Ukwahlukana kweziNxalenye zoMvuzo: Ukuthanda, Ukufuna, nokuFunda (2010)

Umvuzo: Izimvo – Eli qela linezifundo ezininzi kunye nophononongo oluvavanya i-neural substrates yokufuna vs ukuthanda. Ithiyori yangoku icebisa ukuba iindlela ze-dopamine ziyathanda kwaye iindlela ze-opioid ziyafuna. Umlutha ufuna kakhulu kangangokuba uqhubeka nokusebenzisa nangona ujongene neziphumo ezibi.

Isifundo esipheleleyo: Ukwahlula amalungu omvuzo: 'ukuthanda', 'ukufuna', nokufunda

Curr Opin Pharmacol. 2009 ngoFebruwari; 9(1): 65–73.

Ipapashwe kwi-intanethi ngo-2009 ngoJanuwari 21. doi: 10.1016/j.coph.2008.12.014.

UKent C Berridge, uTerry E Robinson, kunye noJ Wayne Aldridge

Idilesi yeSebe lezeNgqondo, kwiYunivesithi yaseMichigan, eAnn Arbor, 48109-1043, eU.SA

Umbhali ohambelanayo: Berridge, Kent C (I-imeyile: [imeyile ikhuselwe])

Abstract

Kwiminyaka yakutshanje kuye kwenziwa inkqubela phambili ebalulekileyo echaza amacandelo engqondo yomvuzo kunye neendlela zabo ezisisiseko ze-neural. Apha siqaqambisa ngokufutshane iziphumo kumacandelo amathathu engqondo anokudityaniswa nomvuzo: 'kuthanda' (impembelelo ye-hedonic),'efuna' (i-incentive salience), kunye nokufunda (unxulumano oluqikelelwayo kunye nokuqonda). Ukuqonda ngcono amacandelo omvuzo, kunye ne-neurobiological substrates, kunokunceda ekuyileni unyango oluphuculweyo lokuphazamiseka kweemvakalelo kunye nenkuthazo, ukusukela kuxinzelelo ukuya kukuphazamiseka kokutya, ukuba likhoboka leziyobisi, kunye nokunyanzeliswa kokusukelana nembuyekezo.

Yiya e:

intshayelelo

ukuyithanda

Kubantu abaninzi 'umvuzo' yinto enqwenelekayo kuba ivelisa amava olonwabo - kwaye ke eli gama linokusetyenziswa ukubhekisa kwizehlo zengqondo kunye ne-neurobiological ezivelisa ulonwabo. Kodwa ubungqina bubonisa ukuba ulonwabo lobuqu luyinto enye yomvuzo, kwaye loo mivuzo inokuphembelela isimilo naxa ungekho uyazi ngabo. Ewe, ukuzicingela ngamanye amaxesha kunokukhokelela ekubhidekeni malunga nokuba imivuzo ithandwa kangakanani na, ngelixa ukusabela kwangoko kunokuchaneka ngakumbi [1].

Okona kugqithisileyo, kwanokungaziva okanye okufihlakeleyo 'ukuthanda' ukuphendula kwi-hedonic stimuli kunokulinganiswa kwindlela yokuziphatha okanye kwi-physiology ngaphandle kweemvakalelo zolonwabo (umzekelo emva komboniso omfutshane wembonakalo yobuso eyonwabileyo okanye idosi ephantsi kakhulu yecocaine efakwa emithanjeni) [2,3]. Ngoko ke, nangona mhlawumbi iyamangalisa, imilinganiselo yenjongo 'yokuthanda' impendulo kwimivuzo ngamanye amaxesha inokubonelela ngokufikelela ngokuthe ngqo kwiinkqubo ze-hedonic kuneengxelo ezizimeleyo.

Eyona njongo iphambili ye-neuroscience kukuchonga ukuba yeyiphi i-substrates yobuchopho ebangela ulonwabo, nokuba yeyiphi na okanye injongo. Izifundo ze-Neuroimaging kunye ne-neural zokurekhodwa ziye zafumanisa ukuba imbuyekezo ukusuka kwincasa emnandi ukuya kwi-cocaine ene-intravenous, ukuphumelela imali okanye ubuso obuncumileyo kusebenze izakhiwo ezininzi zobuchopho, kubandakanya i-orbitofrontal cortex, i-anterior cingulate kunye ne-insula, kunye ne-subcortical izakhiwo ezinje nge-nucleus accumbens, i-ventral pallidum, i-ventral. tegmentum, kunye ne-mesolimbic dopamine projections, amygdala, njl.I-4,5,6,7 ••,8,I-9,I-10,11-13].

Kodwa yeyiphi kwezo nkqubo zobuchopho eneneni ebangela uyolo lomvuzo? Kwaye zeziphi iintshukumo endaweni ezinxibelelene nje (umzekelo ngenxa yokusasaza ukusebenza kwenethiwekhi) okanye iziphumo zolonwabo (ukulamla endaweni yoko eminye imisebenzi yokuqonda, yenkuthazo, yemoto, njl.njl. enxulumene nomvuzo)? Thina kunye nabanye siye safuna i-causation yolonwabo kwizifundo zezilwanyana ngokuchonga ukukhohlisa kwengqondo okwandisa impembelelo ye-hedonic [6,14 ••,15,16,I-17,18-22].

Ukufunda iinkqubo ze-neural ezinoxanduva lwempembelelo ye-hedonic yemivuzo, thina kunye nabanye siye sasebenzisa iinjongo 'zokuthanda' iimpendulo kwimivuzo yencasa eswiti, efana nembonakalo yobuso yeentsana ezisandul' ukuzalwa kunye nokusabela kobuso obufanayo beeorangutan, iichimpanzi, iinkawu, kunye nokuba iimpuku kunye neempuku [I-4,18,23,24]. Iilekese zivuselela inkangeleko 'yokuthanda' ebusweni kuzo zonke ezi (ukukhothwa imilebe, ukuvela kolwimi olunesingqisho, njl.njl.), kanti incasa ekrakrayo endaweni yoko ivuselela amabinzana 'angakuthandi' (izithuba, njl.); Umzobo 1; Imuvi eyongezelelweyo 1). Oko 'kuthanda'–'ukungathandi' ukusabela kulawulwa luluhlu lweenkqubo zobuchopho kwimpembelelo ye-hedonic kwi-forebrain kunye ne-brainstem, kwaye ziphenjelelwa zizinto ezininzi eziguqula ubumnandi, njengendlala / ukuhlutha kunye nokukhethwa kwencasa efundiweyo okanye ukuzonda.

Umzobo 1

Umzekelo wokuziphatha 'ukuthanda' reactions kunye nobuchopho hedonic hotspots ulonwabo luvo. Phezulu: Iimpendulo ezilungileyo ze-hedonic 'zokuthanda' zibangelwa yincasa yesucrose evela kusana olungabantu kunye nempuku yabantu abadala (umzekelo, ukuvela kolwimi olunesingqisho). ...

Ziinkqubo ezimbalwa ze-neurochemical ezifunyenweyo ukuza kuthi ga ngoku zokwandisa 'ukuthanda' ukusabela kwincasa eswiti kwiimpuku, kwaye kuphela ngaphakathi kweendawo ezimbalwa zobuchopho ezisokiweyo. Iinkqubo ze-opioid, i-endocannabinoid, kunye ne-GABA-benzodiazepine neurotransmitter zibalulekile ekuveliseni ukusabela okumnandi [14 ••,15,16,I-17,25,26], ngakumbi kwiindawo ezithile kwizakhiwo zemilenze (Umzobo 1 kwaye Umzobo 2) [15,16,I-17,21,27]. Ezi sayithi sizibize ngokuba 'ziindawo ze-hedonic' kuba ziyakwazi ukuvelisa ukwanda 'kokuthanda' iimpendulo, kunye nokuthelekelela, ulonwabo. Enye i-hedonic hotspot yokuphucula i-opioid yolonwabo lweemvakalelo ikwi-nucleus accumbens ngaphakathi kwe-rostrodorsal quadrant yeqokobhe layo eliphakathi, malunga ne-cubic millimeter ngomthamo [14 ••,15,28].

