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    Neuroscience of Personality Cloniger and Others

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    Neuroscience of Personality Cloniger and Others

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    Chapter 10 The Neuroscience of Personality oS ‘THE SCIENTIFIC STATUS OF PERSONALITY RESEARCH Personaly copies « peclar postion within che diipines ‘of maccin-pjchology ‘The Pelee of ming in Seay nguage shows how Fequendy fe topic oP sonality crops up in conversation, andthe ples er of dosnyouselt personaly ivenobe Poplar mogaine confis the wel exashed place that personality ocupis a poplar tanks fog and care By Contrary tour perception «bac pessoal secich often seconded ow scientific sus by some experimental Prycholo fis and is somewhat uoderepeseted Ite Fanking jurrals Mach ofthe responsi be the sets quo mus lie th penalty eeacy iselt In sdaion to the inevable copie nature of personaly studies, te disspling foe seve other fetes that tend to eats dade anongst card-carrying expenmentit, Thee include the cena rl of abject lumens (Ga. selfrepor menue), the nator ten dency to cexsibe pesondity stucues rites than to acemp redactions explanations a the song links berween persnsliy and ane ing theory inthe infomation proceing es Ta thls chapter we wl oy to lanes tae eck Alan D. Pickering and Jeffrey A. Gray University of London tal developments in neuroscience over the last 15 yeas so may offer a window of opportu nity for petsonalry esearch to undergo a sien ‘fie makeover. Despite the above difculies, relating co the scenic suis of penalty sauch, Shee hhave been several theorist, such a8 Pavlov and Hi. J, Eysenck, who have arempted to under- stand personalty—and its psychobiological sub- strates—via the usual mel I sc fice Now, modern veins ofthese teas ace being given fresh impetus by our abilies to cary Sut funcional maging ade ote brain, to conduct investigations into the mo: lecular genetics of variztions in human neuro. transmission, and to interpret these findings in telaon to feu inspied compuratonal tmodels of the relevant processing operations ander rain systems. One gil of the prevent chapter is to indicate some of the ways one imighe ty to build a modern, integrated neuro science of personality Iv is abo iimpostant to point out that those who have advanced the neuropsychology of ‘human personality have been motivated by an additional desire to cas light on hurnan psycho. patology. The Fundamental astumption under lying the personality psychopathology linkage, 27 278 INTERFACE WITH OTHER FIELDS aad one that underpins our ova teseuch, is of "ormal personaly vaio yng orton war within da Phe ma a cus example thas of aay Hay eens 6 vay Ia tit ancy which scale degree fo which an ina is goatelh ate pod covard anvious cogtions Sad Sehr In this speci example, the continous ral ‘uppoes that cen nical watery duos e Found in individuals who cathe hgh ple of the “normal” at any dimension. es oo legion pd ache dc eg hops, Lykke, 1957) may ccc in ting, sho et heen ora ee ‘Armed with this continnum view of poche, ego tie a din canna er Sgaing pose pychobilopeal neces of ic porthoputologie trough the tenbed af sues ceporing raed pentralty tert sase ton within heathy suet or as, fee teson and Newman (1993) edrance tee the sadly of exraveson sa means of cucdio, ine Reelin of dang, i heve undepins poychopathy and sated di dex In addon fo obvious udantges of ise avaablity and compliance they ove da Fike behavior ofextraversthe lew dato by negative life events and comorbid tee abuse)... and), theiesoca dake bibion and impviy or spontaney tof estemed or sap. «In contr fopho path offen twoubled childhoods | PISO. ‘ody ther expression of te dacs, Mee caved colegestuens hove nov aap tnced the unvoward ef ofan sntnocal spl (p- 718) Claridge (1987 has mad gos atgumens forthe study of schnophvene ‘ici though she lens of scaonpal geese igre variation in hak bj Ith chaper we wll no aternp a compte hensive eview of cent neuoscnehe Bec and sheotes eonerning s tange of majos fe fotalty dimensions, Innead. e will Hostage the use of neuracence tw undetand penal iy beter by concentrating lngey on he repr cluster of personality 8 her ae be eet {as impulsive seaionseking (SS) In tha shaper we wil proposes Seucobicloged ‘motel of these eras by desing together dee ing