neurobiology of emotional experience.pdf

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JOU RN AL O F N EU RO PSYCH IATRY 439

The N eurob io logy o fEm otiona l E xp en en ceK enneth M . H eilm an , M .D .

T he au thor d iscusses a neuro lo g ica l m ode l o f am odu la r ne tw ork tha t m ed ia te s em o tiona l experi-ence . A ccord in g to th is theo ry , em o tiona l exp eri-ence h as th ree com pon en ts: va lenc e (po sitive an dn ega tive ), a ro usa l, and m o tor a c tiva tio n (ap proa ch ,a vo id , ne ither). In th is m od e l, the cortex is c r itica lin reg u la tin g ac tiv itie s o f th e lim b ic system , b asa lg ang lia , an d re ticu la r system . T he fron ta l lob es areim po rta n t fo r va lenc e: the le ft m ed ia tes positiveem o tions , the rig h t neg a tive em o tio ns . T he rig h them isph ere , espec ia lly th e p arie ta l lob e , is im po r-tan t in a c tiva ting aro usa l sy s tem s, an d the le fthem isphere m odu la tes inh ib ition o f these system s.Th e righ t hem isphe re is a lso critica l in m o tor ac ti-va tio n . T he fron ta l lob es , espe c ia lly the orb ito fron -ta l p ortio ns , m ed ia te avo ida nce beha v io rs , a nd theparie ta l lob es m ed ia te appro ach b ehav io rs . T he corti-ca l a reas d iscussed have rich in terco nne c tion s andare a lso c lo se ly con nec ted w ith the lim b ic system ,b asa l gang /ia , an d re ticu la r system s. Em o tiona le xperience m a y d epend on th e p a ttern s o f neura lac tiva tio n in th is m odu la r ne tw ork .

(T he Journa l o f N europsy ch ia try and C lin ica lN euro sc ien ces 1997 ; 9 :439 -4 48)

A lthou gh alm ost eve ry one know s w ha t em otion sa re, the w o rd emo t i on is a psy ch o log ical con stru ct

tha t rem a ins d iff icu lt to d efine . P e rhaps em o tions ared ifficu lt to de fin e b ecause they a re no t ob jec ts o r th ingsb u t rath er a re com plex , m ulticom pon en tial p rocesse s.

Th ere a re a t lea st th ree m a jo r com ponen ts o f em otion .F irs t, the re is em o tiona l behav io r, inc lud ing ove rt be -hav io r an d chan ges in the v iscera and au ton om ic ne rv -ou s sy stem . S econd , th ere is em o tio na l com m un ica tionth roug h w ord , ton e of vo ice (p roso dy) , and gestu re s ,inc lu d ing fac ia l ex p re ss ions. Th ird , the re is em otion alfee ling o r expe rien ce . Em otion al expe rien ce, a ltho ughsub jec tive , is o ne of th e m a jo r fac to rs in m otiva tinghum an b eh av io r . It is no t on ly th e d irec t exp erience o fem otion th at m otiva te s behav io r , b u t also the k now l-ed ge tha t ce rta in stim u li, s itu a tio ns, and actio ns cou ldin th e fu tu re rep rod uce em otion al sta te s .

In th is rev iew , I exp lo re th e neu ra l basis o f em o tiona lex pe rience . B u ild ing m ode ls o f the neura l basis o f em o-tiona l exp e rience m ay he lp u s un de rs tan d ab norm a lem otion al state s . A lthou gh stud ie s based on the ab la-tion pa rad igm w ere used to he lp fo rm th is m od el, I dono t d iscu ss he re th e psy ch iatr ic a ffec tive d iso rde rs suchas pan ic and depress ion . I f irs t b r ie fly rev iew th e cla ssicfeedback and cen tra l theo ries and th e m ore recen t rev i-s io ns o f these theo rie s , an d I p ro v ide ev idence tha t the seth eo rie s cann o t fu lly exp la in th e n eura l b as is o f em o-tio na l exp erience. I th en d iscuss a d istr ibu ted m odu la r

From the D epa rtm en t o f N euro log y , U n ive rsi ty o f F lo rid a C ollege o fM ed icine, a nd N euro log y S erv ice , D epar tm ent of V e terans A ffair sM ed ica l C en ter , G ain esv ille , F lorida. A dd res s corre spo nd ence to D r.H eilm an , Box 100236 , U n ive rsi ty o f F lo rid a, G ain esv ille , FL 32610 -02 36 .


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NEUROBIOLOGY OF EMOTIONAL EXPER IENCE

44 0 VO LU M E 9. N UM BER 3 #{ 149}U M M ER 1997

ne tw ork theo ry of em o tiona l exp erience. T he pro -po sed m ode l su ggests tha t em otion al expe rien ce in -vo lves th e com pon en ts o f va lence, a rousa l, and m o to ract ivat ion .

FE ED BA CK TH EO R IES

Fac ial F eedb ack H ypo thesisC ha rle s D arw in2 no ted tha t h e w ho g ives v io len t ges-tu re inc rea se s h is rage . D arw in a lso b elieved th at them eans by w hich w e expre ss em otio ns are in na te. Iza rd3an d Ekm an e t a l.4 pe rfo rm ed c ro ss-cu ltu ral s tu d ie s o ffac ia l em o tiona l ex pre ss ion and fo und tha t the sam eseven to n ine em otion al fac ia l exp re ssio ns app eared tobe u n iv ersa l, the reb y p rov id ing sup port fo r D arw in shyp o thesis tha t em otion a l ex pres sion m ay be inn ate .Tom kin s56 pos ited tha t it w as the feedback of th esefac ial em otio ns to the b ra in th a t indu ced em otion alfee ling . L a ird 7 expe rim en ta lly m an ip u la ted facia l ex -p res sions and fou nd tha t p atien ts fe lt em otio ns, th erebyprov id ing som e support fo r th e facia l feedback hy-po thesis .

T he re are , h ow ev e r, m any unreso lv ed prob lem s w iththe fac ia l feedback theo ry o f em otio na l ex pe rience . F orexam ple , w hen one is fee ling a strong em o tion , thev alence o f th is em o tion canno t be chang ed by a lte rin go ne s facia l exp re ssion , an d a pe rso n can exp re ss o neem otio n w hile fee ling ano the r. I t is p ossib le th at vo lun -ta ry facia l em otio ns cann o t en tirely con tro l em otio na lexp erience because th e inn erva to ry pa tterns o f vo lun -ta ry facia l g estu re s a re d ifferen t from tho se th at occu rn atu ra lly . P a tien ts w ith pseud obu lba r pa lsy m ay ex-p re ss stro ng fac ia l em otio ns tha t they a re no t fee ling ,8 9b u t it is po ssib le th at the se pa tien ts b rain le sio ns a lsoin terru p t feedb ack to th e b ra in . P atien ts w ith d isea se so f th e fac ial ne rve m oto r u n it, even w hen th e p rob lemis b ila tera l, d o no t report a chang e of em otio n . A lthou ghit is po ssib le , a s D arw in sug gested , tha t facia l exp re s-sio ns m ay em be llish em otion s, it rem ain s p ossib le tha tth is in flu en ce is n o t from feedback from facia l m usc le s ,bu t rath er is m ed ia ted to ta lly w ith in the cen tral ne rvou ssy stem by asso c ia tiv e ne tw orks.

V iscera l F eed ba ck H ypo the sisW illiam Jam es# {1 76} p ro posed tha t s tim u li th at p rov okeem o tion indu ce chang es in the v isce ra an d tha t th ese lf-p ercep tion of th ese v isce ra l chang es is w h a t p ro -d uces em o tiona l exp erience.