Oko kukuthi, i-hotspot iquka kuphela i-30% yomthamo weqokobhe eliphakathi, kwaye ngaphantsi kwe-10% ye-nucleus accumbens iyonke. Ngaphakathi kwe-hedonic hotspot, i-microinjection ye-opioid agonist, iDAMGO, iphinda kabini okanye kathathu inani leempendulo 'zokuthanda' ezibangelwa yincasa yesucrose [14 ••,28]. Enye i-hedonic hotspot ifumaneka kwisiqingatha esingasemva se-ventral pallidum, apho kwakhona i-DAMGO ikwandisa ngokunamandla ukusabela 'kokuthanda' kubumnandi [I-17,21,28]. Kuzo zombini iindawo ezishushu, le microinjection iphinda kabini 'ukufuna' ukutya ngengqiqo yokukhuthaza indlela yokutya kunye nokutya.

Umzobo 2

Ukwandiswa kwe-opioid hotspot kwi-nucleus accumbens ngokuchazwa kweendawo 'zokuthanda' ngokuchasene 'nokufuna' iindawo. Luhlaza: lonke iqokobhe eliphakathi lilamla ukwanda kwe-opioid-evuselelwe 'ekufuneni' umvuzo wokutya. ...

Ngaphandle kwezo ndawo zishushu, nakwisakhiwo esifanayo, ukuvuselela i-opioid kuvelisa iziphumo ezihluke kakhulu. Umzekelo, kwi-NAc phantse kuzo zonke ezinye iindawo ii-microinjections ze-DAMGO zisavuselela 'ukufuna' ukutya njengakwi-hotspot, kodwa azinyusi 'ukuthanda' (kwaye nokucinezela 'ukuthanda' kwindawo ebandayo esemva kwiqokobhe eliphakathi ngelixa. isakhuthaza ukutya; Umzobo 2). Ke, ukuthelekisa iziphumo zomsebenzi we-opioid ngaphakathi okanye ngaphandle kwe-hotspot kwi-NAc medial shell ibonisa ukuba iisayithi ze-opioid ezinoxanduva 'lokuthanda' ziyahlukana ngokwe-anatomically kwezo ziphembelela 'ukufuna' [14 ••,16].

I-Endocannabinoids iphucula 'ukuthanda' ukuphendula kwi-NAc hotspot engaphaya kwendawo ye-opioid [16,27]. I-Microinjection ye-anandamide kwi-endocannabinoid hotspot, esebenza mhlawumbi ngokuvuselela i-CB1 receptors apho, ngaphezu kokuphindaphinda inqanaba lokuphendula 'lokuthanda' kwincasa ye-sucrose (kunye nokutya okuphindwe kabini). Le substrate ye-hedonic endocannabinoid inokunxulumana neziphumo zeyeza labachasi be-endocannabinoid xa isetyenziswa njengonyango olunokubakho lokutyeba okanye umlutha [16,29,30].

I-ventral pallidum yinjongo ephambili yeziphumo ze-nucleus accumbens, kwaye isiqingatha sayo sangasemva siqulethe i-opioid hotspot yesibini.I-17,21]. Kwi-pallidum hotspot, ii-microinjections ze-DAMGO kabini 'ukuthanda' i-sucrose kunye 'nokufuna' ukutya (kulinganiswa njengokuthatha). Ngokwahlukileyo, i-microinjection ye-DAMGO ngaphambili kwi-hotspot icinezela 'ukuthanda' kunye 'nokufuna'. Ngokuzimeleyo, 'ukufuna' kuvuselelwe ngokwahlukeneyo kuzo zonke iindawo kwi-ventral pallidum ngokuvalwa kwe-GABA.A ii-receptors nge-bicuculline microinjection, ngaphandle kokuguqula 'ukuthanda' kuyo nayiphi na indawo [I-17,31].

Indima ye-ventral pallidum 'ekuthandeni' kunye 'nokufuna' iyenza ibe nomdla okhethekileyo kwizifundo zokuvula i-neural okubangelwa ngumvuzo. Ebantwini, i-cocaine, isondo, ukutya, okanye imali ivuza yonke into eyenza i-ventral pallidum isebenze, kubandakanywa ne-posterior subregion ehambelana ne-hedonic hotspot kwiimpuku [I-9,I-10,11,21]. Kuphononongo oluthe kratya lwe-electrophysiological lwendlela ii-neurons ezikwi-posterior ventral pallidum ezifaka ngayo imiqondiso ye-hedonic kwiimpuku, sifumanise ukuba i-hotspot neurons ivutha ngamandla kwincasa emnandi ye-sucrose kunencasa enetyuwa engathandekiyo (kathathu ukuxinana kwamanzi olwandle) [7 ••]. Nangona kunjalo, ngokwawo umahluko phakathi kokudubula kwe-sucrose kunye netyuwa akubonakalisi ukuba ii-neuron zifakela impembelelo ye-hedonic ('ukuthanda' ngokuchasene 'nokungathandi') kunokuba, yithi, yinto esisiseko yemvakalelo ye-stimulus (iswiti ngokuchasene netyuwa. ).

Nangona kunjalo, siphinde safumanisa ukuba umsebenzi we-neuronal ulandelele utshintsho kwixabiso le-hedonic le-hedonic yezi zivuseleli xa ubumnandi bencasa ye-NaCl bulawulwa ngokukhetha ngokubangela umdla wetyuwa womzimba. Xa iimpuku ziphelelwe yi-sodium (ngehomoni ye-mineralocorticoid kunye nolawulo lwe-diuretic), incasa yetyuwa enzulu yaba yindlela yokuziphatha 'ethandeka' njenge-sucrose, kwaye i-neurons kwi-ventral pallidum yaqala ukuvutha ngamandla kwityuwa njenge-sucrose [7 ••] (Umzobo 3). Sicinga ukuba uqwalaselo olunjalo lubonisa ukuba, eneneni, iipateni zokudubula zezi neurons ze-ventral pallidal encode 'ukuthanda' kwe-hedonic kumvakalelo omnandi, kunokuba iimpawu ezilula zokuva [21,32].

Umzobo 3

Ukhowudo lwe-Neuronal 'lokuthanda' kulonwabo lweemvakalelo zokungcamla okuswiti kunye netyuwa. Iimpendulo zokudubula kwe-Neuronal ziboniswa ukusuka kwi-ventral pallidum yokurekhoda i-electrode ukuya kwincasa ye-sucrose kunye netyuwa enzulu efakwe emlonyeni wempuku. Mbini ...

I-Hedonic hotspots ezisasazwa kwingqondo iphela zinokudityaniswa ngokudibeneyo kwisekethe edibeneyo ye-hierarchical edibanisa i-forebrain eninzi kunye ne-brainstem, ngokufana neziqithi ezininzi ze-archipelago ezirhweba kunye [21,24,27]. Kumgangatho ophakamileyo wezakhiwo ze-limbic kwi-ventral forebrain, uphuculo 'lokuthanda' zii-hotspots kwi-accumbens kunye ne-ventral pallidum inokusebenza kunye njengembumba enye yentsebenziswano, efuna 'iivoti' ngamxhelo mnye kuzo zombini iindawo ezitshisayo [28]. Ngokomzekelo, i-hedonic amplification yi-opioid stimulation ye-hotspot enye ingaphazanyiswa yi-opioid receptor blockade kwenye i-hotspot nangona 'ukufuna' ukukhulisa i-NAc hotspot kwakunamandla ngakumbi, kwaye yaqhubeka emva kwe-VP hotspot blockade [28].