kinds of sewexcnte Hadiegr A we wrasse oun fat ae pesonlty dimenson—ancety- he beer at 2erted by one of us previously (Gray, 1589) this "has ‘been tetenly‘updaeel (Cry ae MeNaughvon 1996. in pres} ‘The personality wait of Impuliveness (or Im. pulsviy), in its marrow sense, is usually defined 35a tendency to act rapidly without deliberation ‘ot consideration. However, this trait covares With a broader ser of characteristics known as Sensation Seeking, Novelty Seeking, or ver, turesomeness. Sensation Seeking individuals tend ro engage in behaviors that inereate the amount of simulation tat they espenee Such behaviors (e.g. participation in dangerous Spons, drug use, etc.) involve seeking out thei dlanges, and arousal, In this chapter we shall refer tobe vole ce of ned lars ee sentation secking (ISS). ISS ite are typically measured via various standard selfsepore qutstonnaies, Examples inca the IVE (6. GB. Eyck, Peon, Essting, & Allsopp, 1985), especially its “Impul. srenet™ and Nenu nibs “Positive Emotionalicy” from the Muleidimen sional Personality Questionnaire (Tellegen, & Waller, 1996); various versions of Zuckermans Sensation Seeking Scales (eg, Zuckerman, 1979) and associated subscales; an the Novelty” Seeking subscale from the Tridimensional Per~ sonality Questionnaire (Cloninger, 1989). om shore peptic bas been ued that Jmpubiviny along with Anxieoy repre. fear fundamen dents of ores {Gray 1970, 1981, 1982, 1987), Individual ferences along these dimensions ae argued tore. Act variation in the reactivity, oF sensitivity, of 0 basic brain systems to their specific classes oF input stimuli, These systems—the “behavior inhibition system’ (BIS; for antiety) and “be- havioral activation syscem" (BAS; for impulsiv. inp repond a difeing kinds of sonar re inforcing scimuli: the BIS is acivated by novel stimuli and by conditioned simul Punishment or frustative nonceward activated by conditioned stimuli s ward or rele from punishmene. We refer to this account of personality ax ‘ment sensitivity theory (RST), ‘The personality trait label of “Impulsivi when used in relation to che reactivity of pet- son BAS, is really shorthand for impulsive sensation seeing. An individual with a highly ‘active BAS is proposed vo have high scores on ‘ypieal ISS inventories, including ‘those. listed Chapter 10. The NewrocieneofPesonlicy 29 above, RST specially proposes tha subj th hig 8 soe htc toe Boe 3S" subjecs) wil show very song re spans to thore smal hat acca se ns (acs for reward ‘or tli from push nen Gray (1970) argu hac he BAS posse etn, Hae sheer Neurosci (EN) plane of Bena poner, aliy sysem with high 188 sbeebs ee rode exeavers and low ISS cabs ete inwovers, In sbieguentreniong ef ae Enenck seals (eg, EPQ) Hf, Bpecck ne Ensenck, 1975) theimpulvigy cance wear legely inthe EPQ-P seale eae then ERO Gain Reve, 1981 and pia Fests EPQD (parcl = 3) caplet ee ratoae WE inp ea ee (partial R= 05; Dns Fcketng. 999) Bie tae efletig BAS reactivity ems lye snk lesan ER say other EPQ scale Depur Coline iho dae Stn eres oe positive afc) ate lcated on te dngere he tween two orthogonal anc, which the beled Exawenion and’ Consint taker Telest 1982). High 15S subjects ate cher Exerverion and low Constann Gi, BR ek Cont arty EPQLE Depue and Calin anaes rape tee pessoal trai variation raed to BAG as tes along the diagonal besween EPO a EPQP (not EPQ-E and EPQN as orginally gested) TE adlion o standard personaly easures signed to capture vation in Impey nk sensi seeking, Carver &e White 19987 ie recently developed 3 ser of seals che se tendel special to inden the peony xt Vatiation associated with BAS fest They id this by using ms directly sean ea Pychologal ees of evar sa evard ears along with more gene ites conceming el en ed perm sles complement the eating publaged sei Silay designed winder Senet ee iy vation (MacAndrew Be Sele 1S rubis 8 Tebena: 1989) ‘The BAS and the Bi compete with one an- xr fr contol ov: behavior (Gn Set, 1969; Pickering, 1997). When sete tht BAS can be chaste asa “po oe ee scivates ongoing approach belewiog Br ont tras the BIS i 2“sop"ysem tha inher, ging behavior and allons further infra Proceaing of the environment Thea maton appropriate under BIS acivaing conditions in whch gnu 3 pnihmens sr lon of revacds wain of impending neve Cea uence. BIS activation is ale accompani ty sido thee day ad that ae eventual