Th is feedback th eo ry w as ch alleng ed by C annon ,1w ho a rgued tha t th e sepa ra tion of the v isce ra from thebra in , a s o ccu rs w ith cerv ica l sp in a l co rd in ju r ie s , doesn o t e lim ina te em otio na l expe rien ce . H ow eve r , the C an-

no n a rg um en t w as no t fu lly supp orted b y H ohm ann ,2w ho s tud ied pa tien ts w ith sp ina l co rd in ju r ie s an dfound tha t pa tien ts w ith e ithe r h igh o r low sp in a l co rdtran sectio n d id exp erience em otion s as p red ic ted byC annon , bu t tha t p a tien ts w ith low er le sion s repo rtedstron ge r em o tions th an those w ith h ig he r le sio ns.H igh er ce rv ica l lesio ns w ou ld be m o re like ly to affec te ffe ren t con tro l o f the v isce ra an d in ju re the au to nom icne rvou s sy stem s e ffe ren t ou tp u t. T he re fo re, H oh-m anns o bse rva tions p rov id e pa rtial su ppo rt fo r thev iscera l feedb ack th eo ry . I t sh ou ld also be no ted tha t thev ag us ne rv e is the p rim ary v isce ra l a ffe ren t n erve , andce rv ica l sp ina l co rd transec tion w ou ld no t in te rfere w ithth is ne rve s ab ility to p ro v ide the b ra in w ith feed backfrom the v iscera . H ow eve r, even if feed back w as po ssi-b le , w ith a los s o f au to nom ic con tro l th e re m ig h t b e littleto feed b ack .

C annon tho ugh t tha t th e v isce ra h ave insu ff ic ien ta ffe ren t inpu t to th e b ra in to b e im portan t in indu cin gem o tiona l exp erience. A s m en tio ned ab ove , how eve r,the v agus n e rve is m a in ly a v isce ral a ffe ren t ne rv e. Inadd ition , using a hea rtbea t d etec tion p arad igm , K a tk ine t a l.1 3 no t o n ly fo un d tha t som e no rm al su b jects canaccu ra te ly de tec t the ir h ea rtbeats , bu t a lso tha t the su b-jects w ho h ad th e strong est em otio na l re spo nses ton eg a tiv e s lide s w ere the su b jects w ho w ere ab le to de -tec t th eir ow n heartbea ts .14

C annon a lso th ou gh t th at becau se th e sam e v isce ra lre spon ses occu r w ith d iffe ren t em o tions, feedback ofth ese v isce ral resp onses cou ld n o t accoun t fo r th e va ri-ety o f em otion s th at h um ans expe rien ce . H ow eve r , A x5d em ons tra ted th at d ifferen t b od ily reactio ns can b e as-soc iated w ith d iffe ren t em otio ns.

T hus , C annons argum en ts fail to refu te the v isce ra lfeed back theo ry of em otio na l expe rien ce . H ow eve r, w ehave no t y et d iscussed h ow th is feed back system m aywo rk . In hum ans, the neocortex and lim b ic co rtex p laya c ritic a l ro le in th e an a lysis and in te rp re ta tion of v ari-ous s tim u li (see H eilm an e t a l.16 fo r rev iew ). B ecause thev isce ra re sp ond to com plex stim u li (such as speech ),the re m ust be effe ren t d escend ing n eu ra l system s tha ten ab le th e co rtex to con tro l the v isce ra v ia the au to -nom ic n e rvo us system and the end oc rine system .

T he au to nom ic n e rvo us sys tem has tw o com pon en ts ,the sym pa the tic an d the pa ra sym pa the tic . T h e descend -ing sym pa the tic neu rons rece ive p ro jec tio ns from thehypo tha lam u s, an d the hypo tha lam u s rece ives p ro jec -tions from m any lim b ic an d p ara lim b ic a rea s, inc lud ingthe am ygda la . The m o st im po rtan t pa rasym path eticne rve is the v ag us. T he vagu s o rig ina te s in the do rsa lm oto r n uc leu s situ ated in th e b ra ins tem and p ro jec ts tothe v isce ra , inc lu d ing the hea rt. T h e am ygda la no t on lyp ro jects to the hypo tha lam us , bu t also send s d irec t p ro -


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HE ILMAN

JO URNA L OF NEUROPSY CH IA TRY 44 1

je c tion s to the nuc leus o f the so lita ry trac t an d the d orsa lm o to r n uc leu s o f the vagu s. In th is m anne r theam ygda la m ay d irec tly in fluence th e p ara sym pa the ticsys tem . T he am ygda la rece ives neoco rtica l inp u t.

A ltho ugh th e am ygda la m ay be the m o st im portan tpa rt o f th e lim b ic sys tem to in fluence the au to nom icne rvou s sys tem and v isce ra, s tim u latio n o f o th e r a rea s,inc lud ing the in su la and orb ito fro n ta l co rtex , can a lsoind uce au tonom ic an d v iscera l ch an ges, and thesestruc tu res a lso rece ive in pu t from the n eo co rtex . A sm en tio ned , in rega rd to feedback the m ajo r n e rve tha tcarrie s v isce ral affe ren t in fo rm atio n back to the b rain isthe v agus. These v aga l a ffe ren ts te rm in ate p rim arily inthe nu cleus o f the so lita ry trac t. T h is n uc leu s p ro jec ts tothe cen tra l nuc leus o f th e am ygda la , and the cen tra lnu cleus o f the am ygda la p ro jec ts to o the r am ygda lanuc le i an d th e in su la . Th e am ygda la an d insu la, in tu rn ,p ro ject to seve ra l n eo co rtica l areas , inc lud ing th e tem -pora l, pa r ieta l, an d fron ta l lob es .

L uria an d S im ern itsk ay a 7 tho ugh t tha t the r igh them isp he re m igh t be m ore im portan t than the le ft inpe rce iv ing v is ce ral chan ges. T o tes t th is po stu la te ,D av id son e t a l.1 8 g av e a v arie ty o f tapp in g te sts to nor-m a l ind iv idu als . T h ey found tha t the le f t hand w as m o rein f luenced by hea rt rate than the righ t h an d , su gg estin gthe righ t hem isphe re m ay be supe rio r at de tec tingheartbea t. U nfo rtun ate ly , th is le ft h and su pe rio rity inthe de tectio n o f hea rtb ea t has no t been rep lica ted byo th er in ve stig ato rs .

T he re are o the r o bse rva tions fo r w h ich the v iscera lfeedback theo ry cann o t accoun t. F o r ex am ple , M a-ra#{241 }on19in jec ted ep ineph rine in to sub jec ts and then in -qu ired as to th e na tu re o f the em otio n fe lt by thesesub jec ts . E p in ephrine do es no t c ross the b loo d-b rainba rrie r, b u t it affec ts th e au ton om ic n e rvo us sy stem andv isce ra . H ow eve r , M aran on fo und tha t in jec tions o fep ineph rine w ere no t assoc iated w ith g en u ine em o-tio na l expe rien ces, bu t rath er w ith as if feelin gs.

S chacte r and S inge r2#{176 }a lso in jec ted ep in ep hrine in toex pe rim en tal sub jec ts and rep orted tha t p ha rm aco log i-ca lly in duced au tonom ic and v isce ra l ac tiva tion d id no tp rodu ce an em o tion un les s th is arousal w as accom pa-n ied by a cogn itive set. A lth ough S ch ac te r and S inge rsstud y sug gested th at v isce ra l feedb ack com bined w ithcen tra lly m ed ia ted co gn itio n is im portan t fo r em otio na lex pe rience , ob se rva tion s in o ur lab o rato ry d id no t en -tire ly su ppo rt the se f ind in gs.

