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Philip Morris

Some Effects of Limbic System Self-Stimulation Upon Conditioned Emotional Behavior

Date: 19610000/P
Length: 11 pages
2060435243-2060435253
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Brady, J.V.
Conrad, D.G.
Harlow, H.F.
Area
LEGAL DEPT
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PSCI, PUBLICATION SCIENTIFIC
BIBL, BIBLIOGRAPHY
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Bcnj/Produced
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N28
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2060434863/5330

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Named Person
Nauta, Wjh
Author (Organization)
American Psychological Assn
Journal of Comparative + Physiological P
Walter Reed Army Inst of Research
Date Loaded
16 May 2000
UCSF Legacy ID
ikd22d00

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Page 1: ikd22d00
THE JOURNAL OF COMPARATIVE AND PHYSIOLOGICAL PSYCHOLOGY EDITOR HARRY F. HARLOW CONSULTING EDITORS FRANK A. RLAeiI ,U91l11 V. nRAqY P. L. IIRUAUIIURir W. J. nRIK)nnN WILLIAM K. FStlE FRANK W.FINaV.R FRANK A. GNLnARn CLARKNCR H. GRAHAM G. RORnIT C/RICU JmIN F. HALI. MARnARISr K. iiAR1A1W DALe 11. HARRIS Jo11N l. LACRY DUUOLAS H. LAWR8NC1f DUNALO B. Lamnsl.l:T FRAN[ A. fI1nAN MELVIN ii. MARX DIINALn R. MnYF:R FRYDGRICK A. Muw CA1NRAn Ci. MUKI.I.I.R RnnMRT A. I)A'RnN CARL PFAFFMAN KARL PRn1ReM IiAtRUir RUelNnnl.n AUSTIN 11. RI131iN ELIOT STRLLAII HANr-LueAS TevnnR ARTHUR C. HOFFMAN, MANAOwa EDITOR it.uN ().t, ttOMOt10N M.M.NtR VOLUME 53, i96o American Psychological Association, Inc. Copyright sq61 by the American Psychological Association, Inc. 2060435243
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J..•.J y 1•-nn•un...J I•s/r,.11d )SrrLIM/ Yd.11. 6b. 1. I W S(IMIi i:FFECPS OF I.TMRiC SS-S'PIiM SIiLI -5'I'IA4ULA'1'fON UPON CONDITIONED IiMO7'IUNAL BL[IAVIOR )(1SF,1•11 V. Itltdlll' ANU UUNALD O. ('ONRAU+ 11'alfR Rrrd .I nny Lnnotr n/ Rr.rnrd. Sl-vcral rcacnt replrls have dt•w'rilxd ilucr- Aif:TUnn ~ aclion clferts iuvnlving clcclriral ti<If-slimula- (;ruaa! !'rnrrrlurr lian u( x•k•cled braiu arcav :md a widc variety ' uf phvsiuhrgit'aI and Inychohq;iral phenomena. 5'smaleelhi:..rus(120tlaysoklat lhesurt+Il the , ' eqvimmu, uen seannlly nmure mnle cate, .u,l Inur I{aJx•rimrnlal demnnslnniuns ul Jn.ilivcly malure mak rhe.us n.mkn.xrveri as5s. AII anitnals Winlurr im; t•Rcrls Jrrtxhu'c'tl liy inlmt'tanial ~erc lrafnnl in kuR preniuR La lerlh IxainsJvrck to- ad(-NilnuLdiun in the rat (UtdS & AlilnCr, inh,rcen+entamlvwtef(ratslnr6.dlrals.m/n.x,krysl 19-a-1),ral(Itrnd•v,19.i(I),andmunkey(lifady, rn.atl.Alleraatxbleperlurmance.vastwuldishsl,lhe IO.iRLI hace nnw Iw•cn cxlendcd In an analyzis rnxakxul can/I1lwxlh+R pnmlure was sulxximp.M ul",n the kvR-IxeasinR hase Gne Iry rryxalRl I+-urinp uf relatinnships iuvnlviug clt•e Inelc hx.:•fl iun n( a clickin6 nui:e maminl for a haerl interval (d .w 5 (IR+1., 19iG), stimulus inlenaily /Rcynnidx, mind unl IRminstnl cmrtieunusly with a hrief hul . 17iX1, •t'heduh•s u( • rciufurcemcnt (Sidm:cn, Iptinful eltelrJr rh.rk In Ihe udno/'a feet (llunt 8 ISfntl•y, Iloren, C'unrad, & Srhulmnn, 1915), Ilruly, 1951). K•itlr aermlaaiully implanlnl ehrnnk I ekctrrales in the limldc q'µrm d all auinmh, the Lxxl :unl walcr tlrlvlvallon (Ur1uly, 1)ufen, dufaclRiaic.o/Ihe('P:RuRrnmqurmlumlervui.vs ("nnmd, R Sidman, 1957), ~cx !h~urmum•% cundit«ne nf ekctrird xlbstimulatMm and h..l• .x ((0d+, 19.iRaL dnlXs (OI114, Killam, tr Ilat'II•y ,t'llR-rewatlltfl kVel )VeNn/,. lllxxl PMII•klxb o11/R Rlla, 1'/Jfr), mnl sa lial iutl cf$Mtx (( Htls, 195R6). Iclrar:.x.l aletrvaiwn, all nnunak wrre aaai(cnl, and Signi/iranlly, hnn'cvcr, Cmnliun:d nr af(tctivc IlreelRtrnleptacemcnuverifxdhi.lukrylically. limu4cs trlalyd In Ixrsilivrly rcu•artlinR hrain 1lfhaeirrralCnrulilinuinp and 1•relinp !'rnrrAwrr slimlllallun IMVe reraivell lillla t{Irltil t'Y)+efi- '1'M Irainlna PnKehne uanl lo Rtnldish the CER menlal al trnlinn. L•'vrn Ihe few puhlislu•d oh- io the nts haa hlen PrevinuJy tlesr'rihe,l in e.mm de1aR eertaliun, rt•laling cScaJtc Ixrh:tvinr (Rulx•ns, (Itunl&IhaAy,1'150.IIriellv,allsisratswrreiniliallq 1958; Itoscvr & Millet, 19iA) aud )clin-siuuk trailxxl in levn prccdne. f+rst nn. nxuium.n and then t•Ildunnlrl• in an uh\Nrllrlion-Ix:3 ailuallun anyx•rinlicrtinfwrrn+rnl¢hnhde(VI/11ftCl."I"hm, ((lkk, J9iRl') lo electrical scll-nlimulaliun c(- a aRks nl eight enmlfunl canlili.x:htR Idats was uF minislRnl, roch Irial u.osisling td a.1-min.lnextl/ai~w (lr'tx, hmcevrt, sugy,t•sl Ihal lhe delim•alian uf ol adicking :xrix (CS) lerminaul cooliRtuwsh• with a curh "emnlinn:d° inlCmflinn5 may fnnlrihule momentsry elaetric slnrk in tl:e ke1 (iN:rr+sinmtely siptilil'anlly lu uur Imtler5l:tndinR of Sut'It I.S ma.) as the UR, Jrliven.l thnx:Fh Ihc pill ll,.x M htain-In'haviuf ttdnllunshiJK. the pryaralus. Mrwt nf the rm:ditinnhrt uinis x'erc q.eRl sevval davs a).u1; nnly mw rnnt/lihminR Iria7 'ILc• (ut•st•ul reJxul ft•Jut•sa•ulg a tlirt•e'1 :11- ,,.ae Riven tx, any ,me dsy rlar111F Ilre acl(nrAluxt 1lllas I('mJtl IO /lr•Ii11C lhr inlt'r:rCllnlls Ix.•twQ•It Iltlr:l- al the eflRrlmnll. /ht inlerveninS tlayf lhe rata 16 tr:mi:d ~If-nlimulaliull and :nmc relalivcly cdvnl m,rnul rtms 'm the kvR Incvine alTnratua 1'ift 1a/1V'rllw.•rl •lsJxtit nf an arRinRn1'S 1•mtf- •%ith neither clicker n n shu'k in mder lu aAapt uul liunal lx•hac•ior rcJw•rluire. Slxrilir:Jly, Ihis RenetalNA l"Ihl/' ffal'I+M+) \O the eayerinxvttal epace aml Ix•rmit a mme clrnr.rn[ evdualinn of Ihe mp~ t-Slw'rlnlt'nL is C/ttu'cfnttl wlllt IIIC l'Xt'Clti uf I„~nRrRrrlulthrrrmdltlunnlclickRflimuhMaM.