Intsebenziswano efanayo ephantsi 'kokuthanda' ibonwe ngokulandela i-opioid kunye ne-benzodiazepine manipulations (mhlawumbi ibandakanya i-nucleus ye-parabrachial ye-brainstem pons) [27]. Uphuculo 'lokuthanda' oluveliswa lulawulo lwe-benzodiazepine lubonakala lufuna ukufunwa okusisinyanzelo kwee-opioids ezingapheliyo, kuba ithintelwa lulawulo lwe-naloxone [33]. Ke ngoko isekethe enye ye-hedonic inokudibanisa kunye iindlela ezininzi ze-neuroanatomical kunye ne-neuro-chemicals ukwenza 'ukuthanda' ukuphendula kunye nolonwabo.

'Uyafuna'

Ngokuqhelekileyo ingqondo 'ithanda' imivuzo 'eyifunayo'. Kodwa ngamanye amaxesha inokuba 'bafuna' nje. Uphando lufumanise ukuba imivuzo 'yokuthanda' kunye 'nokufuna' iyahlukana ngokwasengqondweni nakwi-neurobiologically. Ngokuthi 'ukufuna', sithetha u khuthazo, uhlobo lwenkuthazo yenkuthazo ekhuthaza ukusondela kunye nokusetyenziswa kwemivuzo, kwaye eneempawu ezahlukeneyo zengqondo kunye ne-neurobiological. Umzekelo, ukuqina kwenkuthazo kwahlulwa kwiindlela zokuqonda ngakumbi zomnqweno othethwa ligama eliqhelekileyo, ukufuna, elibandakanya iinjongo zokubhengeza okanye ulindelo olucacileyo lweziphumo zexesha elizayo, kwaye ubukhulu becala bulamlwa ziisekethe zecortical [34-37].

Ngokuthelekisa, inkuthazo yenkuthazo ilawulwa ziinkqubo ze-neural ezinobunzima obungaphantsi obubandakanya uqikelelo lwe-mesolimbic dopamine, ayifuni ulindelo olubanzi lwengqondo kwaye igxile ngokuthe ngqo kwi-stimuli enxulumene nomvuzo [34,35,38]. Kwiimeko ezinje ngokukhotyokiswa, okubandakanya ukukhuthazwa kwenkuthazo, umahluko phakathi kokunyanzeliswa kwenkuthazo kunye neminqweno yokuqonda ngakumbi ngamanye amaxesha unokukhokelela kwinto enokuthiwa 'ukufuna' okungenangqiqo: oko kukuthi, 'ukufuna' oko kungafunwayo ngengqondo, okubangelwa kukugqithisa. ukunyaniseka kwenkuthazo [I-39,I-40,41].

'Ukufuna' kunokufaka isicelo kwi-innate incentive stimuli (i-stimuli engenamiqathango, ii-UCSs) okanye ukufunda izivuseleli ebezikade zingathathi hlangothi kodwa ngoku zixela kwangaphambili ngokufumaneka komvuzo we-UCSs (i-Pavlovian conditioned stimuli, CSs) [38,I-40]. Oko kukuthi, ii-CSs zifumana iipropathi zenkuthazo xa i-CS idityaniswe ngokufumana umvuzo wokuzalwa okanye 'wendalo' ngokusebenzisa i-Pavlovian stimulus-stimulus associations (ukufunda kwe-S-S). Ukuqina kwenkuthazo kubalelwa kwezo CSs ngeendlela ze-limbic ezitsalela kulo mbutho ngexesha 'lokufuna', ukwenza i-CS ibe nomtsalane, kwaye inika amandla kunye nesikhokelo sokuziphatha okukhuthazwayo kumvuzo [35].

Xa i-CS ichazwa ngobuchule benkuthazo ngokuqhelekileyo ifumana iimpawu 'ezifunayo' ezahlukileyo nezinokumetwa [35,42], enokuthi iqalise xa i-CS iphinda iphinde yadibana (nangona umfanekiso ocacileyo weempawu zomvuzo unokwanela, ngakumbi ebantwini). Iipropathi 'ezifunayo' ezibangelwa zezi mpawu zomvuzo ziquka oku kulandelayo:

  1. Inkuthazo umazibuthe uphawu lwenkuthazo. I-CS ebizwa ngokuba yinkuthazo ebukhali iba ngumtsalane okhuthazayo, uhlobo 'lwemagnethi ekhuthazayo', esondelayo kwaye ngamanye amaxesha ide isetyenziswe (Imuvi eyongezelelweyo 1) [43,I-44,45]. Uphawu olukhuthazayo lwemagnethi lwe-CS lwenkuthazo lunokuba namandla kangangokuba i-CS isenokuvusa indlela yokunyanzeliswa [46]. Umzekelo, amakhoboka ecrack cocaine ngamanye amaxesha 'aleqa iziporho' okanye akrwele emva kweegranule ezimhlophe abawaziyo ukuba asiyococaine.
  2. Inqaku elithi 'ukufuna' kwe-US liqalise. Ukudibana ne-CS yomvuzo kwakhona kubangela 'ukufuna' kwe-UCS ehambelana nayo, mhlawumbi ngokugqithisela inkuthazo yenkuthazo ukumelwa ngokudibeneyo komvuzo ongekhoyo [34,47,48]. Kuvavanyo lwelabhoratri yezilwanyana, oku kubonakala njengencopho ye-phasic yokunyuka okubangelwe kukusetyenzwa komvuzo ongekhoyo (ikakhulu uvavanywa kuvavanyo olubizwa ngokuba yi-PIT okanye i-Pavlovian-Instrumental Transfer eqhutywa phantsi kweemeko zokuphela; Umzobo 4). 'Ukufuna' okubangelwa yi-cue kunokuchaneka ngokuthe ngqo kumvuzo ohambelanayo, okanye ngamanye amaxesha kuchitheke ngendlela eqhelekileyo yokukhuthaza 'ukufuna' kweminye imivuzo (njengokuba mhlawumbi xa abantu abakhobokileyo okanye abaguli be-dopamine-dysregulation bebonisa ukungcakaza okunyanzelekileyo, ngokwesondo. ukuziphatha, njl., ukongeza kwindlela yokuziphatha enyanzelekileyo yokusela iziyobisi) [49,50]. Ke ngoko, ukudibana nenkuthazo yenkuthazo kunokonyusa ngamandla inkuthazo yokufuna imbuyekezo, kunye nokwandisa amandla afunwa ngawo, into enokuthi ibaluleke ngakumbi xa i-cues iphinda iphinda ibuyele kubukhoboka.

    Umzobo 4

    I-NAc amphetamine yokukhulisa i-cue-triggered 'ukufuna.' Iincopho ezidlulayo 'zokufuna' umvuzo we-sucrose zibangelwa yimbonakalo ye-30-s ye-Pavlovian sucrose cue kuvavanyo lwePavlovian-Instrumental Transfer (CS +; ekunene). ...
  3. Uphawu lokuqinisa olunemeko. Ukuqina kwenkuthazo kwenza ukuba i-CS ibe nomtsalane kwaye 'ifuneke' ngengqiqo yokuba umntu uya kusebenza ukuze afumane i-CS ngokwayo, nangona ukungabikho komvuzo wase-US. Oku kuhlala kubizwa ngokuba yi-instrumental conditioned reinforcement. Ngokufanayo, ukongeza i-CS kwinto ezuzwayo xa isilwanyana sisebenzela umvuzo wase-US njenge-cocaine okanye i-nicotine, kwandisa indlela abasebenza ngokukhawuleza ngayo, mhlawumbi ngenxa yokuba i-CS yongeza i-target 'efunwayo' eyongezelelweyo [51]. Nangona kunjalo, siqaphela ukuba ukuqinisa okusemgangathweni kubanzi kunokuba 'ukufuna', kufuna iindlela ezongezelelweyo zokudibanisa ukufumana umsebenzi wesixhobo. Kwakhona, iindlela ezizezinye ze-SR zinokulamla uqinisekiso olunemeko kwiimeko ezithile ngaphandle kokubaluleka kwenkuthazo konke konke. Oku kwenza imagnethi yenkuthazo kunye neempawu 'ezifunayo' ezivuselelweyo zibaluleke ngakumbi ekuchongeni ubukhali benkuthazo egqithisileyo.