R ecen tly , w e te sted the au to nom ic -v is ce ral feedb ackth eo ry to lea rn if the r igh t h em isph ere p lay s a dom inan tro le in pe rce iv in g v isce ra l ch anges. U sing a sho ck an tic i-pa tion p arad igm , w e fo und tha t w hen com pared w ithn orm a l con tro l su b jec ts , p atien ts w ith rig h t hem isph ereles ions had a redu ced au to nom ic re spo nse . A lth ough

the ir au ton om ic re sp onse w as red uced , they show ed nod ifferences in the ex pe rience o f an tic ipa to ry anx iety .2 1

In add ition , the strong est a rg um en t ag ain st the co g-n itive -v isce ra l feedb ack theo ry o r a ttr ibu tio n theo rya rise s from the observa tion s o f pa tien ts w ho havestron g em o tions (su ch as fea r) a sso cia ted w ith m ed ia ltem pora l lob e or am ygd ala se izu re s. In m any of theseep ilep tic pa tien ts , the em otio na l exp erience is o ften th efirs t sym ptom . Th e re fo re , in these p atien ts co gn itiv e setand au tonom ic /v isce ra l chang es com e afte r the ex pe ri-ence ra the r than be fo re th e expe rien ce . T he attr ibu tio ntheo ry o f S ch acte r and S inge r canno t accou n t fo r the seobservat ions.

C ENTRA L TH EO R IES

T o accoun t fo r em otion al ex pe rience , C annonprop osed tha t a ffe ren t stim u li en ter the b ra in an d a retran sm itted to the th a lam us, w h ich ac tiva te s th e hy po-tha lam u s. Th e h ypo tha lam us can in tu rn ac tiva te theend oc rin e system and the au ton om ic n ervo us sys tem ,and it is the se sy stem s tha t indu ce th e phy sio log ica lchang es in th e v isce ra . A ccord ing to C annon , au to -nom ic and v isce ra l ac tiva tion a re p rim arily ad ap tiveand a id in th e su rv iv a l o f the o rgan ism . Em otion al ex -pe rien ce, acco rd ing to C annon , is indu ced by the hy po-tha lam us feed ing back to the co rtex . M ore recen tly ,L eD oux e t a l.22 h ave m od ified C annons tha lam ic -h y -po tha lam ic em otion circu it to in clu de the am ygda la inf ea r c on di ti on in g.

C annon and L eD oux d id no t p ro pose a c ritic a l ro le fo rthe co rtex in th e in te rp re ta tion of stim u li. A lth oughsom e cond ition ed stim u li, s im ila r to th ose u sed by Le-D oux , m ay ind uce em otio n w itho u t co rtical in terp re ta-tion , the re is ov erw he lm ing ev id en ce th at in h um ansthe n eocortex is c ritic a l fo r in te rp re ting the m ean in g ofm any stim u li, e spec ially stim u li tha t a re com plex andre ly on past learn ing . F or exam ple , sp oken and w rittenstim u li m ay eng en de r stro ng em o tions . T hese stim u lineed phono log ica l, o r tho grap h ic , lex ica l, and sem an ticp rocessing th at is m ed iated p rim arily b y the le ft ce reb ra lco rtex .

R ecen tly , R oss et a l.2 3 p osited tha t the rig h t hem i-sph ere is im po rtan t fo r p rim ary em otion s (such as ange ro r fear) an d the lef t is im po rtan t fo r so cia l em otio ns(su ch as em barra ssm en t) . T hey b ased th is theo ry o n theob serva tion tha t som e pa tien ts u nde rgo in g se lec tivehem isp he ric anesth es ia ( the W ada te st) chan ged th eirem otio na l resp onse to traum a tic even ts tha t had oc -cu rred ea rlie r in the ir live s. F o r exam p le, o ne of th eirsub jec ts to ld o f an inc iden t w he re h e w as ve ry fr igh t-ened , b u t w hen h is r igh t h em isphe re w as anesthe tized ,


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NEUROBIOLOGY OF EM OTIONAL EXPER IENCE

44 2 V OLUM E 9 #{ 149}U M BER 3 #{ 149}U M M ER 1997

he sta ted tha t he fe lt em barra ssed . H ow eve r , R oss andco-w o rke rs d id no t dem onstrate tha t le ft hem isp he redy sfu nc tion led to a loss o f soc ia l em o tions . In th eabsen ce of su ch ev idence , th ere a re a lte rn ativ e exp lana -tions o f R o ss and co lleag ues o bse rv atio ns, and thesew ill be d iscussed b elow .

C annons d iencepha lic -h ypo tha lam ic m ode l, L e -D ouxs d iencep ha lic- lim b ic (am ygd ala ) m ode l, an dR oss and co lleag ues le ft- rig h t soc ial em o tiona l theo rie sn o t o n ly fa il to accoun t fo r h ow hum ans exp erienceem o tion in re sp onse to com plex stim u li, bu t a lso fail toexp lain how hum ans can exp erience a va rie ty o f em o -t ions.

M OD U LA R THEO RY

There are tw o w ays in w h ich th e b rain m ay m odu late av arie ty o f em otion al expe rien ces.

O ne poss ib ility is th at the b rain m ay con tain spec ial-ized em otio na l system s (devo ted system s) , so th at the rew ou ld be a specia l system for fea r, o ne fo r ange r , onefo r h ap p iness , an d so fo rth . Th is w ou ld im ply tha t eachem o tion w as un iq ue ly m ed ia ted . A second possib ility isth at each em otion is no t un ique ly m ed ia ted -tha t then eu ra l ap pa ra tus th at m ed ia tes on e em o tio n m ay no to n ly p lay a ro le in o the r em otion s, bu t m ay a lso m ed ia teeven non em otion a l func tion s (no nd ev o ted system s).

Th e nondevo ted sys tem s v iew is cons is ten t w ith th ed im ension al v iew o f em otio n favo red b y m any cogn i-tive em o tiona l theo ris ts . W und t24 p ro posed th at em o -tion a l expe rien ces v ary in th ree d im ensio ns: q ua lity ,ac tiv ity , and exc item en t (arou sal) . S im ilar ly , O sgood eta l.25 pe rfo rm ed fac to r ana lyses o n ve rba l a sse ssm en ts o fem o tiona l jud gm en ts and fou nd th at the va riance cou ldb e acco un ted fo r by th ree m ajo r d im ension s: v alen ce(pos itive /nega tive , p lea san t/u np lea san t) , a ro usa l(ca lm /ex cited ) , and con tro l o r dom inance (in con tro l/o u to f con tro l). U sing th is typ e o f m ultid im en siona l scale ,o ne can d e fine the d ifferen t em otion s. F o r ex am ple , fearw ou ld be u np lea san t, h igh a rou sa l, and ou t o f con tro l;sadn ess co u ld be unp lea san t and low arou sa l. F rijd a a lso exp lo red the co gn itiv e struc tu re o f em otion andfound th a t ac tion read iness w as an im portan t com po-n en t. S tu d ie s u sing a va rie ty o f techn iq ues w ith norm a lsu b jects27 h av e sup ported th is d im ens io na l v iew .

Th e m odu la r theo ry po sits tha t the co nsc ious ex pe ri-ence o f em otion is m ed ia ted by ana tom ica lly d istrib u tedm odu lar ne tw o rk s. B y modu lar , w e m ean th at a systemw orks in dependen tly . H ow ev er , th is does no t m eanth at m odu le s a re en capsu la ted in such a w ay th at theycann o t be in flu en ced by o the r sy stem s. B y ne twork , w em ean the sum to tal o f m odu les th at a re cr itic al in p red i-

catin g a behav io r o r expe rien ce . If the v io lin is t is them odu le , th e o rchestra is the n etw ork . Th e propo sedne tw ork co n ta ins th ree m ajo r m odu le s, one th at d ete r-m ines va lence (p leasan t ve rsus u np lea san t), an o the rtha t con tro ls a ro usa l, and a th ird th at m ed ia te s m oto ractiv atio n w ith approach o r avo id an ce b eh av io rs .