+ eletttita) mll-ptimulaliou in st•Ictlfd IimLir- Ihe n+imai's lehavino With lhret• nl the nb IK-d1, s)'~tt'm trKions nf Ihe braiu ulnnr Iho a(quisi- K.13,am1(:F2R),IheU':Rnnslrsleslablislrnltlurirryl limt :md mainlCnanoc nf a candilioncd cmu- kvR-preadn[srui,n.lun+atRrdnfortrn+enu:xllhee linn:d re•rlxmV' (CFIC) nf Ihr "(rar'• or •'aux- INed with the annn• animJs lever preadnK lor en invatnnisl elRlrital slimulue rt•wad. For Ihe n- Icl)'" IS'Jx• (Itnuly & lfunl, 19.15) in I:Ilnrtnlui}' maininR Ihrve rnls IGI'rJ, W-fr, ana OF-55), the - raK. t:tl., antl mmokcys. r:nM4n.al rn.diUmdna nrurniure waa iniiially tarrled out rlurina Iever pre.<ine e+•.•Iros fnr lhe electrical ' Oe xiJt ta ecprnn aur rmliluAe ln lV J II. Nauu hrain ntlmulus reoanl and Ia1R ustnl wilh Ilm samr lorhi•es(,rrla••ntm.ceinlhrprq+arsliwtandlnterpre~ aninalnlevRlnesin.fnrnaler, tatnm .d the hisluhx(tcal nuoerid ratalred (nc thix A rlmilar uniniult Vr.xcvhue vns folluwnl tn eslah~ snul.. IBh Ihe fER in lhe cats and au,nkevs. With lhcu 125 2060435244
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IIRAIN Ar1Rllrl.A't'f(IN ANII l'I1R CIiR 121) hllrx Ian Rr+Mqn, I+nwevrr, ail auhnala aa•rr lailialir mcutd rexai+o. IM arklili+n, aqnratr r n+la M e re Irained in kver IueainR /nr o fM.l renanl lmilk (xr 1h+• ror+Ml f+r +ach unlividual clkkrt imrrral l,t n+. Inr rlia a1MI anRar IIrIIetS hn 1)If I+MMIkY)'a). f+r\I +nl a C/M- ra15, S min. I+N nta arNl n.mkm)• aMl /m au hurvvaI IimNnn and Ihen on a vannlde.inlerval rci+dXrr+•mmnt uf Iheun+eJuntaxrinm+nlialelvpraenimRea<hdiAe. a[hniuklYl(OSrc.).'flmll!ReasealahlishnlouJ¢ inrrrval'ILisrsmrdinRpntnlurrPmvhk+ll.wqum+tili In.f re.arA kvrrpreaanR luu• linc iqlnrY iruralutlimi ntkmaf thf ('6R in tnm++d m"in/leclirm rmx+" ea- nf the hrain stimnlalinn. "I'Im roa+liliN++ina )u+cnlure pr+.unR h++w m+Nh an+1 in wGNll +lirs+xeo Ii+e di+ker wilh 1.16 the cats mu1 (he nnnkecc invuknl a Sarin. c1+anRnl the nlqnrt nl lerer reywwsrn uMlrr thr+liRer ireaenlmim+u/ the Cti IcRckinc iN.i.ael tc•aWnatnl mm rnt onrdithnn u( rrinf+rrfenrtnl (hrain dNSA rrr /•xx1/ liqarlrslv u'ith a d-ina. eln'Iric shnt lu Ilrc larl fnr ualo/ usni in Ihis ealnrin+rnl (atc Ilm+t. jrn+lnrR• & 01 sec. Vur the cata. lhie eh+rk uaa drllre.nl Ihrouah Rrndp, Iv52, (nr a r1MMe Cl/IIrI•kte rI1N'InaMM +J thia the Rrill f4.n nl a n.xli(cal 5kiur+n IMrt (2 fl h, rahId. All rrenulinp nf Ihe aninuk In~InvuM and 2)1. by 2 il.) runairuclnl mith tiNre aluminum ssArs• a Praltmmo+inR ni the es).rimental prnenlutrs wr+e ar- Rri11 hdrt, aud a/daas tq.. With Ihe nemkeya, all Inim canqlfisiN.+i aulmnauraliy hy timns, mapnrlm rnnmrra, inR aMl I+sN++R was acamqdl.haal in a q•a•clnlly cuu- cun+n4lhcu+xk rrr+x.irr., and aa ru Ir+l rrlar nlrrctnl primale chair prevenn/v +/arri)wd in vmre cirruils, rletail IRlnrm, Iv5R), uul nlMrk una+klirrtnl/hrmlRh dalnak.crw+uetnllnthesfalamllmlrsas.IlnlhllM: (IPe/ufittand 11i.alrAnRirnl !'rncnlurf cal. aml mrnknT frenuentlv nceiverl mnh'ryrk IlairinRe drikkaand.Imck+lnriuRaumtkkvrrq+raafrnRa.xMx+ 1'he rkrtr+xle v+mslmrli+x+ alNi implmuxli•m pn~IhrnuRhrrut ac.ryisilbw lrainlnlG Im the l'IiR. With «+lure u.nl nillr the ralx mvnlvnl a IiIMJnr sl.vink>s- I/N+e1w++Rrrul+s+Jauinnla,arqaiutm+..d th<erM+tN.mi stml A+rtrnle uMI Luatile )•nlnlal hkck lx'+Ihx+nl nud req.arae Maa mx limilnl tu a livni mmdnr af Iiiala aa frtnl in plare aa rkvrilnl I+r Ubb aml hll)nar I IaGil. wil6lhe raln, Inlt crmlinunl ualil alahk anNxqawn+ nl 51 ilh IIm rnl., a mnlliknd ekrlnxk arnn' .a • a lever Ixnwna in Ilm presenre nf Ilm rlickfr nns nuah. ulasirally im)Janlnl saud+aR tu a pranhue )ursr- fe11nl.Th.n.alllheanln+alsueret~nlfrnllveRrclsnf mnh- drr<rihc+i hy IkIRatk. 119521. rbnl wilh Ihe inlraenNa) ekctrimi seli-stimu)aliun ul•ul Ihe ('IiR ninrkeyn, a multi'kad ekfltm)e. /x+kaa) u s dpidlc AurinR rsl•'^mfnlal raaiwwn inruicinR allrrnalinR ( ni in dm indiratni kwntFina by a nrrth•.d r+'rn+/y rer'..lani krer t+rc.v11R fM (.xl reM'atrl v+MI f+n I+rnlu- rk.rrrlMVl ln' Shnlr (I'J41)..SII surRrrr u'na I+rrlnrl+M+l Clack rnnlMMllerla. I)'INVMN)', rlYrlnR Ihi. InlinR nsqeieallp un+kr Nem4rtal aneerhetia• mid ILr ani- Idsam n/ the olw^nN'nl. .Mi min VerWls nf vartahiv~ nmh anrln'rmiltnl a pM+qnralia'e rmuc.rn Ixri+xl 4 inlerval (md-rdnlnrYenrvni krcr pressiuR allnnaiul alhaslnrvernldayatXa.rcrklx~h+rcln~harunaln~IMe rkrlnxlr pl.+rrmrma wllY dll.min. INri+Nle iJ variahkiNlaval luain eWrk- IliYUk~Ri+ai verifaali•+n uf the reward Ir'ver pres.inR tlrnwRMwl a+•aai,an+J 1 l0 2 he uaa •JnairN+l kn all 12 animal. ury.n omry.lnkx+ -d 4' kMIR. I'rMrtat:Nl d the c&krf anJ h.+t JMni JurinR hnvknal /+<ti+ra lLe m.inul. ++rre In'duw+l a i+h <a<h rA lhex Ml min. ryrlk. Inovidnl n rrliahk 1+.1 n( plymnl•lRhnl ~~I4nr• fnil•nral I•r IIY',. h.r+naliu. i IlKefkclsa/ hrain etinnikllhm upni Ihc ('I!R uhirh Ihc brain. oerY Ihen lias+l lur •xv n• I••ur orrk• 'fhe ekrlrival hmin slimalrrs nvrl Iu rrinhxm IIn• Iufore rn+In•rldinR in Iwallia S+.iai urlinn •+ r rnt kver~pravina rry. Xxrw• in Ih< v~ILqim+dalnw Ixnlbnn a1 ISa an•l alainnl rilb U+i•n+1n. 'I'hr rh.'trnlr plarc- d Ihese nlNrimenle .us Innvklnl hy a'I"rklrm~is notsMCrermmieruclnlall+•rmirn•.n~~h.•r.uniimli~~n stinluhtlinR un4t nenlll+erl In Rive . Idphaek pulae M all )+rains, aml ldnunmionn;mpha d.A+rle+l urlbu+a airnilar In Ilul dr:<rilw+l hr IAIc• Ilutahes, Alvord, aud hu rarh anin+:d ..rrr n+a~Ir. Gakin (iv5p1. l'I+e I+^rx+m~l+ra d lhi. folraeranial I'Intr.+4•ylarn.+eats(+nnlll2nnimahlntldaa)udv ekelriCal slimulus wcre usun/l) kelrl rnmlanl Inr endl xre r xatrminl i r Ihr Ph•. L.mirn•er.qd+. +Inxur in aniaW IbrtniaM+ul the murR• d the es)annn+rl esrq~l Fignrt'a I, 2, and .1. 