(uthors Techni)ue Threshold "esign Experimental tas *aseline +ontrast (rea
*edn' et al!
$600:%
fMRI !01 corrected
Event,
related
;emantic
matching
Identit'
.udgement on
pseudo,#ords
Nouns,3erbs
2 inf! -rontal g'rus $R5I
anal'ses%
3erbs,Nouns
2 sup! Temporal g'rus $R5I
anal'ses%
"avis et al!
$6007%
fMRI !001 uncorrected
Event,
related
5ne,bac
s'non'm
monitoring
2etter identit'
matching
Nouns,3erbs No area
!01 corrected 3erbs,Nouns 2 post! Mid Temporal g'rus!
@able et al!
$6006%
fMRI !01 corrected *loc ;emantic
association
$pictures%
3isual matching 3erbs,Nouns 2 and R lateral
5ccipito,Temporal regions
$#hole,brain anal'ses%4 R and
:
2 postero,lateral Temporal
region& R and 2 5ccipital
sulcus $R5I anal'ses%!
;emantic
association
$printed #ords%
3isual matching 3erbs,Nouns
No area $#hole,brain
anal'ses%4 2 postero,lateral
Temporal region $R5I
anal'ses%!
2onge et al!
$6008%
Event,
related
Pleasantness
.udgement
-ixation
Noun stems,3erb
stems
No area
3erb stems,Noun
stems
No area
Inflected nouns,
Inflected verbs
No area
Inflected verbs,
Inflected nouns
2 post! mid Temporal g'rus
Palti et al!
$6008%
fMRI !01 uncorrected *loc ;emantic
.udgment
Passive
listening of
#ords pla'ed
3erbs,Nouns 2 inf! -rontal g'rus
8
bac#ard
T'ler et al!
$6001%
PET !01 corrected *loc
;emantic
categoriAation
2etter
categoriAation
tas
Nouns,3erbs No area
3erbs,Nouns R inf! Temporal g'rus
+on.unction
anal'sis $#ith
lexical decision%
Nouns,3erbs No area
3erbs,Nouns No area
T'ler et al!
$600=%
fMRI
!001 uncorrected
$!01 at cluster level%
Event,
related
;emantic
categoriAation
2etter
categoriAation
tas
Nouns,3erbs $tools% No area
3erbs,Nouns $tools%
No area $normal threshold%4 2
inf! -rontal g'rus and 2
Insula $threshold B !01%
Nouns,3erbs
$biological%
No area
3erbs,Nouns 2 inf! -rontal g'rus& 2 mid
C
$biological% Temporal g'rus
T'ler et al!
$6007%
fMRI
!001 uncorrected
Event,
related
;emantic
categoriAation
2etter
categoriAation
tas
Nouns,3erbs No area
3erbs,Nouns 2 inf! -rontal g'rus
T'ler et al
$600C%
fMRI
!001 $uncorrected%
*loc
Pleasantness
.udgement
Passive vie#ing Nouns,verbs 2 mid Temporal g'rus
3erbs,nouns No area
$b% ;imple effect anal'sis
T'ler et al!
$600=%
fMRI
!001 uncorrected
Event,
related
;emantic
categoriAation
2etter
categoriAation
tas
3erbs$tools%,
baseline
2 inf! -rontal g'rus& R ant!
+ingulate g'rus& 2 -usiform
and Parahippocampal g'rus&
2 2ingual g'rus& R inf!
-rontal g'rus& R sup!
temporal g'rus& R
+erebellum!
/
3erbs$biological%,
baseline
2 inf! -rontal g'rus& 2 sup!
Temporal g'rus& 2 -usiform
g'rus& 2 Parahippocampal
g'rus! 2 mid! Temporal
g'rus& 2 inf! Temporal g'rus&
+erebellum bilaterall'
T'ler et al!
$6007%
fMRI
!001 uncorrected
Event,
related
;emantic
categoriAation
2etter
categoriAation
tas
Nouns,baseline
2 inf! -rontal g'rus& 2 Insula&
2 mid -rontal g'rus& 2 inf!
Temporal g'rus& 2 -usiform
g'rus
3erbs,baseline
2 inf! -rontal g'rus& 2
Parahippocampal g'rus& 2
-usiform g'rus& 2 and R
+ingulate g'rus
T'ler et al
$600C%
fMRI
!001 $uncorrected%
*loc
Pleasantness
.udgement
Passive vie#ing Nouns,baseline
2 inf -rontal g'rus& 2
mid Temporal g'rus& Putamen
3erbs,baseline 2 inf -rontal g'rus& 2
10
mid Temporal g'rus&
Putamen& R +erebellum
11
Table =: PET and fMRI studies on nouns and/or verbs using a picture naming experimental tas! "irect comparisons bet#een nouns and verbs are listed in
part $a%& #hereas simple effect anal'ses are illustrated in part $b%!