executed (nsating fake Aight) are carried out with inctened foe et vigor From the peupecive of RST. the decison to ngug in tht dangerous but pleat behets iors character of high ISS ubjore en] in tecms of an_approach-avodance tan ee which is eesolved by the ineracdons Leones BAS and BIS, Ina highly BAS tease ck 1S5) subject, the BAS-contoled approach Oe Poneat will end wo predominate ntsc subject wil exhib aloof tay. sensor ing behaviors In arnioussbjec, wih high oe, eh of BIS reactivity, the avoidance comme wilted hold vray, and such asbjoce eal Binal avoid danger and eek At this point, we should briefly note some of the pschopathelogial sates thas have bene lied to extemes ofthe ISS personaly dine, Sion, Quay (1988) has hyposheine’ thatch hood behavioral diordet. such ax seers defi disorder with hypeeccivigg and comics disorder ae in par the result af dyusenod (overactive) BAB. Impulsnry ¢ 2 codon ee tute of these childhood disorden, Corpora ing dsiohibted peychoparholgis in clans have been silly actibed fo an orsreee BAS (Gorenstsin &€ Newman, 1980) or oils cule in interrupting or modulating BAS cor soled behavior Paterson & Newns, 1999) Finally we note that RST bas marly Beer see pling he thar np chophysiologicaleponsesto reward and peren met ce in aisha pani Prtionlty wc score. Subjects performance og 4 tak is asesed under 2 conul condaien oe Which reinforcements are ncither andere ace ssceved, Ths is compared wih pdomanes ey condition in which subjects know tar ther se Spa es prt centre ally guns or loss of all sims of meses) Der Spe the exter of any ma Pee Cot, Powel, Kumar, Thorston, 8 isp (199%, aepued thatthe sheary bas ot been adkquaceh tested in most if no al ofthe publabed weal ons Ie was ao pointed out that within the Feaion of te sy may (apne ings—fom dear supporting idence ‘hrosph sopsigniicant rent to cet constudcion oot Aenccmhave been seporced, Pas hoc sper 280 INTERFACE WITH OTHER FIELDS 2s nad net Soot dct i ln scien ase ranes aap Coase eae RELATED THEORETICAL ACCOUNTS Influential acount of personaiey by Cioninger (Cloninger, Svraki,& Praybeck, 1993), Zuce- ‘rman (1991), and Depue (Depue & Collins Press) resemble RST in proposing a basic dimen- sion of personality that shaves features with ‘he BAS-telated ISS dimension hypothesized by RST. Ta Cloningee’ theory thee are four strongly biologically determined temperament dimen. sions and three more environmentally deter mined character dimensions. One of the tome Perament. dimensions—Hatm Avoidance closely akin to the Anxiety dimension of RST. By exploring the corecatons between Clonin. ee temperament dinenson eel ta hen the EPO! Com Pci & Ga Go) oe lined possible links berween BAS Function and ‘wo of Cloningers other temperament dizmen- sions: Novelty Seeking and Reward. Depend: ence. Novelty Seeking showed the positive so. ciations with both EPQE and EPQLP thar Depue and Collin (in pres) have shown to bes consent fenure of 188 tais, There resale were confirmed ina larger study by Zuckerman and Cloninger (1996). Subjects with high Novelty Secking scores are shart in a iia ay co subj with ighly reactive BAS. They are impulsive, ex. BE os ‘xcravagint. OF particular importance for she pissent chapser is Cloninger proposal that Novelty Seeking involves gene ifferences in lopamineygic ‘teuretransmision; as diseased belo theres now dict rides acne this suggestion Depue and Collins (in prs) recent secount Aifers from those of Cloninger, Zuckerman, and RST in suggesting thatthe snes of causal euros biological infiuence lie jindy with Extraversion and Constain, rather than on the ISS diagonal berween these dimensions. Depue and Cllins suggest that ISS traits ae emergent fom interac tions becween the fundamental Extraversion and Constraine dimensions and, as such, would be expected to have heterogentous neurobiological sources of influence. Asa resul they expecrshet research autempting to find neurobiological cor clues of ISS tts waldo Proce weak Inconsistent results. They argue speeifaly ‘i oer ine en ‘dopaminergic functioning should be more con ‘iene wih Extension tar 8S tats, In their review of some of the pertinent evidence Depue and Collins concede’ that the data are pretty indecisive. Depue's own studies of prolactin and ee blink indies of D2 dopamine tar effects (Depue. 