'I'I+e ret u.'1i+n+s IPIq 1) are rnt ~nllhrnvurraaYnswlM'nan•JrorvMcl(mlwnxapkxnl npiwrnimalrlp I b~ 1 nuu. r Imnt ut /hr Inu~.•rrv~ u a/u1MkXa n( the I+rniu-ahwk a)iumhn intnlsilr as Iimi. nl Ihe nmrr:.r+.u+nui.•urr an+1JYU+tlre rln/ndr inPOtnl Irthnr. For the rms, u'ilh r~ialnrre valrrm rr.+fla r.nminR tlnnngh Ibr n+nlial aywlls nl Ihe .plnvirna)ir¢ Slanahne, rarh )naiu~siM•rk rtinhucrIxain. In Ihr,c nf Ihrsr a<9irnn, Ihr atln.nlalinR rlrrnw+u c.m.ialeal ++f a alinwhls u( ItRI eyoles In•r Mr., Ir+nk 11)q Can IY' YTn rll tl+P Ml)nal'l+++Imlaarlml P~r/ Vn1a OLn.ve. puk duralkwy 25 Irak nu. fm a lrain nf the nytal rraim+ 16 .12 :wd K i,it mnl rinpdute +imlk+n ol 11.5 sec. Wilh the mts. nsianurf values unlfe (CF 2RI. l'hv remaininR Ihr+e ra1 ~+Ih•na dxna unRnlletwren.i,(RR)mMl4ppnhnnaMlcarhrllrlri~ the .IMrMk Ii1n.~mnviiat nx+rY ~k"dy nn,I anrallraimsUnwluuriid+xernlrnlenu.ntnl+Jnuinmlus IakMly Vlurnl in Ilre v uf Ihf n+nlinl hnrlxain nf IIRICyCInlrrsrc,I).2 nnaT.)+IIUfrI11ra1MM,N)In.4I IIIIMik Ifi1' h1 aml \5' hl+al++l ihe nXfifNr n1lX/nAfx+ Irat n+a. uith a 415 ve. Irnln +turalkM.. l'ieatw refGF SSI. Rrcnnalnu liim n/ hraiu x.Ynmx /•v Ilrr I+v. tistun< values in the nnn.hev. nly........ a/e+l I,Sn) tu cars us++l i+ Ihia stu+ly arc xMmn i+ Viqure 2. /'at 211111nhnr.. and Ihf hnin-durk alinmlua I.+.+ridrvl h.r tirrl6m ('-Nlfi is rut alqMnsinulrla+llu•xrRh I.nn/nl an electrical alimtJ+n d IIM) cvcln prr ve., 1/.2 rnser. Isvel A 1.5 111unk'r l'Iark rundlnalasl ami sh••us Ihr puhs'doralixn, 31) ina. fnr a I).S.vc, Irain duralksl. clMnnlr Inrl ruurainathruaRh the I+rnd nllhrrarr•Inle - 97m+uRhun dle murw nI the ea/N•rinlenl• renrtrb nu+lms, /he interlxd/u. 6nIL nf the vuerxu +:nmia .em mainwinnl nl Ihr' Inlal nmu)n•r ni Irv+, rfysanea anre, .nd Iho area rrl the nndia) Inrfbr:dn Inunllr• ollh adI rcinlrrrnuenla frw r•arh ani+nal dhlrinR faeh rrls•r+. tie rkrlasl I'enbalinn Iennir11r1inR ahrn.l al Il+a 2060435245
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Isu J. V. Iik,111V ,\NII I) G. IIINkAtI 7 . II-.k µ or-af Fln. I. t-u.r (wwrr phntaakngmphs M hruitl vcln~ns Inr sia mu shnuluR eleetrtale plucrmems in Ihe eqrul raxiln anrl nnletlur Luellrain. lusikv .IU/am lit Ihr Iuafu. ('nl G.~dm C 1077 is rrn alyxu.iuratrl. Ihruuqh Yrnulal 1.1,1 A 14 (IlarJry- ('I:uk I'..r, linalls/ and elarws Ihr Nerlnde trnrl kavinR Ihr hrnd nl Ihe raullale nud..m, cuur.iuR thnelRh Ihr Intvpbr reRilrn and the arra nf uu• nlvlial hreluain bundk. ..ilh Ihe dl•rywal In•nrlnlh.a in lalrral lip nl 1Le qllie chiunl. 'I'hr ntiumlalinR clro In.h-a aw•I oilb Inllr cals in the Prt.rnl rqn rmrnl n tl,. Mad nI t he vvdnle mmle.n. „xl Ix• elrrt r,.lr plxcementa ua.I wilh the Iwlr mnnkqs arv slwnn in FiRUre 3. Srrliml MIV7 i. cu1 al'Prnri. matrh• Ihrnugh 1•nnu.l Irvel A 17 Itluraley.( L rk cn. mJ{~utcs/ aml ahnas Ihe elt•rlrvk Imak cnarslnG Ihnru¢b thr raudate nuckua, Intt•rnal nlwulq nnd rhd.r pllidlr.. nith Il.r sunnrfalinF lip in Ihe arrn ul 1he Ilmolnl lnfrlNalll IIn11,Ik. ,tt'Innl hM.3h la 1'nl aq.nrinmlcly Ihruudl Prnntal Isvcl .1 IS (Ilnralrv- 1'tark rundilminl aml aSain ah,.wa Ihe eleclnvle kav- inR the caudate unckus nnll intrrual cnpsuk In pasa IhrnllKh the .nler6n mrlnniuure an.l ddrnt pdfidln, ailh Iln• slimulatinR tilr tenuiluHnR iu the area uf Ihe m.+lial InmLrain Lundle. Sertiun A1 cl ia cut slunrwhal n.ul• aulrri,uk IhruuRh aplur.at+, Frm,tal I-evcl A SI Illuraleyl'lark er.ndinatr.l an.l ehn.vs Ihe elec lrl.lr Irack rnnrsing IhrnlMh Ov rnrOnr rulfnfu~u, nu•lale nurklls, aud inlrrnal cnlsuk lu Mrminale wllh !hr rlimulatin[ lip in Ihr avra ul Ilm nurlinl Inre6ram Inmdle. I'ilullv, tirrlinn hl 2 ia rul snamohnt Irume Lnlrrhnlv al al•ptnailnalri) fn,nlnl I-rvrl d 16 (Ihwak~('Lrk r.n.linalral au.l dn..a Ihr rlrnr,ak Iralk Io.vin6 Ihr cawdale mulca• arnl Ldrrnul raPallie In 1.•nninaln wilh Ihe aumolaline Iq, in Ihe mr,/ral ay~la nl lhr rhdnrt }nfliAul. t1F501-15 '•Ihr thrt•v rats (R-72, R-,i.i, :utd (7P~1N), iuitially Iruint•1t in Ievre lur.~in(; for a te:ul:r rracard :ulll rxl*latd to tile rrtlnliuu:d cundi- . tiuuing prle'rdura hrfnrc• inlrr.lucliun ut I1te'~C rlcrtricnl hrain .limuhta, alquinvl Ihe "anx- ic•IV" rrcprrnu• liv thc vi(;hlh tundilinninx Irial. All Ihree auimnlz 4nlqs.l pre•.ing Ihc Irvrr duriug dm rlitkrr prrv-nlalinn, culd all dt•la•rnvd during "frials6, l, aud H. VChetl lhtv same unimals wrre sn•iltheJ fnlm .vnlrr In 4rnirrv6nik rt•w:rrd nn t6lr r:uuc c:niaLlo intt•rv:d reinfortcmrnl rhrdldr (avrra(;c (11 src.), Ihc Ircrr-pn•svinti r:uv tvas nlniutnim+l al a c'untparaWr xud uaLlr levrl fur .rvrr:d daily .csaiun. I><furu introllut Iiun nf Ihr t unJi-~ linnld ":ulcicty" <linmlus.'1'Lr first pr•arnla- linn a( Ihe rlilkrr duriug a Ilrain-shulk rrim forremcnl se..inu ccilhin unc lvc•ck :cflrr Ilv riKhltl emnliun:ti rundilinninti /ritd, hmvrvrr, failt•d lo prrrllule vttplnesann nf Illr h•veo- pres.inR ralc in any of Ihv Ihrrv r:ns. "hhi- virlunlly cnnlplclc nnrnnaliun nf Ihv l'I•:It ,unlinurtl for all three r:us 111roughnul righl <ulcrrding p:ririnFS ut tlilkrr nnd pain hluk In Ihr ferl Jnring daily raprlimc•nl:d ar..iml. 2060435246