Va lenceG oldste in2 8 rep orted tha t ap hasic p atien ts w ith le fthem isp he re les io ns o ften ap peared anx ious, ag ita ted ,and sad . H e ca lled th is beh av io ral sy ndrom e th e cata -stroph ic reactio n . G a in o tti2# {176} stud ied 160 pa tien ts w ithrig h t o r lef t hem isp he re s tro kes. L ike G o ldste in , he re -po rted tha t p atien ts w ith lef t hem isp he re le sions h ad acata stroph ic reac tion , b u t he thou gh t th is w as a n orm a lresp on se to se rio us cog n itive an d phys ica l de f icits suchas aphasia o r h em ipa re sis . In co n trast, B ab insk i,3 #{1 76}H #{233} cean e t al.,31 an d D enny-B row n e t a l.3 2 n o ticed th a tpa tien ts w ith r igh t hem isp he re les ions o ften ap pearede ithe r in approp ria tely in d iffe ren t o r ev en eupho ric .G ain o tti29 con firm ed these ob serva tion s bu t su ggestedtha t the se pa tien ts ind iffe rence m igh t be re la ted to theden ia l o r u naw aren ess o f illne ss (ano sognosia ) tha to f ten is as soc ia ted w ith r igh t h em isph ere les ions.

A ltho ugh G a ino tti p ro posed th at pa tien ts psych o-log ica l respon ses to the ir ow n illne ss m ay accoun t fo rsom e o f th e em otion al a sym m etrie s ob se rv ed be tw eenpa tien ts w ith rig h t and lef t hem isp he re les ions, the rea re add itio na l repo rts tha t a psy ch o log ical reac tive th e -o ry can no t en tire ly exp la in . T e rz ian33 an d R o ssi andR osad in i34 stu d ied the em otion al reac tion s o f p atien tsrecov ering from se lec tive hem isp he ric ba rb itu rate - in -du ced anesthesia ( the W ada test) . T hese inv es tiga to rsno ted th at w he reas righ t ca ro tid in jec tions w ere o ftenassoc ia ted w ith euphoria, b arb itu ra te in jectio ns in to thele ft ca ro tid a rte ry w ere o ften assoc ia ted w ith ca ta -stroph ic reac tions. Th e W ad a te st is a d iag nostic tes t tha tcauses on ly tran sien t hem ipa resis and aphasia . I t th ere -fo re seem s un like ly tha t th is p rocedu re w ou ld cause areac tive dep re ssio n tha t o ccu rred d urin g on ly a pa rt o fthe te st. In ad d ition , w e have seen righ t hem isph ere -dam aged stroke p atien ts w ho app ea r to be em otion allyind iffe ren t b u t d o no t d em ons tra te ano sognosia o r ve r-ba lly ex p lic it d en ia l o f illne ss .

T he ca ta stroph ic-dep re ssive reac tion asso cia ted w ithle ft hem isph e re le sion s is seen m os t com m on ly in p a-tien ts w ho h av e an ter io r (fron tal) pe r isy lv ian le sions .3 5 3 6It is p ossib le th at the hem isphe ric em otio na l a symm e-trie s repo rted by G ain o tti an d o th ers m ay be re lated toa comm unica tion d iso rde r ra the r than d ifferences inem otio na l ex pe rience . Tu ck er e t a l.37 an d R o ss an d M e-su lam rep orted tha t righ t hem isp he re -d am aged sub-jec ts lo st the ir ab ility to exp re ss em o tiona l p ro sod y .


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Pa tien ts w ith lef t hem isp he re lesio ns m ay be ap hasicand un ab le to expres s th em se lves u sing propo sition alsp eech . T o expre ss fee lings , they m ay re ly m ore o n righ th em isph ere -m ed iated , n onp ro pos itiona l a ffectiv e com -m unica tion , and the re fo re these pa tien ts m ay in to nethe ir sp eech m ore heav ily and u se m ore em o tiona l fac ia lexp re ssion s. B ecause these le ft h em isph ere -dam ag edpa tien ts m ay have a grea te r need to use these exp re ssiv eem o tiona l system s , th ey m ay app ea r to b e h igh ly em o-tion a l. In con tra st, pa tien ts w ith righ t h em isph ere le -sio ns m ay have m ore d ifficu lty than pa tien ts w ith le f them isphe re d isease in exp re ssin g em otio na l faces andem o tiona l speech prosody . T he refo re, w h en com paredw ith le f t hem isp he re -dam aged pa tien ts , those w ithr igh t hem isp he re d am ag e m ay appea r to be ind ifferen t.

A lth ough hem isph e ric de fec ts o f em otio na l exp re s-sion m ay accoun t fo r som e o f the behav io ra l observa -tion s b y G olds tein ,2 8 B ab insk i,3 #{1 76} and G a ino tti,29 th eycanno t exp la in the re su lts o f G aspa rr in i e t a l.,39 w hoadm in iste red the M inneso ta M u ltiphasic P e rson ality In -ven to ry (M M PI) to a g ro up of lef t and rig h t hem i-sph ere -d am ag ed pa tien ts . T he M M PI is w ide ly used toasse ss em otio na l ex pe rience . T he lef t h em isph ere -d am -aged p atien ts w ere no t s ev ere ly aph asic . In add ition , ther igh t an d le ft hem isp he re pa tien ts w ere ba lanced fo rcogn itive and m oto r de fec ts . T h e M M PI do es no t re -qu ire em otion ally in toned speech o r fac ial ex p ress ions.T he find ings o f G aspa rr in i e t al. sup ported the hem i-sph eric v alen ce hypo thesis . T h ey fo und tha t pa tien tsw ith le ft hem isp he re d isea se show ed a m arked eleva -tion of the depres sion sca le , w he reas pa tien ts w ith rig h them isp he re d is ea se d id no t. T h ere fo re , the righ t-le f td iffe ren ces in em o tiona l behav io r ob se rv ed b y G a ino ttiand o th ers can no t b e a ttrib u ted to em o tiona l ex pres sived iso rd ers o r to the sev erity o f the m oto r o r co gn itiv edefici t .

S ta rk ste in e t al.4# {176}lso s tud ied em otio na l chang es as -soc ia ted w ith strok e and a lso fo und tha t abou t one -th irdo f strok e p atien ts h ad a depre ssion . Th ey no t o n lyfo und tha t th e dep re ssion w as asso cia ted w ith b o th le ftfron ta l and lef t caud ate les io ns, b u t a lso tha t the clo se rto th e fron tal po le the les ion w as loca ted , th e m o resev e re w as the depres sion . M any of the pa tien ts w ithle ft hem isp he re le sio ns an d depress ion w ere also an x-iou s. In con tra st, pa tien ts w ith r igh t fron ta l lesio ns w ereo ften ind ifferen t o r even euphoric . H ow ev er , no t a llinv estig a to rs ag ree tha t a fte r s trok e the re is m ore d e -p re ss io n w ith lef t th an righ t h em isphe re le sions . H ouseet a l.4 an d M iln e r42 co u ld no t rep lica te the em o tiona la symm etrie s foun d in p rio r rep orts . In ad d ition , a s d is-cu ssed abov e, R oss e t a l.2 3 po sited th at th e le f t hem i-sphe re m ed ia te s soc ia l em o tio ns and th e righ them isph e re m ed ia tes all p rim ary em o tions . H ow ev er , it

is n o t clea r th at R oss e t a l. fu lly exp lo red th is rig h them isph e re p rim ary em o tion hypo th esis .