1995, 1996; De Elna, Sat, Colla ee Loon ao ae ated by another group (Netter, Henning & Roed, 1996), provide the clearest evidenss of strong associations with Exteaersion in the ab sence of the significant relationships with ISS traits or constraint, These findings must be set against the cvidence, reviewed below. which finds associations between dopaminergic effects and ISS or Constraintlke tats, athet than ser ‘eas invlin La A final relat of research and theory ems from Newman and his colleagues (Arnety, Smith, & Newman, 1997; Gorenstein 8 New! ‘man, 1980; Paterson & Newman, 1993). There authors have been less concerned with con. structing + theory of personality than with Balding modelo dit tned pehalags Sk a thy. As che eatlier quotation from the work inde, these sts have chat the study of extraverts offers clear insights on ‘hci central question. ‘The remainder ofthis chaprer will build from an analysis of the work of Newman and his col leagues, which wil be considered in some det We will interpret their information-procesing ‘model of disinhibition, and some asociaed ene pirical Findings, in terms of inteactions benween the BAS and the BIS. As we have advocated ele. ‘where (Gray, 1972), this step will lead to the de. velopment of «conceptual nervous system arch. feouce for the systems involved. Pickeing (1997) has suggested thae conceptual nervous system models are nacurlly implemented. a5 computational neural networks, and so we will use our knowledge of neural network sytem to understand the way in wich ou cone tual nervous system might finction. la parce. lar we will consider the effects of systematically varying nctwork parameters to model personal. iny tat variation, The next step wll be to look ata range ofev- dence, fom varying ncurseence medio, hepter 10. The Nereceneof Personality 281 which may provide clues as co where our con- ceptual nervous system may reside in the b Evidence from cel Lecording studies, work with motivaionally disturbed clinical Pelion, lus developments in neuroimaging, and. mo. lar genetics will inpicae the slic slopamine system in BAS function and in ISS personality traits, Next we will ry co map the ‘Shut npu to and output ea target cells of the mesolimbic dopamine path- ays onto the concep nervous gate ge tecture chat we have suggested, By doing this we fan construct a more constrained neural model that we hope wil be of use in understanding the complex relationships ISS personality traits and behavior on experimental tasks We hope, to, chat our genera approach, illustrated this chapter might bean ehectve frameetoh than culd be employed to hop ange ee scurosciene data elated to eer posse trait, THE PATTERSON AND NEWMAN DISINHIBITION MODEL. Patterson and Newman (1993) outline a process through which dsinhibited individual (tach 2 excraverts and psychopaths) may exhibit their characteristic impulsive behavioral style, The first stage of chis process is the acquisition of ¢ dominant response set for approach behaviors. A Very stong approach response set may lead to excessive focus on the behavioral goal, eset informacion. gatheing. and. consierscon of response altematives, producing perseverative ching behavior even afer emvizons imental contingencies have changed. Patterson nd Nevin ges cha tee be Se ences in the ease with which individuals form approach response sets and the intensity with which the activation of such sess i maintained, They argue that exaggerated responsiveness to seed el dh ind eponhle ae hibited behavior. As already noted, they suggest that he relevansindvial ifeence ait Extavertion The second stage ofthe route ro disinhibition occurs when the approach response sec is die ‘rupted by an (aversive) event that stalls oF pre< i further aon, Paes nd Newman Phasize che orienting of atcention toward she uuneepected event and the accompanying. Grease in atousal. These outcomes ae py the output functions ofthe BIS in RST: Further ‘more, Patterson and Newman suggest that the ‘ey individual differences variable for this poo ‘esting stage is Neurticism; Neuroticism islose to the anxiety disnension that RST aligng with BIS reactvigy. (It shouldbe noted that aithough RST has always emphasised the relative closers of Neurotic to the anxiety dimension of RST—see Gray, 1970—the in ba ben con fused somewhat by the simplifying explanatory deve of presenting the amare Ay ing on the 45° diagonal between Neuroticiee and