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IIkA1N till\II'1.,1I'111N SNb'IIII! r'Flt G-1697 G-1697 I•IC. 2. L.wlawct p1a,It..r,iaraFml.ha d Lr:nn a..'~ INyn I,u 1„n . wp slx,e inR rlr, l r,wlr I,I:n wnvnl5 dn•1. 1„ Illr rlm,late mu Irua in ILr ar, a„I I I,t• rnrd4:11 I,nrlnnal 4u.llr, iuvnkinlk rlrtlriral u'lh•limulali,rn. 11'illl nur ctl QK ~.i11, Ihc.v nlnlv..icr daih' pairinl;<n( rlidcvr :11]II paiu .h.x k<ttln•rimle,~.~d ny.ro litr kt•t•i-prc•.inl; h:nc lint- (ur all rLvlrital brain- tlilnlliu+ rrwud rt]nlinurd fur .42 rall'rilnrnlld vvium ttilll na (udir:tliuu uf a r:UV uqyRt'~- ainu ill rryxnl.r lu Illtr I lilkrr. hiVuro -1 dunr. the Iypit;d rr>Inni u( ral 6-S! In I III• "1 Iit krr- du.k" pairing, durinti ILc Itr.l, ILird, xnd ciF11161'ICR atrpli.ilinn Irial. ntlllt-rilnlroM'd ulnn thr Walrr-rt•x'ardcd Irvrrprv,.iul; 6:Lt• lilk, :wd duriuq Ihr •ulrvquvul liral, riy;hlh, and Sitlt.•nll] 1'I{R Ir,l Iri:ll. ailh brain- aimulaliuu rrillLlnrnlrnl fur Ivvr'r prt•..illy;. R:ua F:-33 :Ind GF~IR Iloned c.,i-nli:llly ILt• cmtr rr•ull. I6run911 Illv r•iph111 6rain-.Iirllnla- liuu IcV Iri:d, allrr tt•hirll Ihc Itr.linf; Irrlr- rtrlurt• dcv ribrd Lelo]v tc:l" inilialcd. Pullmsinq Illia u'ril" uf Ivaiu-ainwlalinn k'vt'rpre+•ing o-t••,iuu, anll (•I•:R Iri:ll., ILI~w• 151 ~ • ~_ ~ ~- 1.~ ..A 1 I I•Ill. J. Ina Iw,ut< Idlartau\INraFr:q,llc ,d L,alll a„ Ilinra Il,r I,Nrr rN.lll.l•\a all~r~tr11F 1•Irl"1. , r„u-„la.,1 tlle ~rt•n ul ILr mnhal Iu/rLraln I.ulull. :md .Ldm. p.,lli,hr. Ytlr£F REWCr1U INrprr]Rrw~ro,l nn I -Wr[re' L(./ SrMLnlre/ Ff MrH) Ivr~µwt.~i ~~ rv .°-. I'u d. ti.ll,qdr , ,nl:,lt. ,+Ix, , . .r,,,..1..... n,r ,./llnlli„n Iri.'rla Im Ihr 1-I It a111x' r,1lu...~,t ,Ilu„ r aud iidm Ir.rr Ivr.v„4 Ln .•ari:ddriul„nal 1, 111 namal rlr,lrir:d .IimWmn~, rnl ILr ,.Lli,pn ~•Inl nme 1„di,.uc ILr ,.rl ~,( II,r1o,,,~Lu„o,.l : u.61~,.. aliundua. ninl Ihr ,.LlLlur Iwdrn arr,nu nn6,alr tln 1,'nnnnliun aI Ille I~S enn114nunvl. ,..illl IIn' 1•n.~l u ndllnm,.I Rn,l s6.. A.1 iumhl< b. I b.. Ir.l dw mc r.,1L Id.]I. .:Imr Ihrrr ral. Ik:-32, f:~l.t. :ml (ll' 141 tvrrc tl.nlinnc,l r.n n Ic.liuC ~.L,dldv LLi.h prrrcitlyd Inr uhrmalinl; aal,~r :md Llaln~ .lintnl:lli.m Icrrr pr,'..iag, .r..i„In „u ,uur. 'irc d:lya tvilh daik t'1•:R tri:d.. IIU vcrm mmmix•re11 Ilav.I uh,rn Ihr :ulinval. 11„rkrd in ILv Ivcrrprr..ing app:~ruu. 6,r u:llrr rrtv:l]d. :dl Ilrtar• mlincd. Inutinurd In .Ivnt .uplnr. eiun uf Ilu• Irvrrprr..iu~ ralr• durinC Irm-nl:r linn uf Ikr rli.Arr. Un Ihr ..IJ~nuuJnrrd davc, Lmcorrr, vhrn Ircrr rr~lvnulinl; u~a, rrinfnra~d ttilll Lr:lin •liluulaliun, dll ILrrr ral. tvrrrkrd rig61 IhruuyL Ilu• rli.lrr aud .Luwrd m, iudilali.m nf Illr 1'I(Id, liuallq, nflrr 61 d:y. ua Ibi, nhrrnalt'•3.ic Iryiul: pnnl•rltut•, nnt~ ral (1: t.ll wo• Ir-lnl durinc a 2060435247
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132 ). V. IIRA111- ANII II. (1. CONRAD single 2-hr. rxls:rimental ses~ion wilh allernat- in resl.mse tn the divkcr dcspile rcpemed I hlg .AI-min. In•riuls uf lever prccving for a pairings ul lite audilary slimulus with pam walcr reward and lever pressing fnr an intra- shnck to lhe feet according to the same pro ~ cnurinl rirclriral-stimulalion reward. Figure 9 cedure usal with the waler-reinfnrcerl animaic I shuws dre rr•sults oblaincd when pra•scntminn Following the eighth emotional ennditinnine of lite rlitkcr followed hy pnin chnrk to the trial, two of the Ihrce bmin-slimul:uion mis feel was programmed during carh of lite suc- (W-6 and GF.63) were swilcherl to wnler-re- tes.ive .NI-min. tycles. Although virtually warded lever prexsing and given eight addi• rnmplrle cuplue.sinn of lite Irver-Irressing tinnal CEIL acquisilirrn tri:J.. Allhough Ihr.t rate in resl+mse to Ihe clicka•r (thc i.nliuu of two animals showed nn evidenre uf "Iransfe/ Ihe rurve belwt.•n Ihe two small, heavy hlark czvings" from 1he previous aunditioning trials arrnws in enrll .tp-min. uugmenl nf lite record) during lever-pressing seccinnS fnr brain stimu- r.m lu• v-cn during lite "watcr" I:criwis, lever latinn, Ihcy did develop thc conditinuerl suf. I rrnlwmding rrmlinucs uunf(cclcd Ihruughuul prt.•asimr µzllcrrt in rrs(ronsc to Ihc ditker Ihiscmredirkerpresentationduringtheiulra- during Ilrr eight surcecrfing water-reiidnrra-I uanial rirctrical-slimulalinn pr•rt«(s. menl lever-pressiug tri:As. 'llte one rcntainin6 'I'he renmining three rats (GP-(.l, \V-fi, and rat in Ihis group ((Ih-55) ruutinur•d In rt•rcive (:P-S5) were initially trained'in lever prr•sint•, pairinge of clicker mtd ).rin-chr.k lu lite fert lur lite elerlrical bmin-slimulus rt•word, aud during daily levcr-precaing scssious l(ir hraia nll:udmalsdevelulocrirclalivelyst:ahlerr.(xm+c .tinrulalion. Afler 23 nJdiliuuni 1'ISR trials r.rla•s un a variahle-intr•rval (nverage (A sec.) with Ihis aninral, pnrlial suppressiml of the rrirdurcrmcul acha•dule wilhin a week fulluw- lever-pressing rnte Jirl dr•vclop in re=)nnse to iug rcruvcry from surgical implantation of the Ihe clicker during hrain stimulalion, nnd Irnns elr•clrorles. Iu marked contrast with lite pcr- (er of the couditiomvl "anxiety" rrspunse was Inrmaurc uf the three ralx (K-32, K-3.1, CF-2R) readily apµvrent when water rciuforcemenl inhi:dly trained and enrotiormlly ronditinnr:d was sulnlitulcd fur brnin-slukk reward. on the w:rler-reward schedule, however, nnne 'llre two cats (1G93 and 1697) acquiral (lie nf (lie Ihree hrain-slimulaliun animals (CP-fi3, CKK within 12 pairings of cfirkcr and pnin 11'-G, Gf-55) develolK•d Ihe "muicly" re- shrrk supcrimiroscrl upou thr Icvcr-pre•sinR 5)nna• Juriukr lite iniliai series of eight arrryisi- hn:e line for a variahlc-inlcrval milk rein- Iion Irials. Nunc of Iht•rc iattcr thrre ratv fnrcemcnt. Allhough a slable resinr.e rate shuwr.i upprex+inn nf lite lever-prc+sing rttlr was maintained for the milk rcward durin{ uer m daily exry:rimenlal sessinns, prescntalinn of Ihe clirker during Trials lU, 11, and 12 elicited Ihe typiral Icvcr-prccinQ snppressimt paltcm (Ilunl & Arndy, 1'1511 arrunqxmiul in Ihe two rnls by pilrKrmtiun, dcfecuinn, and immo- hilily. ISnlb animals were Ihen placerl on an alternate-day schedule whiclr invelved exlxri- mcm:d Irvcr-pres.ing scssians fnr an electrical hrain-slimulus rcwnrd on one day and lot milk rt•wnrd Ihc next day. 