*erlingeri et al! $600C% fMRI
!001
uncorrected
*loc
-orced,choice
figure naming
Nouns,3erbs No area
3erbs,Nouns
2 inf! -rontal g'rus& 2 Insula& R and 2
mid! Temporal g'rus& R inf! Temporal
g'rus& R and 2 post! Parietal g'rus& R
and 2 sup! and mid! 5ccipital g'rus&
"amasio et al! $6001% PET
!01
corrected
*loc
5rientation
.udgement of
unno#n faces
3erbs,Nouns 2 and R mid! Temporal g'rus
2il.estrDm et al $600C% fMRI !01 -"R *loc -ixation Nouns $5b.(ct%,3erbs
2 inf Parietale lobule& 2 superior
Parietal lobule& 2 Precuneus& R mid
-rontal g'rus& R superior parietal
lobule& R inf! Parietal lobule
16
3erbs,Nouns $5b.%
2 ;upramarginal g'rus& 2 mid
Temporal g'rus& 2 ;uperior Temporal
pole& 2 sup Medial -rontal g'rus& 2
Precentral g'rus& R mid Temporal
g'rus& R inf Temporal g'rus& R sup
anterior Temporal g'rus& R insula& R
+erebellum
;accuman et al! $600:% fMRI
!001
uncorrected
Event,
related
None Nouns,3erbs
R +uneus& R post! +ingulate cortex& R
+audate nucleus
3erbs,Nouns
No area $normal threshold%4 2 intra,
parietal sulcus& R -usiform g'rus& 2
+erebellum $lo#er threshold%
;iri et al! $600C% fMRI
!01
corrected
*loc None Nouns,3erbs 2 inf! -rontal g'rus
3erbs,Nouns No area
$b% ;imple effect anal'sis
1=
+hao et al! $6000% fMRI
!001
uncorrected
*loc
Passive vie#ing of
scrambled stimuli
Nouns,baseline
2 inf! -rontal g'rus& 2 Insula& 2
ventral Premotor cortex& 2 post!
Parietal cortex
2il.estrDm et al $600C% fMRI !001 -"R *loc -ixation Nouns$5b.%,baseline
2! Precentral g'rus& 2 ;M(& 2 inf!
-rontal g'rus& 2 mid! Temporal g'rus&
2 -usiform& 2 mid! 5ccipital& 2! inf
5ccipital& 2 sup! Parietal lobule& 2 inf!
Parietal lobule& R inf! Temporal g'rus&
R -usiform g'rus& R mid 5ccipital
g'rus& R inf! 5ccipital g'rus& R sup!
Parietal lobule
Nouns$5b.(ct%,baseline
2 inf! -rontal g'rus& 2 -usiform& 2
mid! 5ccipital& 2 sup! Parietal lobule&
2 inf! Parietal lobule& R -usiform
g'rus& R mid 5ccipital g'rus& R
Precuneus& R sup! Parietal lobule
17
3erbs,baseline
2! Precentral g'rus& 2 inf! -rontal
g'rus& 2! Insula& 2 sup! Temporal
g'rus& 2! mid! Temporal g'rus& 2
-usiform g'rus& 2 >ippocampus& 2
mid! 5ccipital g'rus& 2 inf 5ccipital
g'rus& R Precentral g'rus& R ;M(& R
mid! Temporal g'rus& R -usiform
g'rus& R >ippocampus& R inf
5ccipital g'rus
Martin et al! $1//:% PET
!001
uncorrected
*loc Passive vie#ing Nouns,baseline
2 medial 5ccipital lobe $animals%& 2
Premotor area $tools%& 2 inf! -rontal
g'rus& 2 medial Inferotemporal region
$animals and tools%
Tranel& ?rabo#si et al!
$6001%
PET
!01
corrected
*loc
Passive vie#ing of
scrambled stimuli
Nouns $tools%,baseline 2 post! 2at! Infero,temporal
Nouns $animals%,baseline 2 ant! ventral Infero,temporal
11
Tranel& Martin et al!
$6001%
PET
!01
corrected
*loc
5rientation
.udgement on
unfamiliar faces
Nouns$tools%,baseline 2 post! ventral Infero,temporal
3erbs,baseline
2 -rontal 5perculum& 2 mid!
Temporal& 2 post! lat! infero,temporal
1:
Table 7: PET and fMRI studies on nouns and/or verbs using a morphological experimental tas! "irect comparisons bet#een nouns and verbs are listed in
part $a%& #hereas simple effect anal'ses are illustrated in part $b%!
$a% "irect noun,verb comparisons
*erlingeri et al!
$600C%
fMRI !001 uncorrected *loc
?rammatical
class s#itching
-orced,choice
figure naming
Nouns,3erbs
2 inf! -rontal g'rus& 2 inf!
Parietal lobule& 2 Precuneus& 2
mid! and sup! 5ccipital g'rus!
3erbs,Nouns
R and 2 sup! -rontal g'rus& R and
2 Precentral g'rus& R and 2 ;M(
*urton et al
$600/%
fMRI
!01 corrected for
multiple
comparisons
*loc
Morphological
.udgment
Tone
discrimination
Nouns,3erbs 2 inf -rontal g'rus
3erbs,Nouns No area
;hapiro et al!
$6001%
PET !001 uncorrected *loc
Eord and pseudo,
#ord inflection
Pseudo,#ord
reading
Nouns,3erbs
2 -usiform& R mid! sup! Temporal
g'rus& R Insula& R +erebellum
3erbs,Nouns
2 sup! -rontal g'rus& 2 inf!