Extrversion; ace Pickering, Cors, & Gray, 189 fhe asin) 3 thied. sage of processing according to Rarcrson and Newman follow mediately a. ter the unexpected distuption of approsch be- haviow Nondi ‘en an effortfal switch from automatic procesing under the reward expectancy to controled prove cessing of the circumstances surrounding the un. expected incercuprion. Once again, this precaly icembles the Reina secant of begin inhibition thar we have offered (or example, sce Pickering, 1997), Paterson and Newman prov foie thar disahibied individual chor akg ‘make the automati-to-controlled i switch and s0 show weak modulation of thet overt goal-directed behavior and the undedying ‘esponse st. From the perspective of RST: disine hibited individuals display functionally imped behavioral inhibition. Peterson and Newman refer to the individual differences a chit stage a being differences in “response modulation and sclace this bias to Extaversion. Ie ca likely that failures oF response modulation 3) in exraverts might e responsible for tee proposed cendency strongly to maintain the at ‘log of an approach reponse set (tage Dy ae though the acquisition of «response te might logically be distinguishable from modulatery ‘processes involved in is maintenance, ‘The fourth element of Patterson and New= man's account describes an associative deficie that usually follows disinhibition of appetiive behavior. They suggest chat weak response mod. ulation ofen leads toa failure co pause and proc. See us thas, ough eating, could eve 4 carly warnings signs of the onginally unere pected imceruplnstt vera acdee Pe, Disinhibited individuals therefore ryprally row ‘fbtced eect flowing puishmens, and the resulting impairmenc of leathing skews re- sponse repertoires toward active, goal-drecsed Dehaviot. "Interestingly, Patterson, “Kosson, 282 INTERFACE WITH OTHER FIELDS Newman (1987) have shown that extraverts do fear from theit mistakes when they are forced 0 pause after corrective feedback, presumably because this allows therm time for the kind of re- rocessing desctibed above, time that A CONCEPTUAL NERVOUS ‘SYSTEM MODEL OF BAS-BIS INTERACTIONS We have poined our the close similis be- ‘een pa ofthe above account an he cone Se of RST, shar Neva cs eaplicily acknowledge’ "Fallow fra Sk wt canoe, Following hep to relate the disinhibition model ec RST using a diagram te Figure 10.1. This Spare baued on the orginal Sages hee Gay and Sinth (1969) and shove mldple compa feng both the BAS and the BIS. Each spun oe sponds to its characteristic motivational pps by producing « modulation our (obich ne hibits th other syne and an onus tha hac functionally exciatoryefict ona gener sae system, The tied output of exchtyrem les feyponse selection snd contal mechssloes he BAS output activates and the BIS output inhi is any reponse habits ltd by oxker nina inthe envtonment Figure 10,1 represents 2 simple conceprad etvous system architecture forthe BAS st BIS, and their sociated personaly tate Neve vee need o understand how such au schieotce mi ncn 4 seat noc pc spay be faclitated by interpreting Figure 1011 Afi wa the ah co Meu nesworke 3c simplest verion of RST would ‘wepors that, i responce to a eamed eve sending ee wad, each ofthe outputs fom the BAS ced be stonger fora high 18 subjec than fore 1SS subject (Analogous statements col be anade forthe BIS and Ansty) If one were $6 construct «neural network mocel withthe be auchiccture of Figure 10.1, the implon venina OF RST would medel ISS tits by lowing he threshold function controlling wie outper Rene BAS “eli” o vary across indvidvals igh BAS reactivity = high ISS = low BAS output thresh: old). Similanly, erat Anxiety would be modeled DP agl te heshldenvaling BIS exp {would logically be possible ro model vacations in the sensiviy ofthe BAS or BIS to their re Spratt Somer Roca Piasone ‘Revsar Sistem Isoacton SS SS] FIGURE 10.1. A schematic representation ofthe in ts to and outpuss fiom the behavioral activation ay. ‘em (BAS) and behavior inhibition system (BI), Tae sousal sem is depiced a afecing response section ‘mechanisms, but tik eo have a diet eet onthe ‘exponie stems. Pathways are functionally exctetory {where minus signs denote functionally inhibitory ‘ffs Learned inputs ar denoted by pathmays ending in rounded arowheads. In this chapet we ague dt ing hibition ofthe BIS bythe BAS maybe