Ukongezwa kwenkuthazo

  1. Ukunyaniseka kwesenzo? Ngaphambi kokuba sishiye iimpawu zengqondo 'zokufuna', siyahendeka ukuba sicinge ukuba ezinye iindlela zokuziphatha. iintshukumo okanye iinkqubo zeemoto Isenokuthi 'ifuneke', iphantse ibe sisivuseleli senkuthazo, ngohlobo lwenkuthazo esetyenzisiweyo kwimifanekiso yobuchopho yeentshukumo zangaphakathi endaweni yokubonakalisa inkuthazo yangaphandle. Le ngcamango siyibiza ngokuba 'kukunyanzeliswa kwesenzo' okanye 'ukufuna' ukwenza. Ukunyanzeliswa kwesenzo sicebisa ukuba inokuba yimoto elingana nenkuthazo yenkuthazo, kwaye ilamlwe ziinkqubo zobuchopho ezijikelezayo (umzekelo, iinkqubo ze-dorsal nigrostriatal dopamine ezidibana ne-ventral mesolimbic). Isizukulwana seminqweno yokwenza, mhlawumbi ebandakanya imotor edibeneyo kunye nemisebenzi yenkuthazo ngaphakathi kwe-neostriatum (isakhiwo esaziwa ngokuba nenxaxheba kwintshukumo) sibonakala sihambelana neengcinga ezininzi ezivelayo malunga nomsebenzi we-basal ganglia [52,53,I-54,55].
  2. Ngaba umnqweno unxulumene noloyiko? Okokugqibela, siqaphela ukuba inkuthazo yenkuthazo inokuthi yabelane mhlawumbi ngokumangalisayo kwiindlela ze-mesocorticolimbic ezinoloyiko loloyiko [56,I-57,58,59]. Umzekelo, ukusebenzisana kwe-dopamine kunye ne-glutamate kwiisekethe ze-nucleus accumbens ukuvelisa kungekuphela nje umnqweno, kodwa kunye noloyiko, olucwangciswe ngokwe-anatom njengekhibhodi echaphazelekayo, apho ukuphazamiseka kwezitshixo ezilandelelanisiweyo zasekhaya kuvelisa imixube eyongeziweyo yokuthambekela ngokuchasene nokuziphatha okoyikekayo [I-57]. Ngaphaya koko, ezinye 'izitshixo' zasekhaya kwi-nucleus accumbens zinokujikwa ukusuka ekuveliseni enye inkuthazo iye kwelinye ngokutshintsha ngokwasengqondweni i-ambience yangaphandle echaphazelayo (umz. utshintsho ukusuka kwimeko-bume yekhaya etofotofo ukuya kweyoxinzelelo olukhanyiswe ngokuqaqambileyo nozaliswe ngumculo werock onomsindo) [56].
    Ezo ziphumo zakutsha nje zibonisa ukuba ubuchwephesha be-neurochemicals okanye i-anatomical localizations 'yokuthanda' okanye 'ukufuna' imisebenzi echazwe ngasentla isenokungabonakalisi ugxininiso lweendlela 'zelayini eneelebhile' ezinikezelwe ngokusisigxina apho 'i-substrate enye = umsebenzi omnye'. Endaweni yoko banokubonakalisa ubuchule obukhethekileyo bokuchaphazeleka (umz. iindawo ezishushu kakhulu) okanye inkuthazo-i-valence biases (umz. i-disk-dread keyboard) ye-neurobiological substrates zabo ezithile. Ezinye zezo substrates zinokuba nakho iindlela zokusebenza ezininzi, ngokuxhomekeke kwezinye izinto ngaxeshanye, ukuze zikwazi ukutshintsha phakathi kokuvelisa imisebenzi ngokuchasene nomnqweno ngokuchasene noloyiko.

I-Neurobiological substrates 'yokufuna'

Ukuthelekisa i-neurobiology 'yokufuna' 'ukuthanda', siqaphela ukuba iisubstrates zengqondo 'zokufuna' zisasazwa ngokubanzi kwaye zenziwe zisebenze ngokulula kune substrates 'zokuthanda' [38,53,60,I-61,62-65]. Iinkqubo ze-Neurochemical 'zokufuna' zininzi kwaye zihluke kuzo zombini i-neurochemical kunye ne-neuroanatomical domains, mhlawumbi isiseko sento 'yokufuna' umvuzo ngaphandle 'kokuthanda' ngokulinganayo umvuzo ofanayo. Ukongeza kwiinkqubo ze-opioid, ukusebenzisana kwe-dopamine kunye ne-dopamine kunye ne-corticolimbic glutamate kunye nezinye iinkqubo ze-neurochemical zivula inkuthazo 'yokufuna'. Ubuqhophololo be-Pharmacological kwezinye zezo nkqubo zinokutshintsha ngokulula 'ukufuna' ngaphandle kokutshintsha 'ukuthanda'. Umzekelo, ukucinezelwa kwe-endo native dopamine neurotransmission kunciphisa 'ukufuna' kodwa 'kungathandi' [38,64].

Ngokwahlukileyo, ukukhulisa 'ukufuna' ngaphandle 'kokuthanda' kuveliswe ngokusetyenziswa kweenkqubo ze-dopamine nge-amphetamine okanye iziyobisi ezifanayo ze-catecholamine ezinikwe ngokwenkqubo okanye i-microinjected ngqo kwi-nucleus accumbens, okanye ngokuguqulwa kofuzo okuphakamisa amanqanaba angaphandle kwe-dopamine Ukunkqonkqozwa kwabathuthi be-dopamine kwi-synapse) kwiisekethe ze-mesocorticolimbic, kunye nokuziva kufutshane okusisigxina kweenkqubo ezinxulumene ne-mesocorticolimbic-dopamine ngolawulo oluphindaphindiweyo lweedosi eziphezulu zeziyobisi (Umzobo 3-Umzobo 5) [I-39,I-40,I-61,66]. Sicebise ukuba kubantu abasesichengeni i-neural sensitization ye-incentive salience ngokusebenzisa iziyobisi zokuxhatshazwa kunokubangela isinyanzelo 'sokufuna' ukuthatha amachiza amaninzi, nokuba amachiza afanayo 'athandwa' ngokuhambelanayo, kwaye ngaloo ndlela abe negalelo kubukhoboka [I-39,I-40,42] (Umzobo 5).

Umzobo 5

Imodeli yenkuthazo yokulutha. Imodeli yeskim yendlela 'ukufuna' ukusebenzisa iziyobisi kunokukhula ngokuhamba kwexesha ngaphandle 'kokuthanda' ukonwabela iziyobisi njengoko umntu esiba likhoboka. Utshintsho kwiziyobisi eziqhelekileyo ...

Ukwahlula ukufunda 'ekufuneni': izinto ezixelwe kwangaphambili kunye nenkuthazo yeempawu ezinxulumene nomvuzo

Nje ukuba izikhokelo ezinxulumene nomvuzo zifundiwe, ezo mpawu zixela kwangaphambili imivuzo enxulumene noko kwaye ukongezelela zixhokonxa inkuthazo 'yokufuna' ukufumana imbuyekezo. Ngaba uqikelelo kunye 'nokufuna' kuyafana? Okanye ngaba zibandakanya iindlela ezahlukeneyo? Imbono yethu yeyokuba uqikelelo olufundiweyo kunye nobulumko benkuthazo bunokwahlulwa, kanye njengokuba 'ukuthanda' kunye 'nokufuna' kunokwenza [37,38,I-39,41,46,I-61]. Ukuhlaziya imisebenzi yengqondo kunye ne-neurobiological substrates kubalulekile kwiimodeli zovavanyo lokufunda umvuzo kunye nenkuthazo, kwaye kunefuthe kwi-pathologies, kubandakanya nokulutha. Siza kuchaza ngokufutshane imigca emithathu yobungqina obuvela kwiilabhoratri zethu ezicebisa ukuba iipropathi zokuqikelela kunye nenkuthazo yeempawu ezinxulumene nomvuzo ziyahlukaniseka.