'ilre samr• (A-:er. variuLir-irUrrvnl Iarw nruernmmed lite relm ru[.ruuS torft•mcnt schctlulc nn Innlr nayR, :ow nn F'm. 5. S.rnple cumuletive rry+m.e anve ahn.dnR mtimnlc develu)K•d : umpnrnLle and stahle the dlllere:uial rfleel ul tlre Cltlt pnrrdurr• durinR a Irver-presviug rales under irulh cuuditinm. unak 2 hr. eqsrimrntal rc.emn wilh alternate .MI min. Iwrir+ln ol variaLle-inlenal antrr mrd intrnrraninl •And when Ilte randilirlnrd "anllely" slimulu• chArlual slimulntiun re.urd. llrt uLlirpre s,lid urons was itrln+luced 1lllring Ihe viil(r Ittai11911111ulaindirale IlR uu+et nl lhr rnntlilionel auditurg atinndu., i inll 4•sviull, INrIh rn15 t'onl lnu9d to .hUw t'oln- and thr nhiiqnc hndcn arrrun indiratr the terminmmn f,lclr cupprec~ion uf Icver pl'es.ing in rcs(>tmrr ,J rhr ('S aml:RUmmlr uilh (hrbrid mrr.wJil ional eridshr.4 <Iimulu, lu Ilre fecl durina each ol 1he ahvnsle Itr lite rlirkeri l4durlunatrly, pRmalnrc IrrCnk~ Armin.0 age af lite eicctrorir Ieads at the Ininl yd 2060435248
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tIBAIN 51'tA1111.ATlI1N ANtI TIIN CP:A 1.13 . emcr(,nntc frum lite sralp iu IN.Ih thcsc cals Ihc icel pr/Nlurcd lite lypiral suppn~riun made it inqmc5ihlu lu pursur this testing pru- p:ttlcrn. 1)uring allcrnalc-.L•ry scssions, how- ccdurc Iu Ihe camc cxlt•Ifl as witls Ihc rats. ever, when lever precsing wa% reinfurced by Ilut ou two surtrssivr rn:rasiolls with ('at 1673 and three allernalc-rlay scssinns with Cat 1697, presculaliun of Ihc clicker during Icvrr- pressing scssiuns for hrain slimuhuinn pro- duc(rl complete sufquessinn of lhc reslNmsc rate anJ chlbnralion of lhc lypiral Cf•:N hu- Icm iudislinguixhably similar tn lllr animal's reaction In Ihc rondilinncd "anxicly" stinnrlus during milk-rcinfortcd Icvcr-pressing s[ssinns. AI (casl willl Ihc intlir:ucd rlvrtnnlr plarc- menls in Ihesu Iwo rals, inlrarr:nlinl .el(- stimulalinn had nu appart•nt effcr( ulN,n thc cerz. With lite four rnnnkrys (M-197, '1'hl-.SG, M-53, mtd M-2), rlcvclupmcnA of a.tahle slqr- pr[Rdun lallcnr in resl.rl('. lu Inr=eulation of the [undilunrt.l ":ulxicly" slimulu< requirl.l mllny mnrc pairings of clicker and sllurk, ac- conliug In lite pnat-durc dcsrrilN:J alNlvc, Ihan in either the rntx or r:us. All lite monkcys were initially lrainell in lever prezsing (or a v;viable- inlerval sugar-Ikllcl rrwnrd anJ lite (•EI( supcrinlpo,,ad ulvm this bav- line In•furc imrn- Juclinn of the hrain-slirntdus tcin(nrrcmt•ul. Under a procedure invofving muiliplc rmo- lional condiliuning trials during individual lite vnriablc-inlcrval intrarcnliai alvrlriral stimulus, Ihc cunditioned aullilnry slimultts followad Ily pain-nhnrk to Ihv Itrl prrNlurcd nn Ia•rturhalion in Ihc levrrprc..ing rate and nn overt signs of cmolional dislurh:mu-. 'I'hc rc- xnlls illurlmlcd in Figure (1 wrre nblaincd when thc aninull's behavior xa< oLsr•rvcd during a single G-hr. ralterimonlal sr•ecinn wilh 3 hr. of faNl-rcinfurctrl Irver pns*ing aud 3 6r. uf Lrnin-shrn'k rcward. 1'rr,cumllon of lite dirkcr auJ foot silnrrk on Ihrcc <I-Parnlc rarnsions during Ihc 3-hr. sugnr-In•Ilcl ~gmcnl nf tile .(ssion (inlfiratrJ by tlre Ihrcc pairs nf cm:d1 arrnwe in lite upfa•r sr( lirrn of Fig. 6) prldurrrl virlually t.omplctc supprc••inn nf levvr prrwing with alrnmpmlyinC •ignv of agilaliun (ur Ihr durolinu n( thr :mdilary .limrdns. 1)uring lite lasl 3 hrc rn( Ihr sr..hm, hmrevcr, whrn tltc :mimal vnrked uln Illc ~tme v:viahle-uucrv:rl rrin(onemnll •nc~dule for br:rin sh/nk• pr[scntalinn n( du• lnuJil innrd "anxk•ly" slinmlus and pain-.h/Nk lindil:ned YOIIREY M-19r Y/allr[[ INrrMll R-YrNt Icver-pressing srssinus of a•veral hnur<' dum- lion, nll monkrys JcvelulN.-rl lite rrludhiourd suppression µtllcrn wilhin 42 pairings of clicker and pain-shlx~k lo the ftv;l. Grrns signs of agitalion anrl gcnrral INNlily dislurlNmrc W (piloercrliun, vol:diralion, hylnrmnlilily, mld ° drkcation) usually arcompanicll the dcrreasc f in kvrr-Iuescing rate Juring presenlalion of A, the candilional "ansicly" Ntimuluv. When thc Sy s~ inlsumniaf elcrlrirnl-slimulus rcinfnrrcmrnt Y s~ in lhc indicatcll brain :Irras (Pig. 3) was sulr $oc elillu[rl for sugar Iwllcls and thc cundiliaual :oo rlirkcr slimulus sulwriminncrl rqam Ihc x•If- I".IimrJaliun Irvcr-prvssing ba.r linr, however, o' allt fntlr Inonkry. nlwwt.l allt-uualilm of the ('r.f(. \Vilh Munkry n1-177, lhr- :dlrmaling-rl:rv csllcrimrnl (+crfnrmcd nilh lite m15 was rr•- IN•alcJ Iwd prnviJtvl a mml rnnvinl ing rirmno- drall/llr nf Ih[' Irll['llnYllttg t'(fl'1't\ n( Illr Irrllll .limulalinn un lite CICH. fln lhc evan davs, -ben lite animal Icvl•r-prrvnvl fnr sugar (Ndlrts, pr[•rnlminn uf thc rli/ kcr and pain .Ilurk Irl NlAw $NaM IYtAM rMrMH r1,w0.11 -~T W IX IIY( NI YINUI[5 Irq F'rc. 6. SnmPlr ran,ulnlivc nry.m~~ r nlum{nc Ih. rlillrrv,UN rn~.1 ul I6v ff It pr.Nrlun, •Inrine • nnniuum,a 6h, . rq...,r.n,l:.l u,.i„n oil6 11m M1ra i Lc lw aueur Iwll..t n~ld..rrrm.nl and n- I:rcr I Lr. I•.r Lrnin ah..L . rrl ILr uldl.pu . di.l a.-„dH~ , x tr 11 r umrl A 11 r runJilionrvl wdibn c a nuuln• 1 a.ul Ihr nldiqur /n.. 4.u nn..n. iudi.av Ilu~ 1. nninminn ~d Ilm 1~\ nan~ennll•1\ ,.ilIl lhr INU,nnlnum„•I cr„I M..k duuclrn 1.. th.~ Irrl .IurinR raih ..1 Il,, n.o I. N.I. 2a6o435249