isigniicane ‘Ercan am of iin synaptic efficiency (or weights) of the input Pathway to each ten Teed, Pee (1997) used his method as a computatong, convenience. However, for reatons made clear below, we do not advocate this posbiliy here, According wo RST, Excaversion isa personal ig dimension ta ener fom te ttpiay between BAS and BIS. The personality trie a seed wih BAS od Beaty (SS and Anxiery, respectively) are therefore argued to be causally and biologically more fundamental than Extcavesion, Asa result, one would not sede to model Excaversion by varying one specie fe ture within a framework like that in Eguee 10. Testead, Extaversion would be modeled by si. 1 mulrancous vacations in features of both BAS and BIS. In RST, extravers are considered ro be relatively high in Impulsivity (high BAS outpag) and ako relatively low in Anaiery (low BIS out, pur). We (Pickering, 1997) have previously used 4 neural neework to explore the equllbrany see ofthe mucually inhibitory BAS and BIS syems Chapter 10. The Neurone of Peronatiy 283 dire both izveinpus, Foran exavere. ume fom the BAS even under conditions that ac de the simple version of RST, our previo acu Bago steougly than the BIS. Bagatn showed that che ouputs fram the “How wecld ae eee ‘model explain facil. BS we te wile he outpts om the aon of RT in ee se anishment Pama be iacive This ype of model docs Cue? “The nar Gy oe BIS-activating cue En apie cortices AGRS) dominated behay- would real hom the feta oe of the Hane a casiten with che acions of ds BIS aroua ocr ‘opposed both bythe redue= Nbied individuals, but ic does no fic easly with tion is BAe eee roduced by the BIS iieren mA Rndings om Newrars modltionewpur aed Sree, BS laboratory BIS inhibition ouput on reponse selection, acc sep nits our neural ne implemensa- ing in opostion oi ofan oe BAS acd. tion of BAS-BIS interactions Sut when atin tpt an cxtavr of high ISS subs Fre REN cut experienced during appe- ject here sme dee BIS reactivity. Under Sho pote behave (ge 2 af Ftenon ASI: Nanner eag ee dlory Bas onmats Geinhibiton mode). For she with BIS ceacivay ee Exeavetson, and 1S$ Has teactve, BbSunteactive (cxraver) subjecn Waits ae hype orthogonal o the BIS neual net model predicts that there will be intone of anxiety. Therefore, in Extavert of ‘Tpsialfcanc BIS ousputs and thus litle observ= high SS subjece © BlS-activating cue wil pro see apanse the pusshment cue. The neural duce a degre of BS ean Sek modified Bisa apres that the behavior of the account of RST, ust ee ni ae activation by conte BASuneacive (iron) subject, wll nov be supped Se ge oy stronger Py conast wil be affected by the combined ef BAS Scivation fee tect, high 18S sub- Bis ote ibton and sroual ousputsaf he jes) produced Drage BAS-activat- DIS. However, we have pointed out je Picker. fog ea she resulking modest BIS modulation roteencie Gtk 1293) tacit might behard output will nt have sashes Be activae ‘0 determine what the com ed efecs af thee ion andthe mods BIS ia ep wl Fe ks we ties Tees Bredctions not have much succes in oppecag ig Rene Senedisvaiietndag (needa nee 2 song BAS acai ouput eh response ‘mpan, 1986; Partercon et al, 1987) that show lection Hamtscn the modes: BIS arousal ourput CML CRiaret! tesponse cies (RT) ace fc all able wees eet responses di- [ated following punishment relative to RTS fol. rectly and tuneppased leading roan increase in Ueiog teva with the oppose pattern being response spent "Oe ‘comparison, introverted or sbeited for ivroverts. This punihmencrelated tntaus subjects Ch nee gecater BIS re Eciation of Rs in exraerts implies theexit- acres (2 well elect a reactivities), spores one uP from the BIS in se and othe BIS manllnien me ee Cvered nbpenthiene cue, even for an ex: puts may schiow a market etenn any Bisumeaca st who viewed as being Snguing BAS medina alien fee ey BIS-unreaceive. pf a punishment cue on appettively motivated feacriet fo explain extraver’ RT faciiation behavior wil be determi eet by these BAS- Fpeing Panihmens, within she concet af opposing ete ieee ci the effects PERE LOT itis neesary thet the BIS can re- oP he Bis sone oe increase in RT more acon Sry hee the BAS is activated may therefore be obeebed inet, 8 RE sare auene Such a state of affirs will not sponse to punihorrees, tecee BAS rong competitive inceractians be- We have now comple our conceptual aerv- PIpeBAS and BIS