Umzekelo wokuqala uvela kwiimvavanyo ezibonisa ukuba ii-CSs zinokufuna indlela - oko kukuthi, zisebenza 'njengezibuthe ezikhuthazayo', zitsalela umntu kuzo. Iimvavanyo ezininzi ziye zafumanisa ukuba xa uphawu okanye 'uphawu' (CS), olufana nokufakwa kwe-lever eludongeni, ludityaniswe nokuboniswa kwe-US enomvuzo, njengokutya, izilwanyana zikholisa ukusondela kwaye zibandakanye inqaku [43,I-44]. Isitshixo sokwahlula ingqikelelo kwimpembelelo silele ngokuyinxenye kubume bempendulo elungiselelwe umntu (CR) [43].

Ezinye iimpuku ziya kusondela kwi-lever ngakumbi nangakumbi ngokukhawuleza kwinkcazo nganye kwaye zize zibandakanye i-lever ngokufunxa, ukuyihlaba, kunye nokuyiluma - kubonakala ngathi izama 'ukutya' i-lever.Imuvi eyongezelelweyo 1) [45]. I-cue eqikelela umvuzo we-cocaine isondela ngokufanayo kwaye izibandakanya nepateni yayo yokuziphatha okuchulumancisayo [I-44], ezinokuthi zibe negalelo ekukwazini kweempawu ezinxulumene neziyobisi ukuba zingasebenzi kakuhle, zitsalele amakhoboka kuzo. Ezo CRs ezijoliswe kwi-CS ngokwayo zibizwa ngokuba 'kukulandela uphawu'.

Nangona kunjalo, ayizizo zonke iimpuku eziphuhlisa i-CR yokulandelela uphawu. Kwakwimeko efanayo yovavanyo ezinye iimpuku ziphuhlisa iCR eyahlukileyo - zifunda ukusondela 'kwinjongo' (itreyi yokutya), hayi i-lever, xa i-lever-CS iboniswa. Le CR ibizwa ngokuba 'kukulandela iinjongo'. Ke, ngamava abaxhasi benjongo beza kufikelela kwinjongo ngakumbi nangakumbi ngokukhawuleza kwinkcazo nganye ye-lever-CS, kwaye baqala ukubandakanya i-tray yokutya ngokukhawuleza, ukuyibetha, kwaye bayiluma [43,I-44,45]. Kuzo zonke iigundane, i-CS (ukufakwa kwe-lever) inokubaluleka okulinganayo okuqikelelwayo: ibangela zombini ii-CRs zokulandelela uphawu kunye ne-CRs yokulandelela injongo.

Umahluko kuphela apho iCR iqondiswe khona. Oku kuphakamisa ukuba kwii-sign-trackers i-lever-CS ichazwa njenge-incentive salience kuba kubo inomtsalane, kwaye oko kuxhaswa luqwalaselo lokuba ii-sign-trackers ngokukodwa nazo ziya kufunda ukwenza impendulo entsha ukufumana i-CS (okt instrumental conditioned). uqinisekiso) [46]. Kubaxhasi benjongo i-CS iqikelela ukutya, kwaye ikhokelela ekuphuhlisweni kwe-CR, kodwa i-CS ngokwayo ayibonakali ibonakaliswe ngenkuthazo yokukhuthaza ngezi ndlela (endaweni ukuba kukho nantoni na, injongo 'ifunwa') [43,46]. Ezo ziphumo zihambelana nesindululo sethu sokuba ukubikezelwa komvuzo okanye ixabiso lokudibanisa le-CS efundiweyo linokwahlulwa kwixabiso layo elikhuthazayo, kuxhomekeke ekubeni ngaba oko kuthatyathwe ngenkuthalo ngenxa yokuqina kwenkuthazo [46].

Umgca wesibini wobungqina bokucazulula uqikelelo oluvela kwizifundo 'zokufuna' iikhowudi ze-neural, ngakumbi emva kokusebenza kwengqondo okunxulumene ne-dopamine (nge-amphetamine okanye uvakaliso lwangaphambili). Ukuphakama kweDopamine kubonakala ngathi kuphucula ngokukodwa ukudubula kwe-neural ye-limbic kwiimpawu ezifaka i-incentive incentive salience (Umzobo 6) [I-61]. Ngokwahlukileyo, ukusebenza kwe-dopamine akuzange kuphucule imiqondiso ye-neural ebonisa ingqikelelo enkulu [I-61].

Umzobo 6

Ukwahlulwa kwexabiso lenkuthazo ye-CS (efunwayo) ukusuka kwixabiso le-CS eliqikelelwayo (ukufunda) ngokusebenza kwe-mesolimbic (eyenziwe kukwazisa okanye ulawulo olubukhali lwe-amphetamine). Olu hlalutyo lweprofayili yeepateni zokudubula kwe-neuronal kwi-ventral pallidum ibonisa utshintsho ...

Umgca wesithathu wobungqina uvela ekubuyiseleni umva 'ukufuna' kwe-CS ngelixa ubambe uqikelelo olufundiweyo olungaguqukiyo. Umzekelo, inqaku eliqikelela ubutyuwa obugqithisileyo lidla ngokuba 'lingafunwa' kodwa linokujikwa libe 'kufunwayo' xa kusenziwa umdla wokutya ityuwa ngokwasemzimbeni. Akukho sifundo sitsha, kwaye ke akukho tshintsho kuqikelelo olufundiweyo, ekufuneka lwenzeke ukuze ukuguqulwa kwenkuthazo kwenzeke. Ngaphaya koko, imeko yomdla engaqhelekanga ayizange ibonwe ngaphambili, kwaye i-CS ayifuni ukuba yayanyaniswa nencasa 'ethandwayo' ngaphambili. Nangona kunjalo, i-CS yangaphambili engalunganga ngequbuliso iba 'yifunwa' kwimeko entsha kwaye ikwazi ukuvuselela iipateni zokudubula eziqhelekileyo zokuqina kwenkuthazo. Kwizilingo zokuqala kwimeko yetyuwa, i-CS ngokukhawuleza ikhupha izibonakaliso zokudubula ze-neural ezifaka 'ukufuna' okulungileyo, nangaphambi kokuba i-UCS yetyuwa ibe incasa 'njengokuthandayo' [67]. Uqwalaselo olunjalo lubonisa ukuba ixabiso lentelekelelo ye-cue lahlukile kumandla ayo okufumana 'ukufuna', njengoko le yokugqibela ifuna ukuzibandakanya neenkqubo ze-neural ezongezelelweyo ukuvelisa amandla enkuthazo kunye nokubonisa 'ukufuna' kwithagethi yenkuthazo.

Uphando oluthe kratya luya kufuneka ukumisela ukuba 'ukufuna' xa kuthelekiswa nokufunda kunye noqikelelo kucalulwa njani na ngaphakathi kwengqondo. Nangona kunjalo, ubungqina ukuza kuthi ga ngoku bubonisa ukuba la macandelo anezazisi ezahlukeneyo zengqondo kunye ne-neural substrates ezahlulakalayo.