T o learn if the re a re d is cre te phy sio log ica l changesassoc iated w ith d ep re ssio n , s ev era l g rou ps o f inv es tig a -to rs h av e conduc ted fun ctio na l im ag ing stud ies o f pa -tien ts w ith p rim ary depre ssion . S ev era l o f the seinvestiga tio ns h av e no ted a d ecrease o f ac tiva tion in th elef t f ro n ta l lo be as w ell a s th e le f t c in gu la te gy rus .4 3 4 4H ow eve r , D rev ets and R a ich le4 5 found in creased ac tiv -ity in th e lef t p re fron tal co rtex , am ygda la , ba sa l gang lia ,an d th alam us.

D av idso n e t al.46 and T ucke r4 7 inv estiga ted the h em i-sphe ric v alence h ypo thesis by study ing n orm a l sub jec tsw ith elec tro phy sio lo g ical tech n iqu es. T he ir s tud ies co n-f irm ed the ab la tion stu d ie s , sug gesting th at the fro n ta lreg ion o f each hem isp he re m akes asym m etric co n trib u -tio ns to em otio na l ex pe rience : g rea te r rig h t than le f th em isph ere ac tiva tion is a sso cia ted w ith neg ativ em ood s, an d grea te r le f t th an righ t hem isp he re ac tiva -tio n is a sso c iated w ith p ositive m ood s.

U n fo rtun ate ly , it is n o t k now n how the rig h t an d le f th em isph ere s m ay in flu en ce em otio na l v alence . F ox andD av idson sug gest tha t le ft hem isp he re -m ed ia tedp ositiv e em otion s a re re la ted to ap proach and righ th em isph ere -m ed iated nega tive em otion s a re re lated toavo idance b eh av io rs . H ow eve r , recen tly in o ur labo ra -to ry w e stu d ied the re latio nsh ip o f em otion s an d ap-p roach -av o idan ce b ehav io r . W e fou nd tha t nega tiveem o tio ns can be assoc ia ted w ith bo th ap proach andavo idance behav io rs (G . P . C ruc ian e t a l.: D issoc ia tionof b eh av io ral actio n and em otion al va len ce in th e ex -p re ssion o f a ffec t. P re sen ted a t the ann ua l m ee ting ofthe Am erican A cadem y o f N eu ro lo gy , B oston , M A ,A pril, 19 97) . F o r exam ple , fear an d ang er bo th h av e aneg ativ e v a len ce , bu t fea r is a ssoc ia ted w ith avo idanceand an ge r is as soc ia ted w ith approach . In add ition , th isapp roach -avo id an ce m od el does n o t ex p la in how th etw o h em isph ere s are d iffe ren tly o rgan ized su ch th atthey m ake oppo site con tr ibu tio ns to m ood . T h is ap -p roach -av o idan ce theo ry d oes no t ex p la in h ow othe rem o tio ns are m ed iated , and it a lso does no t exp lain thero le o f o the r a rea s in the b rain , such as th e lim b ic sys tem .

Tuck er and W illiam son49 th ink tha t hem isp he ric va -lence asym m etrie s m ay b e re la ted to asymm etrica l con -tro l o f n eu ro pha rm aco log ica l system s , the le f them isp he re b ein g m ore ch o line rg ic and dop am ine rg icand the righ t hem isph e re m ore n orad rene rg ic . M aybe rge t a l. 5# {1 76 }ep orted tha t ph arm aco log ic chang es in the tw ohem isphe res m ay be d iffe ren t a fte r strok e. U s ing PETim ag in g , they rep orted th at w he reas strokes in the righ them isphe re app ear to in c rea se se ro ton e rg ic recep to rb ind in g , le ft hem isp he re strokes low er sero to ne rg icb in d ing . T he low er th e se ro ton erg ic b in d ing , th e m ore


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44 4 V OLU M E 9 . NU M BER 3 #{ 149}UM M ER 1997

seve re th e d ep re ssio n . A lth ough it is w ell know n fromc lin ical p sych ia try th at neu ro tran sm itte r sys tem s m ayhave a pro fo un d in fluence on m ood and tha t ch angesin these system s m ay in duce e ithe r eup horia o rdy spho ria , the m echan ism by w hich the p ha rm aco log i-cal ch an ges in duce m ood rem a ins u nknow n .

The fro n ta l lo bes also have stro ng p ro jec tion s to th eliinb ic sy stem . T he do rso late ral an d m ed ia l fro n ta l lobep ro jec t p rim arily to th e c ing u la te gy rus; the o rb ito fro n -ta l co rtex , v ia th e un cin ate fascicu lu s an d an te r io r tem -p ora l lobe , h as stro ng co nnec tions to th e am ygda la.A lth ough lim b ic struc tu res such as the am ygd ala areim po rtan t in nega tive em o tions su ch as fea r, them eans b y w h ich the rig h t and le ft fron ta l lob es se lec-tive ly in fluence these lim b ic stru c tu re s rem a in un -k n own .

A ro u sa lThe te rm arousa l , lik e the te rm s at tent ion an d emo t i on , isd iff icu lt to d efine . B ehav io ra lly , w he reas an a rousedorg an ism is aw ake , a ler t, and p repa red to p rocess stim -u li, an u na ro used organ ism is leth arg ic to com a tose andno t p repa red to p ro cess stim u li. A ro usa l h as seve ra lphy sio log ic de fin ition s. In th e cen tral n e rvo us sy stem ,arou sal usu ally re fe rs to the ex c itato ry sta te o f neu ro nso r th e p rop en sity o f n eu ro ns to d isch arge w h en ap pro -p ria te ly ac tiva ted . In func tio na l im ag in g , a rou sa l is usu -ally m easu red by inc rease s o f b lo od flow , an delec tro phy sio log ica lly it is m easu red by desyn chro n iza -lio n of th e EEG or by the am p litude and la ten cy o fev oked po ten tials . O u tside the cen tra l ne rv ou s sy stem ,arou sal u sua lly re fe rs to ac tiva tion o f the sym pa the ticne rv ous sy stem and the v isce ra, inc lu d ing the hea rt.

A rou sa l an d a tten tion a re in tim a te ly link ed an d ap-pear to be m ed iated b y a cortica l lim b ic re ticu lar m odu-lar n etw ork .51 Th is a rtic le p rov id es an o ve rv iew of th ism odu la r n etw ork , bu t fo r d eta ils on e sho u ld re fe r to theo rig ina l ar tic les . M uch of w ha t w e know abou t thean atom y of th is ne tw o rk in itially cam e from stud ie s o fm onkeys an d hum an p atien ts w ith d is cre te b rain le -sions . H ow ev er , m ore recen tly fun ctio na l im ag ing h asco nfirm ed m uch of th is ab la tion re search . In hum an s,lesio ns o f th e in fe r io r pa rieta l lo be are m ost o ften asso -cia ted w ith d iso rd ers o f a tten tion and a rousal,55 5 6 an din m onk ey s tem po rop arie tal ab la tio ns a re a lso asso ci-ated w ith atten tio na l d iso rd ers .57 P hy sio lo g ical re -co rd in gs from neu ron s in th e pa rieta l lob es o f m onkeysap pear to sup port the p ostu late tha t the pa rie ta l lo be isim po rtan t in a tten tion . U n like neurons in the p rim arysen so ry co rtex , th e se a tten tiona l neu ro ns in the pan -eta l lob e, a sup ram od al a sso c iatio n co rtex , show anassoc iatio n b etw een the ra te o f fir ing and th e sign ifi-can ce o f the stim u lus to the m onk ey-th e relev an t stim -

u li be ing asso cia ted w ith h igh e r fir ing ra te s than theun im portan t s tim u li. 5 9