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1.14 J. V. IfKd11Y .1ND It. G- IXINKAU l6r lite Llninn< of Ihc curve bolwtcu Ihr Dl-5.i, Ihc rrilirat Icxls fnr Iltis t•Ifca uf hmin thrtt• p;;irs n( smnll arrmvc in Ihr ImvtK a•ttinn ~Ii;nulntianou lite ('I•:R werccondurlt+lduring u( Pig. 6), pnahltcd uu dt•prt•ssinn in Iht• thrtc cx)x•rimrnlaJ u•scious invnlving a Inla) Icvcr-preccinK rate anJ nu gencralis.rl rmn- nf 16 CF.R tri:d+, R during lever presdng lur tinn:d Ji.durhaute despilc Ilu• ubvinu.ly Itighcr sugar (x•Ilcls and 8 durinK lever prccaing for lever-pressinq rate during the sugar-)x•Ilel lite inlrtcrmtial electrical-stimulus rewan(. ptlrlionc uf Ihc rcrurd Ih:ut tfuring Iltc brain- ll'itlt ht-2 nnly ;t single rxilcrimcnlal scsaimt slimulalion (wriutl. It is alur of snmc imrrest involving Iwo C6R trials during hrain stimu- to mtt• Ilte slutrp (nrrt•asc in Ilte lt•vt•r-prt•ssing Ialion and Inn l'f{fi trials during sugar-Ix•Het ralc u'hit h ap)x•ars immavlinlely (olluwing Ihe lever (1reRtlllg could ILL' relnt(>tl•IIYf IIChHt• l,Ri pain-Juak lo Ille feet during thv lower "brain- of cleeltale cunlinuily imerrupltYl further shutk" 1"ulimr uf thc rernrd' ((he churt seg- lcsling. mrnta n( thc cutvc fulluwing carh uf the Ihrt•c With Iwu mnnkt•ys, onc.S showing virluallv hnrkt•n arron•s in Ihc lower ecclimr uf Fig. 0). cumplclc allenuatinn uf lite CRR durinR No +utlt rate thangc fullnwiug hxrl xhot-k is hrain slinndmiun iu lite mct(ial (orchmin aplnuelu during lhc sugur-(ecllvl lwrliun of buntllc (h1-197) aod ottc S shuwing only lite rctnrd. tmrlial allcntuuion of thc (:I:R during brain "17te rentainiug three monkeys shnwtrl sumc- slimulaliom more amvriurly itt lite aren of the what Ies, drantotic Irnl etjually tunvioring ul(aetury tulxrclt• mtd mtvlial ftneJlrein bwtdk c(4+IS uf Ihc hr.tin sliululalinn ul.m Ihc "anx- ane olhcr clcclrnlc plarcmcnl was icly" re<lxtn:c. With lhese anima(s, all lite Ie.tt•d for interaction efkcls wilh Ihe CF.R. tr(tiral It•.IS far hrain-stimula/iun cOn~ls ulwm hlnnkcy kf-097, will, nn atidilionnl pair of lite (-1•:R wcrc rundutltd tlurinK ca(x•rimenlal elttlrtx(t•x in lite ttnlerinr nuclcus of Ihc lltala- lrrt•r-prr..iug st•xcinns nf scvt•nd hnud dura- nrus, showed mt equally slahln Lul stmtcwhat tiuu wilh allrrltnting .AI-min. (><rirnk of sug;tr lower levcrnpressiug rate for a variaLlc-inlcrval fKllcls nnd nf hmin:shtxk rcinftncnlmcnl. eltt•triral-slimutus rcward in thix area rnm, Alnnkec 'f\I-.46 was Ies1cJ dnring twu .ttoh parad willt the ;mimal's (wr(onnanrc (nr hrain t•+Ix•rinremal xs•iuus involving.ix ('6fL Irials stimulalinn in lhe ntt•tlial farchrnin hundk nn(xtinq.is.'d upm lever prtsing (nr hrain area. JSuI nn hvn CF.R lest Irials tluring sel(- .ha k and six ('(•:(t tvi:dx .u)krienl.,vd ulnn slimuhniun in Ihc nnlcriur Ihalnnms, Ihn lever prc..inK (frt sugar Iltdklx. lLis ;utinraf mmnkcy ehuwnl complrlu supprt•ssiun uf Iht >lmwvd no suppres•tinn of lite rcclwuse rate It•vt•r•pn'ssing rate in rec(xmsc to Ihe dickct in du• •iv ,litker Ix•ri.xis during hr.titrshrKk- rnmµir•thle In Ilie wel6eetahlichtrl CP.R rtal- reiu(unc,l lcvvr prassing hut a rumplcle sup- tcrn nhscrvrd during su/;ar-(xllcl (a•rfals. prt•«iun o( lever prtxsing in res)xnlse lu all Similarly, Alunkcy AI-53 shuwtvl sLtble ra•1(- six rlitkcr prc:cnlalioux during lite sugar- climulalion lever pressing tlunuglt un atldi- Ix•llcl -n•tcnrd Ix•rirnl.. pulh Itf-i.( anJ T1-2, linnal eleclrede pair in lite anterim Ih;damus, hoervcq shawvd only p:vli;d tutrnmtlinu of hut nn a single Clilt Icsl Ire(orc luss nf vlt•c- the ('I•:It eluring brain-shrrk rcinlurrcmcnt Irale t'unl;u'1 rnmplcle suppressinn of lite Irri,.I+ as indicaletl Iry a cunrislt•nl rrduclinn rr<pmu• rate during Ihe clicker prtvntalinn (:dlhuugh nult complele suppressim) of Ihe indicatttl little, if ally, elicct o( lhe hrain locer-prrssing rate in rt•s(ronsc to thu clitker shork in Ihis :nca ul.m thc runtliliwud enxr during thcsc Irortinns of lite cxlKrinu-nl;J liunnl Ir•hnvinr. ('learly, Ihe efictts of brain >e..iunc I/uring the sugar prllct pcritxls, nf shtxk ulxm thc C(SR in Ihesc two animals at ,uunc, bolh animals shnwt•tl cnnlplete sttp- leact, dil3t•red marknlly ax a fum•linn of the prrs.'ern uf lever prt•ssing ("iullt•climl rmios" clet1reKle plnu•ment dtaing sel(-stimulalion. .d -I IXl) amf grns Inefily aKilalinn Ihrough- uul Ihrrli.krrpresenlalinn.I)urinKtbcl4rain- UlscusSIOK FIInInIat111111Kr11xIS.I111wCvPr,'dInl(lYt1U1Ir1Iln' Illl' rl'$Ilhi of Ihis ezlKrimcnl indiralt rducc ,yltproximalinK -.51t tvcrr tnn.islt•nlly quite tletinilcly Ihzl electrical st•If-stimuialinn rrtunh'd hu Ihr.c Lvu auitnal<, indir:ning ;u u(theLrnininstietle~llimbic:yslcroshnclnrr Irasl puliul allenunlino uf Ihe ('I{K. 1Cilh t-nn markcdl)• aut•numc a well-t'stahliJial 2060436250
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IIkdIN S'I'IMOL.1llUN :1NU'fllh: Ctat Ili rmlinnal revlMlnRe in labnralnry aoimolc aamr aninml are IrslrYl for iNeroalinn rRtvlv 1\'ith Ixuh thc r:stc nnd monkryn studietl in a•ilh Ihe ('(r.R. Although wnvplivrh'al al Irmm- the cxlKrimcnl, a Iut•viuusly ruudiliancd lims of Ihc cmntional rrqxmw• 1\:Is cvidrnl aversive slimulus /ailnl In prcxhlrc ils usual (luring elrclrical scl(-sfimulalinn of Ihr mrdial suf4nesaing ef(rci nfwn responac rale when fort•brain hundle, Ibe CIiR aplx•nrvd in fullv k•ver pressing was mainlaincJ Gy brain-sbor'k dr•vclu)w.-tl form when prt-ruletlinn c:f lhr rcinforcumrnt. When thc Icvcr-pra:csing hc- tnndiliomvl aversive slimnlns wa. w)x'rim- havior was maimuim.l hy Ihe murc rnnvcn- Iw:etl uprm lever prc.xing fur bmin zh:xk in lional fucel or walrr reward, however, rumplclc Ihe anterior d)alamus. 