-rontal g'rus& R cerebellum
18
$Nouns,3erbs%
and
$Pseudonouns,
Pseudoverbs%
R mid! sup! Temporal g'rus& 2
-usiform g'rus
$3erbs,Nouns%
and
$Pseudoverbs,
Pseudonouns%
2 sup! -rontal g'rus
;ahin et al!
$600:%
fMRI !01 corrected
Event,
related
;entence
completion #ith
inflected #ords
-ixation Nouns,3erbs
Intraparietal sulcus
3erbs,Nouns No area
;hapiro et al!
$600:%
Event,
related
Regular/irregular
abstract/concrete
#ord inflection
-ixation Nouns,3erbs 2 -usiform g'rus
1C
3erbs,Nouns
2 Prefrontal cortex& 2 sup!
Parietal& 2 sup! Temporal cortex
Palti et al!
$6008%
fMRI !01 uncorrected *loc
Morphological
.udgement
Passive listening
of #ords pla'ed
bac#ards
3erb,Nouns 2 Premotor area
"e "iego
*alaguer et al!
$600:%
Event,
related
Eord and
pseudo#ord
inflection tas
Repetition 3erbs,baseline
+erebellum& R Parahippocampal
g'rus& R ;ensorimotor cortex
$regular verbs%4 2 inf! -rontal
g'rus& 2 dorso,lateral Prefrontal
cortex& R sensorimotor cortex& 2
+erebellum $irregular verbs%!
1/
Marangolo et al!
$600:%
fMRI
!01 $!01 corrected
at cluster level%
*loc "erivation Repetition Nouns,baseline
2 and R inf! -rontal g'rus& 2 and
R mid -rontal g'rus& 2 Precental
g'rus& 2 Insula& 2 ;M(& 2 mid
Temporal g'rus& 2 and R inf!
Parietal 2obule& 2 and R (ngular
?'rus& 2 and R sup! Parietal
lobule&
3erbs,baseline
2 inf! -rontal g'rus& 2 Precentral
g'rus
Inflection Repetition Nouns,baseline
R ant! (nd mid +ingulum& R sup!
Temporal g'rus
3erbs,baseline
2 inf! and mid -rontal g'rus& 2
Precentral g'rus& 2 Paracentral
lobule& 2 ;M(& 2 inf! and sup!
Parietal lobule& 2 (ngular g'rus&
2 inf! Temporal g'rus
60
;ahin et al!
$600:%
fMRI !01 corrected
Event,
related
;entence
completion #ith
inflected #ords
2 inf! -rontal g'rus& 2 ant! Insula&
2 ;M(& 2 Intraparietal ;ulcus
3erbs,baseline
2 inf! -rontal g'rus& 2 ant! Insula&
2 ;M(
61
Table 1: PET and fMRI studies on nouns and/or verbs using a fluenc' experimental tas! "irect comparisons bet#een nouns and verbs are listed in part $a%&
#hereas simple effect anal'ses are illustrated in part $b%!
Earburton et al!
$1//:%
PET
!001
uncorrected
*loc ;emantic fluenc' Rest 3erb,Nouns
2 inf! -rontal g'rus& 2 ;M(& 2
Precentral sulcus& 2 inf! Temporal
sulcus& 2 Temporo,Parietal .unction& 2
inf! Parietal lobe
-rings et al!
$600:%
fMRI
!001
uncorrected
*loc 3erb generation
3erb reading&
Noun reading
3erbs generation,
3erb reading
2 inf! -rontal g'rus& 2 sup! Temporal
g'rus& R +erebellum
3erbs generation,
Noun reading
No area
66
Ro#an et al!
$6007%
fMRI
!01
corrected
*loc 3erb generation
Passive
listening
3erbs,baseline
2 inf! -rontal cortex& 2 and R -rontal
operculum& 2 and R Insula& 2
Premotor cortex& 2 ;M(& 2
;upramarginal g'rus& 2 inf! Temporal
g'rus& 2 and R post! ;up! Temporal
cortex& 2 Putamen& R +erebellum!
Earburton et al!
$1//:%
PET
!001
uncorrected
*loc ;emantic fluenc' Rest Nouns,baseline
2 inf! -rontal sulcus& 2 ;M(& 2 and R
Precentral sulcus& 2 and R -rontal
operculum& 2 sup! -rontal sulcus& 2
ant! +ingulate g'rus& R sup! Temporal
sulcus& 2 post! Temporal lobe& 2
Temporo,Parietal .unction& 2 +audate
nucleus& 2 and R Thalamus
6=
3erb,baseline
2 -rontal 5perculum& 2 inf! -rontal
g'rus& R Insula& R ;M(& 2 Precentral
sulcus& R ant! +ingulate g'rus& R sup!
Temporal sulcus& 2 post! Temporal
lobe& 2 post! Inf! Parietal lobe& 2 and
R +audate nucleus& 2 and R Thalamus
67

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Nouns and verbs in the brain

A PLACE FOR NOUNS AND A PLACE FOR VERBS?

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