S enso ry in fo rm a tion p ro jec ts to th e tha lam ic relaynuc le i. F rom th e tha lam us, th is m oda lity -spec ific sen -so ry system pro jects to th e p rim ary senso ry co rtice s .E ach o f these p rim ary senso ry co rtices (v isua l, ta ctile ,au d ito ry ) p ro jec ts o n ly to its a sso c iatio n co rtex . F orexam ple , B rodm ann area 17 , the p rim ary v isua l co rtex ,p ro jects to B rodm ann a rea 18 . S u bsequ en tly , the se m o-da lity -specif ic assoc iatio n a reas conv erge on po lym od ala rea s such as the fron ta l co rtex (pe ria rcu ate , p rea rcua te ,and o rb ito fron tal) and bo th b an ks o f the su pe rio r tem -po ra l su lcu s.6 #{176 }B oth of these senso ry p o lym oda l con ve r-gence areas p ro jec t to th e supram od al in fer io r p arie tallobe.6

A ltho ugh the de te rm ina tion of the nov elty o f a stim u -lus m ay be m ed ia ted by m oda lity -spec ific senso ry as so -c iatio n co rtex , de te rm ina tion o f the sign ificance o f astim u lus requ ires know led ge as to bo th th e m ean ing ofthe stim u lus and the m otiva tio na l s tate o f the o rgan ism .T he m o tiva tiona l sta te is depend en t on a t lea st tw ofac to rs: imm ed iate b io log ica l n eed s and lo ng-te rmgo als . It ha s been dem on strated tha t po rtio ns o f th elim b ic sy stem , tog e the r w ith th e hypo tha lam us , m on i-to r the in te rna l m ilieu an d d ev elo p d riv e sta te s . Th e re -fo re , lim b ic in pu t in to reg io ns im portan t in de term in ingstim u lus s ign if ican ce m ay p rov ide in fo rm atio n abou timm ed ia te b io log ica l n eeds . R eg a rd in g lo ng -te rm goa ls ,the fron ta l lobe has been dem onstra ted to p lay a m a jo rro le in go a l-o r ien ted behav io r an d set d eve lopm en t,6 2 63and fron tal in pu t in to the a tten tion al-a ro usa l system sm ay prov id e in fo rm atio n abou t g oa ls tha t are no t m o ti-v ated by imm ed iate b io log ical n eeds. S tud ies o f co rticalcon nectiv ity in m onkeys h av e d em ons tra ted tha t th etem po ro pa rie ta l reg ion no t on ly has stron g conn ec tion sw ith po rtio ns o f the lim b ic system (c ingu late g y ru s) b u ta lso w ith the fron ta l co rtex .

S tim u la tion o f the m esen cep ha lic re ticu la r fo rm atio n(M RF ) in an im als ind uces behav io ra l an d p hysio log ica la ro usa l.M In con tra st, b ila tera l les ions o f the M RF in ducecom a, an d un ila tera l le sions cause th e ipsila te ra l hem i-sph e re to b e b eh av io rally an d physio log ica lly hypo-a ro used .65 T he po lym oda l and sup ram oda l co rtexd is cu ssed abov e n o t on ly de te rm in e stim u lus sig n ifi-cance, bu t also m odu la te a rou sa l b y in fluenc ing th eM RFP T he exac t m eans b y w hich these co rtica l a rea sin f luence th e M RF and the M RF in flu en ces th e co rtexrem a in unknow n . H ow ev er , the re a re at least th reep ossib le m ech an ism s by w hich the M RF m ay in fluenceco rtica l p ro cess ing . F irs t, S hu te and L ew is67 desc ribe anascend in g ch o line rg ic re ticu la r fo rm a tion . Th e n uc leu sb asalis , w h ich is in th e b asal fo reb ra in , receiv es in pu tfrom th e reticu la r fo rm a tion and has cho line rg ic p ro jec -


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tion s to the en tire co rtex . Th ese ch o line rg ic p ro jec tio nsappea r to be im portan t fo r in creas ing n eu ro na l re sp on-siv ity .6 8 S econd , the M RF m ay also in f luence the co rtica lactiv ity th roug h th alam ic p ro jec tio ns. S ter iade andG len n69 d em ons tra ted tha t n onspecif ic th alam ic nu cle isu ch as cen tra lis late ralis and pa racen tralis p ro jec t tow idesp read co rtica l reg io ns an d these tha lam ic n uc leican b e ac tiv a ted by stim u latio n of th e m esen cep ha licreticu la r fo rm atio n . T he th ird m echan ism th a t m ay he lpacco un t fo r co rtical arou sa l in vo lv es th e th a lam ic nu-c leu s re ticu lar is (NR ). T h is th in nuc leus en ve lops thetha lam us and p ro jec ts to a ll the sen so ry tha lam ic relaynu cle i. P hy sio log ica lly , N R inh ib its th e tha lam ic re lay o fsenso ry in fo rm atio n .70 H ow ev er , w hen co rtica l lim b icne tw ork s d e term ine tha t a stim u lus is s ign if ican t o rnov e l, co rtico fuga l p ro jec tion s m ay inh ib it the in h ib i-to ry NR , the reb y allow in g th e tha lam ic sen so ry n uc leito re lay sen so ry in fo rm a tion to the co rtex .

T he lev e l o f activ ity o f th e pe rip he ra l au ton om ic ne rv -ous system u sua lly m irro rs the leve l o f arou sa l in th ecen tra l n ervo us sy stem . O ne m ean s o f m easu ring pe ri-phe ral au ton om ic a ro usa l is to m easu re hand sw ea tingby using the ga lvan ic sk in re sp onse . T o lea rn if the rew ere d ifferences in the hem isphe ric con tro l o f sw eatin g ,H eilm an e t al.71 stud ied p a tien ts w ith rig h t and lef them isp he re d am age as w e ll as n o rm a l co n tro l su b jects .T hese in vestiga to rs p re sen ted these sub jec ts w ith no -c icep tiv e stim u li (e lectr ic sho ck ) and dem on stra ted th a tpa tien ts w ith rig h t hem isphe re les ions h ad a red ucedarou sal re spon se w h en com pared w ith n orm a l sub jec tsan d le ft h em isph ere -dam ag ed con tro l sub jec ts . S ub -seq uen tly , o the r inv es tig a to rs repo rted sim ila r f ind in gs.F o r ex am ple , M orrow e t al.72 an d S chran d t e t a l.73 a lsofou nd th a t r igh t hem isph e re -d am aged p atien ts h ad aredu ced sk in re spo nse to em otio na l stim u li. H ow eve r ,H e ilm an and h is co -w orke rs71 repo rted ano the r in te r-estin g find in g . W hen com pared w ith no rm a l sub jec ts ,pa tien ts w ith lef t h em isph ere le sion s ap pea r to have agrea te r au ton om ic respon se . U sin g changes in h ea rt ra teas a p eriphe ral m easu re o f a rou sa l, Y okoyam a e t a l.74ob ta ined resu lts s im ila r to tho se usin g sk in re spo nse .U sin g physio lo g ica l im ag ing , P e ran i e t a l.75 also fo undth at in case s o f r ig h t hem isphe re stroke th ere w as a lsom etabo lic dep re ssion of the le ft h em isph ere . U n fo rtu -n ate ly , le f t h em isph ere -d am ag ed co n tro l pa tien ts w eren ot re porte d.