116% linrling woultl sr•rm elaboration of Ihe CER was observed even tn provide strmrg supfarl for Ihc cno:iu.imt though a close lemfnral rt•lalinnc6ip existed Ihat the rcinfnrciug profx•rlics uf elrtlrital between Ihe brain-shrx k and Lxxl-rcwnrd slimulalinn of different Parls of thc brain may tesls.'fhere appcnml In be na Jurahir rhange dcrive frum quite dilferent hvhaviur:d :unl in the CI{R which oullaxled Ihc ac1u:J Ircricxl physinlogical clfcc'Is. (-ertninly, Ihe evidr•vice o( clct'Irir:sl scll-slinnd:uion of Ihcsc s<icelal frommuhiplrmtimalcxlx•rimenls(Illd.,l'lSAel limbic syslcro structures. imlicalcs that lhc rewarding effects of hr.rin 7Lc available dala, however, provide un shotk rovary with a wiclt• variety of mmdiliun< definitive accounl of Ilsc failure of braiu nlimu- (hmtger, sex, drugs) us a(uuc/iun uf slimnla- Lilion (n allenuale tlm CER in Ihe Iwu rnls liun.ile. And the hndings of Ihr (ursrm studv studied in Ihis expcrimenl. llu mcnl obvious wuuld seem lo cnnfirm the cunrlu•ion Ih:tt explanaliou of the different results nblaimd rlvclrieal sclf-slimulnlinn prnb:rbly d... nol with the eals is suggesll'rI by ml :ulalyslS of have tl)c aime general significaurv h.r Ihc Ifle ekclrofe lnvalions. 1•:Ieetricad s•1(-slimuho- organism regardless of llse annlomi. al I.- u• J tinn in Ibsse two nnimals was canlinr.l to Ihe Ihe rewarding cleclrrmle. head of the c:wdale nucleus• None of Ihe six II is still nol clcar, nf caurv, rilhrr Ir:mr ntsorfourmm)kcysprnvidalsclf-.Iimld;Uims- thG exfreriment or frnm previnus audir., jus CER in4mvtiun rlala fnr similarly 1)larnl wlml 1.¢havioral signi6alnrc is to In• ntlrihmrd ekcirodec-'fl)c Fact, huwevt•r, Ilml twn of lhe In this rewurding Lrnin-vlimulalinn ph.-nnmv- monkcys ttid not show aucnu:ninn of Ihc (:RR non. It would hr lcmpling lo,Ix•. ulnlr, i.u Ihr durinF elcrlriral vI(-Mitnululinn in Ihe an- haais o( Ihr Presenl limliu);s, ahnW 11u• LuWn- lelior lh:dumus (allhough lltr suue animals lial "auxiety-rrvlwing" prap•ni,-I ul 111111 sbowal a markccl n.lut'linn in Ihre slrcnglh nf br:rin aimulntinn, at Irasl in vIrrled limhil the emutinmtl IKlmvinr during slimnlnlinn in system areas. ('crlniniv, Ihr cvidrn: r: from Ihi. lfro arca uf (he medial lurchmin bunrlle) dca•s ec(wriment Irndatrnn/;w)plxrrl tn,urrrnl (he' lentl some sulqmrt to fhc (xKsibility Ihsl Ihc urrtiral fnrmularioac (IlraJy, I')iAa) vi.iJl rk.clrtxlc Itxalion may have In•vn lhc vrilic:d (x)aulalc an iutim:ur rrlaliun•hip Ixlwrrn Ihr lactor in Ihc failure nf thc cals In show allcnu- fmwliunal r hnr.trlerislits ot Ihr liuddr ~c•wm ation of thc CTiR. (If ruursc, Incscnlly availa- :uul Ihe fnn)x•rlit•c of an urt;auieu)'+ rnu,tiun:J Ide data do nnl Ix•rmil ruling out a hcncl of behuvior• A(urlhrr indicatiuu nl Ihv - h.rta./vr olher faclnrx iurluding .Ix•rit•s tliRerencev, of Ihis relalinnship, a, well as n arnvr:J onor- Ixamviural Ir:slingprcnrdurt.,:ucJ Ihclikc,hul m:ninn of Ihe prc•cnl IinJing., w:c~ ..61aim•d Ihe nvidcncc is dcnr frnm Ihi, aud olhcr in nn cxix•rimrul in ohi.h a('h:(t .ca- 1uIea- 4lxfics (Ultts, 1956) Ih:n Ihc analomical hn'us iml.)setl tqtnn Irvrr presciul( for watrr rr..:vd of the ch•rlnnlc is a rrilical drlrrmin:usl nf using a(b51ItV1'ly rrlntnnlnl, hrain Ju„k a• INain-slimulaliun r(fcrls. the US fn fdare af Ilrr a.ual paiu(ul hrn, k 1-, Perhalr.c Ibr mnsl zlriking ohurc:qiun de- Ihr ft•e/. •I'he pnx.ahtrr invulvnl prrwnlulinn ricnl front Ihi, cludy is thc diffcrn•wial effects of Ihe rnnelilioutd cli.ktr ~tiundn<Lrc rrl.-:urd a( inlrat'raui:d vI(-,limulalinn u).m Iht• ('RR 5-min. iutvrvals whilr thr:u)imal levrr-,rew~d as a(uurliun uf t•Ictlnxlr plara•mrnt. \Inukry !ur a vari:dJr~i,u.rrv:d tvnlrr rrcv n,l 1':::, 1, ?6197 pruvidcs Ibr.9varrsl picturc of Ihc rrla- stimtdus pr"rnlnlimr .va. lernriunlr I."mlicu' Iku,shi)nwhith.'unublaintvhc•nlw..l.nilivrly uu,lvwilh:c~iu4lri:ur,ur.u,ial rlc.ni~ •I:.i.k rea:rrdiug elntrrxle hnuliun• in unr ;nld d:v dvliverrvl ti:t a rrcv:uJinc rlr.Innhpln:rul 2060435251
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136 J. V. I{HA111• AN11 U. 1:. CUNH.II1 in Illr srptnl reginn. l'hr tat had no prvvimic ro.(:xnv In Il.c +li+kor Juriug t•ighl addiliuual hi,b+r)' of Irvcr prt•seing (ar braiu .h,vk, and ('I<It Iri:d. Sn(K(Illl)qvv( u(nm w:uc•r•rt'm~ dirt.i 1+urrelaIiun ielwet•n x kvvr rrs(nmt xnd IurvvJ Irvrr prrwing.'lllc rcinfnrtiug electrode Ihe intr.xranfn( clerlrir:d slimulus wa1 prr ptaremenlx in Ihe xic ral. wrre in Ihe u•plal venlc.l bv rcquiring nn inlcrval u( al (casl rrgilm aud in Ihe arva nf Ihe mnii:d (orcbrain 2'l'V. fallowillH a rcv(xalsc M•(urt Ilx• hrain buwullu. . xlxn k awld IK• dc(ivcnd and Ihc t li+kcr lcrmi• 'lllc two r:rts :u'rluirtvl Ihe (-4R within 12 nalwl. hl luulrasl with thc usua( suppresvixg pairin>;v uf tlirkcr and (win-shock su(Mrtla. e/(ral x( dlc rnndilionc•tl clirker slimu(us u(xm laet•rl u(.+u lever pressin); fur milk tcward. Ihc rrNl.mse mte u•hrn pa4n-sh+w k to lhe (vet When a(m.ilivcly reinfnrrinH hrain <hrlrk wur is uvrl. the slnpc of lhc Icvcr-prczeing curve suhsliluWrd fnr ntilk In mafnlnin lever procsint. shnwal a sh:rrp iocrcace duriuR tllc 5-min. bnth vale shuwed aomplclr retenliun o( the clicker iulervals as t'um(.zred willl lhe ha,,r-liue l'(ili. •!'he recwarding vlertr«Ie plucemcnlz ia , r.ue% drvelopcd durinti Ihc 2U-min. inlvralickcr lhesc twu anim:dc wvre (nund lu Iw in llte hend iu/crc:dc, Indced, (wvitivrly rcittforring brain of Ihe cmldale nutleus. shak ucetl as the US in Ihis "cxmlilionc•d With 111c four maukcys cnntlilioniuG of a anxietv" p:lradigm nould sccm lu prrslutc a stahlc supprc•s,.iou lrUlcrn (CER) taryircd a. 1'RR nf r(nile Ilw op(rnite tharaetct. A r'learer large ntlnllMr /1( Iqtlrlnr,5/t( rlirkcr mld shnrl ~ pitlurc of Ihe Ihentcliral and pmclical im{+(6 (up to 42 in tlne animalL All fnur munkeya cnlinnc ul Ibt•sn 