that we hae previously em- ous system analjon ere possible gross neural a- Bye Pikig, 1997) ic eold eis fom check bag Rene Beale Fee ge EE 82 (modulation) output from action, We have se Understanding of Ee BAS that acted on he BIS diet. In this mutually nhiony cee dynamics ‘be- tach BAS outpace seas the eel of tween systems gatered Rom exernann yeh SEARS outpud would be a function ofthese. neural pemwork Segal ie PE teformulae SIT inputs and he inhibitory modulation tion of RST to secoteee findings of (ead sac the BIS. whereas BIS actvation Newman and college In tems of Figure 10.1 {and ousputs) would be solely determined by iu we rae simple modification: to drop the a ot be dven to rro fo raul therfore tow denaing rec inition of he Bee de ‘ot be driven to ze0 by inhibitory modulation BAS. Thi wharreclane Fepresents # more 284 INTERFACE WITH OTHER FIELDS dented specification of the response modula- tion account of disinhibition suggested by Pat. terson and Newman, We now turo to evidence ‘lacing to the neurobiology of the BAS more di. ict by exploring real Rear nce brain, and how they function under appesiive motivation, we may be able to map the abstract representations of Figuce 10.1 onto actual wer ware in order to consteuct a more consrafned and realistic model, THE NEUROBIOLOGY OF THE BAS Evidence for Dopaminergic Invalvement in Appethive Metvation We have prevoutly given a thumbnail serch of the neurobiology of the BAS (Gray 198), 8 censtal oe for dopaminergic nouovansr ick ib BAS fincion'& mandated given the money dnc inpleaig rel and meen dopaminergic patwaye in tewarlcieced fen eth & Wise, 1981; Robbins te veri, 1996). Thi isnot to argue that dope, tnnetgicneurouansrision im moatimblc rok stays excuely linked to postive meta totvatonl eect, at we hate emphased co, whee (Gray, impress: Gray, Kuma, arses, Noung, in pre) Ie i cer for example thy aversive stimuli increas malinbic erin B newroutansmission (ce Slomones [394 fr 5 review Iemay terete be more compete argue thatthe common denominator une hing the capaciy ofa stimals to cc dopumiee ot lesen mesic pathwaya ita tienes ce Gray et lyin pres), Nometeles, we apc Bee that one ype a salience land hence one outs tmeolabie dopamine reese) ba produce of the ation of te BAS: A major gol of thr ace, ‘er iso highligh he pouible Rpusane see pathways eee ‘A particulacy pert: series of suis has been summa by Schule Romo, anes Mirenowice, Holletnan, 8 Dicanaae (Ooee They" report call rcodings from monkey dopamine neuron in he suas nig (ON, and ventral tegmental sea (VTA) thee ee that dopaminegc activity atest nape tapi revs, Aer ing howe te cal respond vo conditioned seul (C5) tea predic the primary rorard exer an he prieary reward ea, Such CS aeeodig to ST, represent one ofthe case of stl en activate the BAS Houk, Adams, and Barto (1995) ofe an in- tetpretation ofthe results described by Schuler et al, chat is emttely consistent with RST. In the schematic neutal architecture of Figure 10.2, an sppettive unconditioned stimulus (US) leads to increased firing of SN or VIA dopaminergic ges, Dopamine seas the al get cls S in Figure 10.2) of thee dopaminessic neurons i argued co be an addoced ed factor invelved in a special form of long-term Potentiation (LTP). This LTP isthe basis of con. dlcioning in the glutamatergic conical afferents carrying information about CSe to the seat calls, Aer conditioning, «CS i able o eli co bust fing ofthe star eucon inthe bene of the US. The outputs from the staal cells send feedback signals that contel the dopamin: «ttic ourput fiom che SNIVTA neurons, A CS ‘an produce cwo types of striatal feedback signal: gas indie ld 3 freon eaten of dopaminergic fing (by inhibition ofthe inh toy inp fom exe uel vo TA ‘neurons); Houk etal. (1995) argue tha there i aie a moch lowes dtc signal (nt shown in Figure 10.2) thae inhibits dopaminergic ii 484d :0 opposes he increase procuced By theese suing US. In this chapter we are going to ema: Phasize the indirect pathways, The other cells receiving dopaminergic input include the pre: frontal cortex (PEC in Figure 10.2). These te. gions are involved in seleerion and control of Such behavioral oucputs as learned stimulus fepans (SR) babi As well a pole ke in ‘modulating learning in these pathways, dopaminergic transmission may also play + rect role in selection of the stimulus inputs that