T he m echan ism unde rly ing the asymm etrical h em i-sp he ric con tro l o f a rou sa l rem a in s unknow n . B ecau selesio ns re str ic ted to th e rig h t hem isphe re cou ld no td irec tly in te rfe re w ith the le ft h em isphe res co rtico fug alp ro jec tion s to the re ticu lar system s or the re ticu lar sys-tem s co rticip eta l in f luence of the lef t hem isp he re , o new ou ld hav e to p ro po se tha t th e rig h t hem isp he re s con -

tro l o f a ro usa l m ay be rela ted to p riv ileged comm unica -tio n tha t the rig h t hem isphe re has w ith th e re ticu la rac tiva ting sys tem . A lte rna tive ly , p o rtions o f the rig h them isph e re m ay p lay a dom inan t ro le in com putingstim u lu s sig n if icance . Th e inc rea sed a rousa l a ssoc ia tedw ith le ft hem isp he re le sions a lso rem a ins unexp lain ed .P e rh aps the lef t h em isph ere m a in ta ins som e type ofinh ib ito ry con tro l ove r the r igh t hem isphe re o r the re-ticu la r ac tiva ting sy stem .

M oto r A c tiva tion and A pproach -A vo idanceSom e em otion s d o no t ca ll fo r actio n (fo r exam ple , sad -n ess o r sa tisfac tion ), bu t o the rs do (an ge r, fear , joy ,su rp rise ) . W hen em o tions a re as soc iated w ith ac tion ,th is ac tion m ay b e tow ard th e stim u lus (ap proach ) o raw ay from the s tim u lus (avo id an ce ) tha t ind uces theem o tion . A ltho ugh one w ou ld like to av o id em o tionstha t a re un p lea san t and ap pro ach situa tion s th at in ducep lea san t em o tions, w hen w e d iscu ss ap proach andavo idance w e a re d iscuss ing th e b eh av io r a sso cia tedw ith the em otio n and no t o ne s p lans fo r s truc tu r ingo ne s behav io r in re la tion to on es env ironm en t. F o rexam ple , w he reas on e w ou ld like to av o id situ atio nstha t indu ce ange r , w h en on e d oes becom e angry , o nehas a p rop en sity to ap proach th e stim u lu s th at is ind uc -ing th is em otio n . Joy , a positiv e em otio n , is a lso as soc i-a ted w ith app ro ach beh av io rs .

P rib ram and M cG uin es s76 u se th e term act ivat ion todeno te the ph ysio log ica l read iness to respond to stim -u li. W e hav e posited tha t m o to r activ atio n or in ten tio nis m ed ia ted by a m odu lar ne tw o rk th at in clu des p or-tions o f the ce reb ra l co rtex , b asal g an g lia , and lim b icsystem (see H e ilm an e t a l.54 fo r a de tailed rev iew ). Th edorso late ral fro n ta l lobe appea rs to be the hub o f th ism oto r p rep ara to ry n etw ork .52 P hysio log ica l reco rd in gfrom cells in the dorso la te ra l fron ta l lo be revea ls neu -ro ns tha t have enh an ced ac tiv ity w h en the an im al isp resen ted w ith a stim u lus tha t is m ean ing fu l and pre-d ic ts m ov em en t.7 8 Th e dorso la te ra l fro n ta l lo bes rece iveinp u ts from th e c ingu late gy ru s and from th e pos ter io rco rtica l a ssoc ia tion a reas tha t a re m od ality -sp ecif ic ,po lym od a l, and supram od al. Inpu ts from th ese poste -rio r neocortica l a rea s m ay p ro v ide th e fron tal lob es w ithin fo rm a tion abou t th e stim u lu s, nc lud ing its m ean ingand its spa tial loca tion . Th e lim b ic sy stem (fo r exam p le ,th e cin gu la te gy rus , w hich is no t o n ly pa rt o f th e P apezc ircu it b u t also rece ives inpu t from Y ako lo v s basa l lat-e ra l circu it) m ay prov id e in fo rm a tion as to the o rg an -ism s m o tiva tiona l sta te . T he d orso la tera l fro n ta l lob ehas n onrec ip roca l con nectio ns w ith th e basa l gang lia(fo r exam ple , w ith th e cau da te) , w h ich in tu rn p ro jec tsto th e g lobu s pa llidus, w h ich pro jects to the tha lam us ,w h ich pro jec ts b ack to the fron tal co rtex .7 9 T he dorso la t-


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e ra l fron tal lobes also have ex tensive con nectio ns w iththe non spec ific in tralam in ar nuc le i o f the tha lam us(cen trom ed ian and pa rafa sc icu lar is) . T hese in tra lam i-na r nu cle i m ay ga te m o to r ac tiva tion b y th e ir in f luenceo n the basa l gang lia , e spec ially the pu tam en , o r byin fluenc in g the tha lam ic po rtion o f m o to r circu its (y en -tra lis la tera lis pa rs o ra lis) . L astly , the do rso la tera l fron tallo be has stron g in pu t in to th e p rem oto r a rea s. T he o b-se rva tion tha t le sio ns o f the d orso la te ra l fron tal lob e, thec ing u la te g y rus, the b asal gang lia , the in tra lam ina r n u -c le i, and th e ven tro la te ra l tha lam us m ay a ll cau se aid -n esia sup ports th e p ostu la te tha t th is system m edia tesm oto r activ atio n .

The righ t hem isphe re ap pea rs to p lay a sp ecia l ro le inm oto r activ atio n or in ten tio n . C osle tt and H e ilm an54d em onstra ted tha t r igh t hem isp he re lesio ns a re m orelike ly to b e asso cia ted w ith co n trala te ra l ak inesia thanth ose o f th e le ft hem isp he re . H ow es and B olle r8 m ea -su red reactio n tim es (a m easu re o f th e tim e taken toin itia te a re sp onse) o f the hand ipsila tera l to a hem i-sp he ric le sion and dem onstrated tha t r igh t hem isphe rele sion s w ere asso c iated w ith slow er reactio n tim es th anle ft hem isph e re le sions . H ow ev er , as p rev iou sly d is-cussed , th is f ind in g m ay be re lated to th e im portan t ro leo f the r ig h t h em isphe re in m ed ia ting a ro usa l. H e ilm anand V an D en A be1182 m easu red the red uc tion o f reactio ntim es o f norm a l sub jec ts w ho rece ived w arn ing s tim u lid irec ted to e ith er the ir rig h t o r lef t hem isp he re . Th eyfound tha t, ind ep en den t o f th e hand used , w arn ingstim u li de live red to th e righ t hem isp he re reduced reac -tion tim e to m id line stim u li m ore than d id w arn ingstim u li de live red to th e le f t h em isph ere .S om e em o tions a re a ssoc ia ted w ith approach beh av -io rs , o the rs w ith avo id an ce behav io rs . U nfo rtuna te ly ,th e portion s o f the b ra in th a t m ed iate app ro ach andavo idance b ehav io rs have no t been en tirely e luc id ated .D enny -B row n and C ham bers83 sugg ested th at the fron -ta l lobes m ed ia te avo idance behav io rs an d the pa rieta llo bes m ed ia te app ro ach behav io rs . Th ese au th o rs alsosu ggested tha t ap proach and av o idance behav io rs m ayb e recip ro ca l, so tha t a lo ss o f on e behav io r m ay re lea seth e o th er behav io r.83 Th e re fo re , becau se the fron tallo bes m ed ia te av o idance behav io r, fron ta l lo be le sionsw ou ld cause in ap propria te approach behav io rs , andb ecause the pa rieta l lob es m ed ia te ap proach b eh av io rs ,p ar ie tal le sio ns w ou ld in duce avo idance . In support o fth is p ostu la te , on e can see pa tien ts w ith fron tal le sion sw ho dem on strate a va rie ty o f ap proach b ehav io rs , in -c lud ing m anua l g ra sp re flexes, v isua l g ra sp re f lex es ,roo tin g an d su ck ing re spo nses, m agne tic ap rax ia, u tili-za tion behav io rs , and d efectiv e respon se inh ib ition . U n-fo rtuna te ly , w h ich specif ic area o r a rea s w ith in th efro n ta l lob es, w hen dam aged , cause ap proach beh av -