Yiya e:

isiphelo

Izifundo zeneuroscience ezisebenzayo 'zokuthanda', 'ukufuna', kunye namacandelo okufunda embuyekezo ziye zatyhila ukuba ezi nkqubo zesayikholoji zimephu kwiinkqubo zomvuzo zobuchopho ezahlukeneyo zeneuroanatomical kunye neurochemical ukuya kwinqanaba eliphawulweyo. Olu lwazi lunokukhokelela ekuqondeni ngcono indlela iinkqubo zobuchopho ezivelisa ngayo umvuzo oqhelekileyo, kunye nokungasebenzi kakuhle kweklinikhi yenkuthazo kunye neemvakalelo. Ezo zicelo zibandakanya ngakumbi indlela uvakalelo lweenkqubo ze-mesolimbic ezinokuvelisa ukunyanzeliswa kwembuyekezo kubukhoboka beziyobisi kunye nokuphazamiseka kwentshukumisa ehambelanayo ngokugqwetha ngokuthe ngqo 'ukufuna' umvuzo.

Yiya e:

Izinto ezongezelelweyo

Ividiyo ye-Hedonic 'yokuthanda'

Yiya e:

Imibulelo

Uphando ngababhali luxhaswe ngezibonelelo ezivela kwiZiko leSizwe lokuSetyenziswa kweZiyobisi kunye neZiko leSizwe leMpilo yeNgqondo (USA).

Yiya e:

Isihlomelo A. Idatha eyongezelelweyo

Idatha eyongezelelweyo ehambelana neli nqaku inokufumaneka, kwinguqulo ye-intanethi, kwi doi:10.1016/j.coph. 2008.12.014.

Yiya e:

Izalathiso kunye nokufunda okucetyiswayo

Amaphepha anika umdla, apapashwe kwisithuba sophononongo, aphawulwe njenge

• onomdla okhethekileyo

•• Inzala ebalaseleyo

1. Umfundi we-JW, uMauss IB. Ukonwaba nokuyazi: amava kunye nokwazisa ngemeta ngolonwabo. Ku: Kringelbach ML, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
2. Winkielman P, Berridge KC, Wilbarger JL. Iimpendulo zokungabikho zingqondweni kubuso obonwabileyo ngokuchasene nobuso obunomsindo ziphembelela indlela yokuziphatha yokusetyenziswa kunye nezigwebo zexabiso. Pers Soc Psychol Bull. 2005;31: 121-135. [PubMed]
3. Fischman MW, Foltin RW. Ukuzilawula ngokwakho kwe-cocaine ngabantu: imbono yelabhoratri. Ku: Bock GR, Whelan J, abahleli. I-Cocaine: Imilinganiselo yeNzululwazi kunye neNtlalo. ISimpoziyam yeSiseko seCIBA; Wiley; 1992. iphepha 165–180.
4. I-Kringelbach ML I-cortex yomntu ye-orbitofrontal: idibanisa umvuzo kumava e-hedonic. Nat Rev Neurosci. 2005;6: 691-702. [PubMed]Ngokucacileyo nangokucacileyo ichaza indima ye-orbitofrontal cortex kulonwabo ebantwini.
5. Leknes S, Tracey I. I-neurobiology eqhelekileyo yeentlungu kunye nokuzonwabisa. Nat Rev Neurosci. 2008;9: 314-320. [PubMed]
6. I-Wheeler RA, i-Carelli RM. I-neuroscience yolonwabo: gxila kwi-ventral pallidum yokudubula iikhowudi umvuzo we-hedonic: xa incasa embi ijika ilungile. J Neurophysiol. 2006;96: 2175-2176. [PubMed]
7. Tindell AJ, Smith KS, Pecina S, Berridge KC, Aldridge JW Ventral pallidum ukudubula iikhowudi umvuzo we-hedonic: xa incasa embi ijika ilungile. J Neurophysiol. 2006;96: 2399-2409. [PubMed]Olu phononongo lubonelela ngobungqina bekhowudi ye-neuronal 'yokuthanda' njengenxalenye yenjongo yolonwabo lomvuzo ngokusebenzisa iipateni zokudubula ze-neuronal kwi-ventral pallidum ukuya kwincasa ye-sucrose kunye netyuwa.
8. I-Knutson B, i-Wimmer GE, i-Kuhnen CM, i-Winkielman P. I-Nucleus accumbens activation idibanisa impembelelo yeempawu zomvuzo ekuthatheni ingozi yemali. Neuroreport. 2008;19: 509-513. [PubMed]
9. Beaver JD, Lawrence AD, van Ditzhuijzen J, Davis MH, Woods A, Calder AJ Umahluko womntu ngamnye kwi-reward drive uqikelele iimpendulo ze-neural kwimifanekiso yokutya. J Neurosci. 2006;26: 5160-5166. [PubMed]Ibonisa ukuba iisekethe zenkuthazo zenziwe zisebenze ngomvuzo wokutya ebantwini ngeendlela ezinxibelelene neempawu zobuntu (BAS) ezinokunxulumana nokufuna imvakalelo.
10. Pessiglione M, Schmidt L, Draganski B, Kalisch R, Lau H, Dolan R, Frith C Indlela ubuchopho buguqulela ngayo imali ngamandla: isifundo se-neuroimaging ye-subliminal motivation. Sayensi. 2007;316: 904-906. [PubMed]Ibonisa ebantwini ukuba iisekethe zenkuthazo yobuchopho ezibandakanya i-ventral pallidum zenziwe zisebenze nangomvuzo ocacileyo ohlala ungaphantsi kokuqonda, kwaye ziyakwazi ukukhulisa isenzo esikhuthazayo somvuzo.
11. Childress AR, Ehrman RN, Wang Z, Li Y, Sciortino N, Hakun J, Jens W, Suh J, Listerud J, Marquez K, et al. Ingabula-zigcawu kwinkanuko: ukwenza i-limbic isebenze ngokusetyenziswa kweziyobisi 'ezingabonakaliyo' kunye neempawu zesondo. PLoS ONE. 2008;3: e1506. [Inkcazelo yamahhala ye-PMC] [PubMed]
12. I-DM encinci, i-Veldhuizen MG, i-Felsted J, i-Mak YE, i-McGlone F. Ii-substrates ezihlukeneyo zokulindela kunye ne-consummatory food chemosensation. Neuron. 2008;57: 786-797. [Inkcazelo yamahhala ye-PMC] [PubMed]
13. I-Tobler P, i-O'Doherty JP, i-Dolan RJ, i-Schultz W. Ikhowudi yexabiso lomvuzo eyahlukileyo kumngcipheko wesimo sengqondo esinxulumene nokungaqiniseki kweekhowudi kwiinkqubo zomvuzo womntu. J Neurophysiol. 2007;97: 1621-1632. [Inkcazelo yamahhala ye-PMC] [PubMed]
14. Peciña S, Berridge KC I-Hedonic indawo eshushu kwi-nucleus accumbens iqokobhe: Ingaba i-mu-opioids ibangela phi ukonyuka kwempembelelo ye-hedonic yobumnandi? J Neurosci. 2005;25: 11777-11786. [PubMed]Ichonga i-cubic-millimeter 'hedonic hotspot' kwiqokobhe lenucleus, apho iisignali ze-opioid zibangela 'ukuthanda' ukomelezwa kweemvakalelo zolonwabo lwencasa eswiti. Olu phononongo lukwabonelele ngobungqina bokuqala bokwahlulwa kwe-atomical 'yokuthanda' i-opioid ukusuka kwindawo esulungekileyo 'efunwayo' kunye ne-coldspot ngaphandle kwe-hotspot.
15. U-Peciña S, uSmith KS, uBerridge KC. Amabala ashushu eHedonic kwingqondo. Neuroscientist. 2006;12: 500-511. [PubMed]
16. Mahler SV, Smith KS, Berridge KC. I-Endocannabinoid hedonic hotspot yolonwabo lweemvakalelo: i-anandamide kwi-nucleus accumbens iqokobhe iphucula 'ukuthanda' komvuzo omnandi. Neuropsychopharmacology. 2007;32: 2267-2278. [PubMed]
17. Smith KS, Berridge KC I-ventral pallidum kunye nomvuzo we-hedonic: iimephu ze-neurochemical ze-sucrose "ukuthanda" kunye nokutya. J Neurosci. 2005;25: 8637-8649. [PubMed]Olu phononongo lubonise ukuba i-ventral pallidum iqulethe i-cubic-millimeter 'hedonic hotspot' kwi-ventral pallidum yokukhulisa i-opioid 'yokuthanda' iimpendulo kubumnandi, obubekwe kwindawo yangasemva.
18. UBerridge KC, uKringelbach ML. I-neuroscience esebenzayo yolonwabo: umvuzo ebantwini nakwizilwanyana. Psychopharmacology (Berl) 2008;199: 457-480. [Inkcazelo yamahhala ye-PMC] [PubMed]
19. Pecina S. Umvuzo we-opioid 'ukuthanda' kunye 'nokufuna' kwi-nucleus accumbens. Physiol Behav. 2008;94: 675-680. [PubMed]
20. I-Kringelbach ML. Ingqondo ye-hedonic: i-neuroanatomy esebenzayo yolonwabo lomntu. Ku: Kringelbach ML, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
21. USmith KS, Tindell AJ, Aldridge JW, Berridge KC. Iindima ze-ventral pallidum kwimivuzo kunye nenkuthazo. Behav Brain Res. 2009;196: 155-167. [Inkcazelo yamahhala ye-PMC] [PubMed]
22. I-Ikemoto S. Dopamine umvuzo wokujikeleza: iinkqubo ezimbini zokubonisa ukusuka kwi-ventral midbrain ukuya kwi-nucleus accumbens-olfactory tubercle complex. Brain Res Rev. 2007;56: 27-78. [Inkcazelo yamahhala ye-PMC] [PubMed]
23. Steiner JE, Glaser D, Hawilo ME, Berridge KC. Ukuthelekisa ukubonakaliswa kwempembelelo ye-hedonic: ukusabela okuchaphazelayo kwincasa yeentsana ezingabantu kunye nezinye iiprimates. Neurosci Biobehav Rev. 2001;25: 53-74. [PubMed]
24. Grill HJ, Norgren R. Uvavanyo lwe-taste reactivity test. II. Iimpendulo zeMimetic kwi-gustatory stimuli kwi-thalamic engapheliyo kunye ne-chronic decerebrate rats. Res Resin. 1978;143: 281-297. [PubMed]
25. Jarrett MM, Limebeer CL, Parker LA. Impembelelo ye-Delta9-tetrahydrocannabinol kwi-sucrose palatability njengoko ilinganiswe ngovavanyo lwe-taste reactivity. Physiol Behav. 2005;86: 475-479. [PubMed]
26. UZheng H, Berthoud HR. Ukutya ulonwabo okanye iikhalori. Curr Opin Pharmacol. 2007;7: 607-612. [Inkcazelo yamahhala ye-PMC] [PubMed]
27. USmith KS, Mahler SV, Pecina S, Berridge KC. I-Hedonic hotspots: ivelisa ulonwabo kwingqondo. Ku: Kringelbach M, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
28. Smith KS, Berridge KC. Isekethe ye-Opioid limbic yomvuzo: intsebenziswano phakathi kwe-hedonic hotspots ye-nucleus accumbens kunye ne-ventral pallidum. J Neurosci. 2007;27: 1594-1605. [PubMed]
29. USolinas M, Goldberg SR, Piomelli D. Inkqubo ye-endocannabinoid kwiinkqubo zokuvuza kwengqondo. Br J Pharmacol. 2008;154: 369-383. [Inkcazelo yamahhala ye-PMC] [PubMed]
30. Kirkham T. Endocannabinoids kunye neurochemistry of gluttony. J Neuroendocrinol. 2008;20: 1099-1100. [PubMed]
31. UShimura T, u-Imaoka H, ​​u-Yamamoto T. Ukuguqulwa kwe-Neurochemical yokuziphatha kokungena kwi-ventral pallidum. Eur J Neurosci. 2006;23: 1596-1604. [PubMed]
32. Aldridge JW, Berridge KC. Ikhowudi ye-Neural yolonwabo: "Iiglasi zeRose-Tinted" ze-ventral pallidum. Ku: Kringelbach ML, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
33. Richardson DK, Reynolds SM, Cooper SJ, Berridge KC. I-endogenous opioids iyafuneka ekwandiseni ukukholeka kwe-benzodiazepine: iibhloko ze-naltrexone zithintela ukwanda okubangelwa yi-diazepam kwe-sucrose-'ukuthanda'. Pharmacol Biochem Behav. 2005;81: 657-663. [PubMed]