undergo learning (Schulte et al,, 1995) and in the psychomotor activation of the selected be- havioral ouxpucs (Wickens, 1993), In terms of RST, the outputs from the swiatal calls, elicited in response to an appropriare CS, correspond to che outputs of the BAS. Impulsive sensation secking subjects, whom RST supposes to have a reactive BAS, should therefore produce strong striatal feedback signals in response to CS chat hat previously been associated with t- ward, We noted earlier that RST suggests that this ineeased BAS output resules from 3 lowered ‘oucput threshold of the cell concerned. ‘The Houle ec al, model, summarized in Figure 10.2, describes a form of classical conditioning in cor. Eicostriatal pathways by which inially neuttal Ss can become a secondary positive reinforcer and clicit conditioned dopaminergic cell fring Chapter 10. The Nearoxiene of Penal, 285 be] — l. us FIGURE 10.2. Outline sketch of part of «neural archi ‘tecture forthe behavior! activation sysiem (BAS), Pah ways ae exciatory except where minus signe denote inhibitory connections or Yahaped connetions {alse ‘marked DA) denote modulatory dopaminergic synapses, Round-headed arows denote modifiable syarper tow. beheaded atts indicate reciproal connecrons, Some ‘connections have been omied for clatty The indice pathways from arial neurons to SN/VTA cane! vs lida nd subthalamic neurons, which ae eft lank the figure. S, medium spiny siatl neurons, SNAVTA, dopamine ells inthe susan niga or ene ep ‘mental area PFC, preftonal eorex, US, appetve us ‘condivoned stimulus; CS, conditioned simula, {and hence striatal dopamine seleas) after pai ing with an apeiive US. RST has argued es vatiaions in BAS reactivity are nota finetion of the ie wich which such C85 acquit shi se ondary ceinforcing properties (see Zinbarg Mohlnan, 1998, for ah oxclene Geet Hence, there should be no relationship berwees ISS (BAS.related) personality traits and. the serength of the corticostratal synapses, formed in the way described by Houk era A link be {beeen ISS traits and the output threshold of the striatal cells is much more in keeping with RST emphasis on the behavioral effects of tablished secondary reinforcers. From the foregoing account, a CS reviously associated with reward will nis the tees, seraal ouput utes, initially eicc high lovee of dopaminergic fing to an’ impale eet ew we Wl develop 2 moc dead ah tyodel of the BAS," bung ‘ant fhe a shetched to date. We wil wggen dee sul upg nd de oes crease in dopaminergic fing, can pea a behavioral sesvaon fear ase ena ns the BAS. We wil aio make a specie ogroice eee te bere for ‘ical ouput ofan impale senses seching subject At this point we boul cen Gate our cals sateen Nee a Kinde of simul can lc dopunipe tet. aaa calls Inthe more complet acdc gee fenced below, we wil bry sopses hee sive and other events can elicit. dopamine release inthe venta suru, Some of ee eae a be interpreted at vesling fom te renee of BIS outputs with BAS procesing, Emphasic. ing ths point is imponant 50 th, the dots no mistakenly infer any spe eave Berween dopamine rise sn epee vation, We have sted eewhee tee gah view is at odds with a great deal of evidence (Gray et al, in press). Sth eee OA a function comer Rot Hadley of aa EOE clnal shnormain of appecine mena Yowell and her claps ive developed toe ile experimental mesure of tewanl ematn, the CARROT (Card ArrangitgRewied Ree siveness Objective Tes), sine for week clinical groups. The CARROT is x repeaed. Imes tesa which subject ate teins perform avery simple cand sorting ak SG icky as posible on for separnte conse rials (T1-T4). On one trial (13) the subjects auc oflered a fasncal incentive for ft soning (10 pence for every five cards sored) snd dog soring rate on th evanted tad coy "arin nonevarded walrcon ducted either side of the rewarded trial (T2 and 14) yields “rewad esponsvense ink ‘These researchers ae ted the CARROT £2 54 patients with ogni bin ine anno whom were characterized by cll gestae impairments of appetitive motivation (AL Adzwi, Powell, & Greenwood. 1998), The oe waudtesponsivene index coveted singh with independent ented raspps of moneene dlring therapy sesions (r= Gk p< O01) aed 286 INTERFACE WITH OTHER FIELDS this reflected neither a general efect of mocor slowing (peed in the wonrewarded tial was une

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