io rs h as no t been en tirely elu cid a ted . A n im als w ith fron -ta l le sion s show an inc rease o f agg re ssiv e b ehav io r .P a tien ts w ith le f t do rso la te ra l fron tal le sion s (w h ichsho u ld ind uce an em otio n of n eg a tive va lence, in -c rea sed a rousa l, and approach behav io rs) a re a lso p ron eto hos tility and ange r.8 4 P a tien ts w ith o rb ito fron ta l le -s ion s a re p ron e to irritab ility , an ge r, inapp rop ria te sex -ua l adv ances , and euphoria -jo cu lar ity ,85 and stud ie s o fan im a ls w ith v en tra l fron ta l lesio ns a lso dem onstrateabno rm al approach b eh av io rs . Th e o rb ito fron tal co rtexis clo se ly co nnec ted w ith a rea s o f th e lim b ic sy stem suchas the am ygda la tha t a re k now n to be im portan t inem otio na l ex pe rience .

D enny -B row n and C ham bers83 d em onstra ted tha t, incon tra st to th e m anua l g rasp re spon se as soc ia ted w ithfron tal le sion s, p atien ts w ith p an eta l les ions m ay dem -on stra te avo id ing respon ses. P a tien ts w ith p a rie ta l le-s ion s, e spec ia lly o f th e rig h t sid e, m ay no t on ly fa il tom ov e or sh ow a de lay in m ov in g the ir arm s, h eads , andeyes tow ard a pa rt o f th e sp ace tha t is o ppo site thepa rie ta l le sion , bu t th ese pa tien ts m ay even dev ia te the ireyes, h ead , an d a rm s tow ard ipsila te ra l hem isp ace . Inadd itio n , un lik e p atien ts w ith fron ta l le sio ns w ho can-no t w ithh o ld the ir re spon se to stim u li, p atien ts w ithpa rie ta l les ions m ay be un ab le to re spond to s tim u li(neg lec t) . T hese avo idance re sp onses are m ore seve rew ith r igh t than le ft hem isph e re les ions.

S U MMA R Y

In add ition to rev iew in g som e o f th e fo rm er feed backand cen tra l th eo rie s o f em o tio na l exp erience, I h av ed iscus sed a cen tral m odu lar th eo ry . A lth ough I a rg ueaga inst the p ostu la te tha t feedback is cr itic al to theexpe rien ce o f em otion s, I do suspec t tha t feedb ack m ayin fluence em otion s. Em o tio ns m ay b e cond ition ed andm ay use tha lam ic lim b ic c ircu its , a s p ro posed b y L e-D oux . H ow ev er , m ost em otion a l behav io rs an d exp e ri-ences are ind uced by com plex stim u li th at an iso latedtha lam us cou ld no t in te rp re t. T he ce reb ra l co rtex o fhum ans has com p lex m odu la r system s tha t ana lyzestim u li, deve lop p ercep ts , and in te rp re t m ean ing .

W e hav e propo sed tha t the expe rien ce o f em otion s isd im en siona l. A lm os t a ll p r im ary em otio ns can b e de -scr ibed w ith tw o or th ree facto rs , inc lud in g v a len ce ,a rou sa l, and m oto r ac tiva tion . T he de term in atio n ofva lence is based o n w he the r the s tim u lus is bene fic ia l(po sitive ) o r d e trim en ta l (nega tive ) to a p e rso n s w e ll-be ing . W hereas th e righ t fro n ta l lobe and its sub co rtica lconn ec tion s app ea r to b e im portan t in th e m ed iatio n ofem otion s w ith n ega tive v alence , th e le ft fron ta l lob e andits sub co rtica l con nec tions m ay be im po rtan t in the


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m edia tion of em otio ns w ith po sitive v alence . D ep end-ing on the na tu re o f the stim u lus, som e po sitive andsom e nega tive em o tions (such as joy and fea r) a re a sso -c iated w ith h igh arou sa l, and o the rs (such as satisfactio nand sadn es s) are a sso c ia ted w ith low aro usa l. W hereasth e righ t pa rie ta l lobe appea rs to be im po rtan t in m ed i-a tin g arou sal re sp onse, the le ft h em isph ere app ears toinh ib it th e arousal re spon se . S om e positiv e and nega -tive em otion s (such as an ge r, fear , and joy ) are a sso ci-a ted w ith m o to r ac tiva tion , an d o th e rs (such as sad ness)a re n o t. T he rig h t fron tal lob e ap pea rs to be im portan tin m oto r ac tiva tion . Th e m o to r ac tiva tion assoc ia tedw ith em otion s m ay b e eith er ap pro ach or av o idan cebehav io rs . A pproach behav io rs m ay b e m ed iated by the

Ref e r ence s

p arie tal lobes; av o idan ce beh av io rs m ay b e m ed ia ted b ythe fron ta l lo bes.

Th e cortica l a rea s w e have d iscussed hav e rich in te r-con nectio ns. In ad d ition , th ese n eo cortica l a rea s h av erich conn ec tion s w ith the lim b ic system , basal gang lia ,th alam u s, and re ticu lar system . T he re fo re, the ana tom icm odu les tha t m ed iate va lence , a rousal, and ac tiva tionsy stem s a re rich ly in terconn ec ted and fo rm a m odu la rn etw ork . Em otio na l exp e rience depend s on the p at-te rn s o f neu ra l ac tiva tion o f th is m odu la r ne tw ork .

This work w as su ppo rted b y the M ed ica l R esearch Serv ice o f th eD epartm en t o f V ete ra ns A ffa irs .

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d isp lays of em otion s. Sc ien ce 1969; 164 :86 -885 . T om kin s S S : A ffect, Im age ry , C on sciousne ss, vo l 1 : Th e Po sit ive

A ffec ts. N ew Yo rk , Spr ing er, 1 9626 . Tom kins S S : A ffe ct, Im ag ery , C on sciousnes s, vo l 2 : T he N egative

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n an ts o f em otiona l sta te . P sy cho l R ev 1962; 69 :379 -39 921 . S lom a ine B S : H em isph eric d iffere nce s in em otion al psych ophys io l-

og y . D oc toral d isserta tio n , U nivers ity o f F lo rid a, 199 522 . L eD oux JE , C icchet ti F , X agoraris A , e t a l: Th e la teral am ygd alo id

n uc leu s: s ensory in te rfa ce of th e am ygd ala in fe ar co ndition ing .N eu rosci 1 990 ; 10 :10 62-1069

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2 4 . W undt W : G ru ndr iss der P sycholog ie (O u tlin e of p sy cho log y].S tu ttga rt, E nge lm ann , 1 903

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27 . G re enw ald M K , C ook EW , Lang P J: A ffe ctiv e ju dgm en t and psych o-p hy sio log ical resp onse: d im ens ion al co-v aria tio n in th e evo lu t ion ofp icto ria l s tim uli . J Psy chophysio l 1 98 9; 3 :5 1-6 4

28 . G oldste in K : Langu ag e and L anguag e D istu rbance s. N ew Y ork ,G ru ne an d S tra tton , 19 48

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3 0. B ab in sk i J: Contr ibu t ion a l# {2 33} tu de de s tro ub les m entaux d anslhem ip leg ie organ iqu e c#{233} r#{233}braleano sog no sie) IC ontr ibu tio n to th es tud y of m enta l p ro b lem s in o rg an ic ce rebral h em ip leg ia (anosog-n os ia)] . R ev N eu ro l 1 914 ; 2 7 :8 45 -84 8

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