34. UDickinson A, uBalleine B. Hedonics: i-cognitive-motivational interface. Ku: Kringelbach ML, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
35. Berridge KC. Ukufunda ngembuyekezo: ukuqiniswa, inkuthazo, kunye nolindelo. Ku: Medin DL, umhleli. IPsychology yokuFunda kunye neNkuthazo. Umqulu 40. I-Academic Press; 2001. iphepha 223-278.
36. Daw ND, Niv Y, Dayan P. Ukhuphiswano olusekelwe ekungaqiniseki phakathi kweenkqubo zangaphambili kunye ne-dorsolateral striatal zokulawula ukuziphatha. Nat Neurosci. 2005;8: 1704-1711. [PubMed]
37. Dayan P, Balleine BW. Umvuzo, inkuthazo, kunye nokomeleza ukufunda. Neuron. 2002;36: 285-298. [PubMed]
38. Berridge KC. Ingxoxo ngendima ye-dopamine kumvuzo: imeko yenkuthazo. Psychopharmacology (Berl) 2007;191: 391-431. [PubMed]
39. Robinson TE, Berridge KC Ithiyori yenkuthazo yokulutha: eminye imiba yangoku. I-Philos Trans Trans R Soc Lond B Biol Sci. 2008;363: 3137-3146. [PubMed]Uhlaziyo lwamva nje kubungqina obumalunga nethiyori yokuba ukukhotyokiswa bubangelwa yinxenye yeziyobisi kwi-neural substrates 'yokufuna'.
40. Robinson TE, Berridge KC Addiction. I-Annu Rev Psychol. 2003;54: 25-53. [PubMed]Uthelekisa uluvo lokuba umlutha ubangelwa kukukhuthaza-ukuvuselela, kunye nokufunda okanye i-hypothesis yomkhwa kunye nokurhoxa okanye i-hedonic yomchasi we-hypotheses yokulutha.
41. Berridge KC, Aldridge JW. Usetyenziso lwesigqibo, ingqondo, kunye nokusukela iinjongo ze-hedonic. Soc Cognition. 2008;26: 621-646.
42. Robinson TE, Berridge KC. Isiseko se-neural sokunqwenela iziyobisi: ithiyori yokukhuthaza ukubonwa iziyobisi. Brain Res Rev. 1993;18: 247-291. [PubMed]
43. UFlagel SB, Akil H, Robinson TE. Ukwahluka komntu ngamnye kubume bokuchaphazeleka kweesisikhuseli kwiimpawu ezinxulumene nomvuzo: iimpembelelo zokulutha. Neuropharmacology. 2009;56: 139-148. [Inkcazelo yamahhala ye-PMC] [PubMed]
44. Uslaner JM, Acerbo MJ, Jones SA, Robinson TE Ukunikezelwa kwenkuthazo ebukhali kwisivuseleli esibonisa inaliti ye-cocaine. Behav Brain Res. 2006;169: 320-324. [PubMed]Ubonisa okokuqala kwimodeli yezilwanyana ebonisa iziyobisi ezifana ne-cocaine ithatha iipropathi 'zemagnethi ekhuthazayo', ukuze i-cues ifune indlela kunye nophando oluchulumancisayo kwi-autoshaping paradigm.
45. UMahler S, Berridge K. Amygdala iindlela zokukhuthaza. Umbutho weeNgcaciso zeNeuroscience. 2007
46. ​​Robinson TE, Flagel SB. Ukwahlulahlula izinto eziqikelelwayo kunye nenkuthazo yeempawu ezinxulumene nomvuzo ngokufunda umahluko ngamnye. Biol Psychiatry. 2008 doi: 10.1016 / j.biopsych.2008.09.006.
47. Wyvell CL, Berridge KC. I-Intra-accumbens amphetamine yonyusa umgangatho wenkuthazo omiselweyo womvuzo we-sucrose: ukongezwa komvuzo "ukufuna" ngaphandle "kokuthanda" okuphuculweyo okanye ukuqinisa impendulo. J Neurosci. 2000;20: 8122-8130. [PubMed]
48. Holland PC. Ubudlelwane phakathi kwePavlovian-instrumental transfer kunye ne-reinforcer devaluation. J Exp Psychol-Anim Behav Inkqubo. 2004;30: 104-117. [PubMed]
49. Evans AH, Pavese N, Lawrence AD, Tai YF, Appel S, Doder M, Brooks DJ, Lees AJ, Piccini P. Ukusetyenziswa kweziyobisi okunyanzelekileyo okudityaniswe nokudluliselwa kwe-dopamine ye-ventral striatal dopamine. Ann Neurol. 2006;59: 852-858. [PubMed]
50. Kausch O. Iipateni zokusetyenziswa gwenxa kweziyobisi phakathi abangcakaza unyango-abafuna unyango. J Unyango olusebenzisa gadalala. 2003;25: 263-270. [PubMed]
51. I-Schenk S, i-Partridge B. Impembelelo yokukhanya okusemgangathweni kwi-cocaine yokuzilawula ngokwakho kwiimpuku. Psychopharmacology (Berl) 2001;154: 390-396. [PubMed]
52. U-Aldridge JW, uBerridge KC, uHerman M, uZimmer L. Ikhowudi ye-Neuronal ye-serial order: i-syntax yokulungiswa kwi-neostriatum. Psychol Sci. 1993;4: 391-395.
53. Volkow ND, Wang GJ, Telang F, Fowler JS, Logan J, Childress AR, Jayne M, Ma Y, Wong C. I-Cocaine cues kunye ne-dopamine kwi-dorsal striatum: indlela yokunqwenela kwi-cocaine addiction. J Neurosci. 2006;26: 6583-6588. [PubMed]
54. I-Everitt BJ, i-Belin D, i-Economidou D, i-Pelloux Y, i-Dalley JW, i-Robbins TW Iinkqubo ze-Neural eziphantsi komngcipheko wokuphuhlisa imikhwa enyanzelekileyo yokufuna iziyobisi kunye nokukhobokisa. I-Philos Trans Trans R Soc Lond B Biol Sci. 2008;363: 3125-3135. [PubMed]Ngokucokisekileyo ubonisa umbono oxhasa umbono wokuba umlutha uvela kwimikhwa ye-SR ebaxiweyo ngenxa yokugqwethwa kwecandelo lokufunda lomvuzo.
55. Haber SN, Fudge JL, McFarland NR. Iindlela zeStriatonigrostriatal kwiiprimates zenza i-spiral enyukayo ukusuka kwiqokobhe ukuya kwi-dorsolateral striatum. J Neurosci. 2000;20: 2369-2382. [PubMed]
56. Reynolds SM, Berridge KC. Imo engqongileyo yeemvakalelo ibuyisela i-valence yolangazelelo ngokuchasene nemisebenzi eyoyikisayo kwi-nucleus accumbens. Nat Neurosci. 2008;11: 423-425. [Inkcazelo yamahhala ye-PMC] [PubMed]
57. UFaure A, uReynolds SM, uRichard JM, uBerridge KC iMesolimbic dopamine kumnqweno kunye noloyiko: ukwenza inkuthazo ukuba iveliswe kukuphazamiseka kwe-glutamate yendawo kwi-nucleus accumbens. J Neurosci. 2008;28: 7184-7192. [PubMed]Olu vavanyo lubonisa okokuqala ukuba i-dopamine ivelisa zombini inkuthazo ekhuthazayo kunye nenkuthazo engalunganga ngokusebenza kunye neempawu ze-corticolimbic glutamate ngendlela ethile ye-anatomically ngaphakathi kwe-nucleus accumbens.
58. Levita L, Dalley JW, Robbins TW. I-Nucleus iqokelela i-dopamine kwaye yafunda uloyiko oluphinde lwajongwa: uphononongo kunye neziphumo ezintsha. Behav Brain Res. 2002;137: 115-127. [PubMed]
59. I-Kapur S. Ii-antipsychotics ziba yi-anti-'psychotic' - ukusuka kwi-dopamine ukuya kwi-salience ukuya kwi-psychosis. Iintlobo zeP Pharmacol Sci. 2004;25: 402-406. [PubMed]
60. UAragona BJ, uCarelli RM. I-Dynamic neuroplasticity kunye ne-automation yokuziphatha okukhuthazwayo. Funda iMem. 2006;13: 558-559. [PubMed]
61. Tindell AJ, Berridge KC, Zhang J, Peciña S, Aldridge JW Ventral pallidal neurons ikhowudi yokukhuthaza inkuthazo: ukukhulisa nge-mesolimbic sensitization kunye ne-amphetamine. Eur J Neurosci. 2005;22: 2617-2634. [PubMed]Umboniso wokuqala we-neural coding wokuba i-dopamine kunye novuselelo lukhulisa imiqondiso 'efunwayo', ezimeleyo 'ekuthandeni' okanye kwizinto zokufunda zomvuzo.
62. USmith KS, Berridge KC, Aldridge JW. I-Ventral pallidal neurons yahlula 'ukuthanda' kunye 'nokufuna' ukuphakama okubangelwa yi-opioids ngokuchasene ne-dopamine kwi-nucleus accumbens. KuMbutho weeNgcaciso zeNeuroscience. 2007
63. I-Abler B, i-Erk S, i-Walter H. Ukuqaliswa kwenkqubo yomvuzo womntu imodareyithwa yi-dose enye ye-olanzapine kwizifundo eziphilileyo kwi-fMRI enxulumene nesiganeko, i-double-blind, i-placebo-controlled controlled fMRI. Psychopharmacology (Berl) 2007;191: 823-833. [PubMed]
64. Leyton M. I-neurobiology yomnqweno: i-dopamine kunye nokulawulwa kwemizwelo kunye namazwe akhuthazayo kubantu. Ku: Kringelbach ML, Berridge KC, abahleli. Ubumnandi beNgqondo. I-Oxford University Press; kushicilelo.
65. ISalamone JD, Correa M, Mingote SM, Weber SM. Ngaphandle kwe-hypothesis yomvuzo: enye imisebenzi ye-nucleus accumbens dopamine. Curr Opin Pharmacol. 2005;5: 34-41. [PubMed]
66. I-Peciña S, i-Cagniard B, i-Berridge KC, i-Aldridge JW, i-Zhuang X. Iigundane eziguquguqukayo ze-hyperdopaminergic zinezinga eliphezulu "elifunayo" kodwa "lingathandi" imivuzo emnandi. J Neurosci. 2003;23: 9395-9402. [PubMed]
67. Tindell AJ, Smith KS, Berridge KC, Aldridge JW. I-Ventral pallidal neurons idibanisa ukufunda kunye nemiqondiso ye-physiological kwikhowudi yenkuthazo yokubaluleka kwezinto ezinemeko; Umbutho weNkomfa yeNeuroscience; ngoNovemba 12, 2005; EWashington, DC. 2005.