6ndings, howcvcr, waultl however, 9howCYl nllUtnnllnn nf Ilrc wcl6 duuhlh•,s in• derivcd most t•xlmrlilioual,v from eslabli.hcd cmolinn:d resprmsc Juring cx(xri- a more prerist• exlKrimental almlysis of sueh n+rnlal sesciuns involving elrclri<'ni ccl(-slimu. complec inlerrelaliun.Mip.. lalinn of Ihe hr;un. When lever prcacing vas SUUUnur maintaintvl by sugar-(x•Ilet reiniorecmenl, Ihe <ulqvt•ssinn a( Ihe rt•sp:msc r.rle JurinH PnaR Si< r:rw, hrn cnls• and four mnnkeyu were talion of Ihe ruudilinned rlickcr slimulus wx Irniurd in It•vcr prt•.rinc fur hnlll Lrain vh:nk conqrlclc.'ll+crcinlxrciug clcttrorlcplntcmenu atwl trnler ur f«nl reward. A tnudilinnevl ctnn- in Itutc o( Ihc munkcys wcrc in Ihe area of Ihr linnal rr=l.mw (l•1•:R) of Ihe ^fcnr" or medial (ureLrain hundle, :u+d in Ihc fuurl4 •,:ulvirly• IyI,a wac su(a•rimlrr:cd u(.ru the mnukt•y, in Ihe mt.linl aslw•ct of Ihc Rlnbn Itverpres.ing base line in all mtimals by rc- pal(idn.r. (n•alnl pairinga of a+litkine nni.e terminated Rl'kF.HHNC7iS luotiHuun•ly wilh a hrie( painful electric sh.xk Howr.e, ti. 1L• & Atnxra, N. E., Hcvardinn ard In the fect. With sterrolaxirally implanled (mniehinR eRmts Irom aimulatinR the ume pua tlrtnnir 1.IrJrlwles m Ihe limhic ryslem. the inlherxr'ehrtin.Lou~p.phY+ld.l'rydid..IVi( J+nr.alrri.lira of Ihr ronJilionrd t•motinnal 61, f!a-671. IK~hnriur wcre t'omparnl under rarinus rnndi- Hanur, J. V. 6matinnl Irhavinr and the nrrvoa xyet<ur. T.aa. NP Acad. S<i., 1954, 16• LAI-I linnc uf cicrlrital Af-slinml:niun nud (axl-nr M2. walt•f-rewmdrd lever pr<..sinF in all anima(5. It.env, J. V. A mmlmative appnmah tn the empen-1'hne uf Ihe six r.us given cight inilinl (7•:A m.•ntd eralysi. nf rn+nl'n,ual IMhari,x. h, P. training trials dnring lever prresiug fnr water Hrch & J- T.ohin R:,IeJ, Iiape.inrenmJ p.yelk tAdnr.. Nc+. Yu.k: Grune arxl Slmt(un. 1957, A rrward dr.elo(w•11 stable su(q+rs.ion Im1lcOrs p t dnr..Iln.rrr, J. V. 7~he t•ulnYMma nMl I,CI:nvNral nxMinin rt+)nrox la the roudilimu•d ditkcr slimulus. thm. In 11. P. Iinrlnw .k C. N. \Y,.drey (IGhI. Whell brain shaek was suhsliluled (ur water Sidnrice! nnd b(urFrMicul baree oJ brhnnm. b1Witn maintnin lever (+rt:ninR, Imwcver, Ihe CIiIL °'^° Univer. 1Viuunuu Pre.u, 195N. tn) L+iieJ lu nplwM.'1'hc remniniug threr r.tlx werc ttauu., J V. l'rnq.ual aod cm,n:mal Inclars rclatnl. lo elMrin ll xlGainmlelion nl tl¢ lind,ir .y.m. inil(:11Fv givcn cighl ('P.IL Irialc su(+erira(,uscd In RrNind.+. J~nnnlinn ol lbe brnin. Ik»I,m: IAUIe lyem a laver-prrscinH basc line ntainlained hy Hrnwn, 1959. (bl hoain-=ha'k lCx:ad. Nxne of Ihc':c Ihrec ttni- 11anv, J. V. Elntivvir,n:Jemulimul f.xta.s vn,i inlm mak drvulolKrI Ihc C(iR during this inlllal rnnial xlfslimulati+mf In 11. 1!. 5h.tt ll'd.l, FJsINmI niemlNi..x n( Ibr b.nin: .F,k,MfN Iraiuiu~' phaer, :dlhnugh Iwu n( (he Ihrvc ralt infepm/i,e tyurnre Ilnuuan: Univer.'1'ean. Prrn, rcadil)' a+yuirtvl Iht• sll(:lReselnn haRCtu in 19rd1- -)o6oa35z52
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14R VN \'1'161t 1.1'I Infi INb 11H f. 19'1t IIMMr\.J.V,IhrxVN,J J,(,~\'YAIr,I11:,fSInV\Y, Ai.'1'hc e0n1 ,A irvxl aml , aN+.hprnnli,m nl•x inlrurun:d .rlf ainmblmn I „rxp phrvd. PTx6d., 145i, 60, 1}I I I:, ItT\In, J. V., 6( II1•NT• iI I'. .In Y\I"'rvlnYnlnl alrlu,rch In Ihr xnahua ul r nn/vxul Ivimveu. i lSyrin/,1955,40,.11.1 Jll ' Ilr.u:.uwr, J. M It. I'rnmmtl~ nl lrnid;urln/tmr l r.mlli kad elartrrd,' in Ibr Lrnu,. Idfr !. I1id. Ilyd., 1952, 21. 351 359. ilnwr, Iff f, & n.nnv, J. V. sn.,Y YRnI. nf eh,trn earlYUldvr Jorsk on n u+„•141i.ma1 r xaxmnl r li.wne ("Avir1Y "). J. nnup, ptrprnl. Paldna., I ros1.44, 88 V8, ,' Ilvrrt, 11. F., Jv.xnnsxc, I'., & Raenv, J. V. '1'Irc r0eetb n( rircmcrmvnidrr .hlck (Pf%1 „ a rnn. d'Yxrr.l rnYMlrrud uyxnlr: 'I'lla rRrrt d Lsb j Iil3 e.lilnti,nl Nn thc rral9namrnr nl llre re { 4aNwr. J. rnmpr plrytid. PryvGd.. In52, 46, .SAn IaLtv, L('.. Ilvrrlrrs, J. K., •\LVnan, P:. ('„ h (eLxIN, 'I'. \Y. Briri mru{njuduux elrctdc rne</„rm /m Himuiaimr ni the bnin. Sri.wae, 1955, 121, IGR. Slavrn, J. W. Rr.ImminR rh:rir fur IIN• eryN+inlenlnl almly ur pamnlra. J. ap/f. rl4yrid., 1450, 12, L91-133. Oun, ). A PrelinrinarY malMenR nl ekelricxl rcinhurinlq c0eeta in Ihe rat brain. J. [emp 'h7rid. 1'tychd., 1n56,49, 2E1-285. OHn, J. hfiaK. n( hangcr nnA nnM Lcs hn.mmne on x14Himnlai,m nf lhc hrnin. L rnw~. PhyTid. Pr/cFnf., 1v5tl, 61,320 324 (n) Lii ITo.. I 5nlamn. .R,.1. vm vi1.al,,.rJ.uv„ ••( IhY M1iSorx wl Uu,.. 1. erll.tlo,llixtn.r, „/ Ihrb.aiu. .Lhnrr. In:x, 12T, d15 12q,ir) tban, J.. h h/nr:r'x. I' 1'.~ilrvr I.iui,.u,+nrnt Pr.,. Jnu.l ln clrclnrnl .uumlalenl nl v.lnal a n,i .,uwt Ne,.n m Ihv ral I,rnin. J r~'xp r~i,v.~r.t. 141rpd, 1941, 47. 411) J27. Ih.ls, 1.. KN.rsY, K. P.. h KlrN.v Rlu. It Srll- .unmixrion a/ tlm Lrau. u...l a. a u urnwn rarr6,d hu Lanqm6iluq ,6ngv $rirmq In51y a24. 255. , 256 Revrnnls, K. 1\'.'1'In,rdntir,nehip Llnrcrntinndahm vnldar .nrJ ral r n/ h( Imlhalxmic .rif ai,nnlnl:,vr in 1 he ml. J. mxrp. plry.na !9 rcLd., 1'FA, 61. In5- Irnl. Knnears, \t'. R'.'tlxlh renmJinF nn.l pnoi•hine rRrae Irom Himulxtim. ni Ixnlrnor Ini•dLxlanm• ol rnl vilh R+me rlnlNxle nl . me /ntmHNl. J. r.m0, p4rrid. Pn<hd., IV58, 61. Am 1//] Slr.un, G. C. 6lultdvad Arcor.lc bnhlrn in Jrrnnir prqrrmirm.. 1n 11 li. tihm (1!d.L f9n/rirnl .nwfulix nJ lo. hrnin: S'dxmfirnf ixbnnnrc nYnrr.. 1lnuslnn: llurYCr.'lesn. 1'res., I'XII Sn)YAN, ,1Lr IIMIrT, J. V., {I„YIN, I. J, l IINa.\n, O, C. 8 Srlmwex, A. Ren.rA uhrdnin xuJ he. hav.N nrxiamined 6v intravnnixl r0 aimalrti•m .4ienrq PA5, 122. 8.tn Pl1. Ue..d.r,l hta, I. lumn 2060435253

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