Umzila ongathintekiyo wezindlela ezibhebhethekisa izidakamizwa kanye nokukhuluphala: Ukungafani nokufana (2013) Nora Volkow

Dardo Tomasi^*,1 futhi Nora D. Volkow^1,2

Amasu we-Neuroimaging aqala ukwembula ukugqama okukhulu emgudwini wobuchopho obangelwa umlutha nokuphazamiseka kwe-dyscontrol ngenxa yokuziphatha okunomvuzo (njengokuphazamiseka kokudla nokukhuluphala). I-Positron emission tomography (PET) ikhombise ukusayina okhubazekile i-dopamine (DA) (yehle i-D2 receptors) ekuluthweni kwezidakamizwa nasekuseni okuhambisana nokuncipha kwesisekelo se-glucose metabolism ezindaweni zobuchopho bezendondo nezendlala. I-imagination ye-magnetic resonance imaging (fMRI) esebenzayo ibike nezimpawu zobuchopho bokusebenzisa ubuchopho futhi obufaka emendweni we-DA-modified striato-cortical pathways. Kulesi sibuyekezo sithola imephu yokuthola okuvela ocwaningweni lwakamuva lwe-neuroimaging olwehlukanisa ukusebenza kobuchopho kumlutha wezidakamizwa / wokudla kulabo abalawula ngaphakathi kwamanethiwekhi wobuchopho asebenzisana ne-ventral kanye ne-dorsal striatum. Sikhombisa ukuthi izifunda ezitholakele njengezijwayelekile ngokulutha kanye nokukhuluphala ngokweqile zivame ukuvela lapho kugqashuka khona indawo yenethiwekhi kanye nenethiwekhi ye-ventral striatal. Izindawo zedical tempial kanye nezindawo eziphakeme zangaphambili ezixhumene ne-dorsal striatum zibonisa ubungozi obukhulu ekukhuluphelweni kanye nasezinkingeni zokudla kunasekuluthweni kwezidakamizwa, okubonisa ukwanda kwezinkinga zokukhuluphala kanye nokuphazamiseka kokudla kunokuba umlutha. Lokhu kulawula ukubandakanyeka kwalezi zinto zombili zangaphakathi (ezihlotshaniswa kakhulu nomvuzo kanye nogqozi) namanethiwekhi we-dorsal striatal (ahambisana nemikhuba noma ukuvuselela impendulo yokufunda) ngomlutha nokukhuluphala kepha futhi kukhomba amaphethini ahlukile phakathi kwalokhu kuphazamiseka okubili.

Izimpendulo ze-Dopaminergic ezidakamizweni nasekudleni

Zonke izidakamizwa eziluthayo zibonisa amandla okwandisa i-DA ku-striatum, ikakhulukazi kuma-nucleus accumbens (i-ventral striatum), ebangela imiphumela yabo emihle (I-Koob, i-1992). Ama-neurons e-DA atholakala endaweni ye-ventral tegmental endaweni (VTA) ne -antiantia nigra (SN) kuphrojekthi ye-midbrain eya e-striatum nge-mesolimbic kanye ne-nigrostriatal pathways. Imiphumela evuzayo neyengeni yezidakamizwa (futhi kungenzeka ukuthi nayo idle) ibonakala iqhutshwa ikakhulu wukungapheli kanye nokwanda kokuqhuma kokuxoshwa kwamangqamuzana e-DA (Ohlakaniphile, 2009) okuholela ekugxileni okuphezulu kwe-DA okudingekayo ukuvusa ama-receptors ama-D1 aphansi kakhulu (UZweifel et al., 2009). Ebantwini, izifundo ze-PET zikhombisile ukuthi izidakamizwa ezimbalwa zengeza i-DA kuma-dorsal kanye ne-ventral striatum nokuthi lokhu kukhuphuka kuhlotshaniswa nemiphumela emihle yokuvuza yemithi [izivuseleli (I-Drevets et al., I-2001;I-Volkow et al., I-1999), i-nicotine (U-Brody et al., 2009), utshwala (Boileau et al., 2003) kanye ne-cannabis (UBongong et al., 2009)]. Izimpendulo ze-Dopaminergic nazo zingadlala indima emiphumeleni eyinzuzo yokudla futhi zinikele ekusetshenzisweni ngokweqile kanye nokukhuluphala (I-Volkow et al., 2011b). Ukudla okuthile, ikakhulukazi lezo ezinoshukela namafutha, kunomvuzo onamandla futhi kungakukhuthaza ukudla ngokweqile (U-Lenoir et al., 2007) ngoba njengezidakamizwa bakhulisa ukukhishwa kwe-striatal DAUNorgren et al., 2006). Ngaphezu kwalokho, ukudla kungakhulisa i-DA ku-ventral striatum kuphela ngesisekelo sokuqukethwe kwayo kwe-caloric kanye nokuzimela kokuqina (de Araujo et al., 2008). Ngenkathi izinhlangano zemivuzo yokudla zazinenzuzo ezindaweni lapho imithombo yokudla yayingatholakali futhi / noma ingathembekile, le ndlela manje isiyisibopho emiphakathini yethu yanamuhla lapho ukudla kuningi futhi kutholakala njalo.

Amanye ama-neurotransmitters kune-dopamine (cannabinoids, opioids, ne-serotonin) kanye nama-hormone e-neuropeptide (i-insulin, i-leptin, i-ghrelin, i-orexin, i-glucagon efana ne-peptide, iprotheni ehlobene ne-agouti, i-PYY) kube nomthelela emiphumeleni yokuvuza yokudla nakwimithetho ye ukudlaAtkinson, 2008;UCason et al., 2010;UCota et al., 2006). Ngaphezu kwalokho, ukunyuka okuhlobene nokudla okuhlobene nokudla kwe-DA kukodwa ngeke kuchaze umehluko phakathi kokudla okujwayelekile nokusetshenziswa kokudla okuphoqelela okweqile ngoba lokhu kwenzeka nakubantu abaphilile abangadli ngokweqile. Ngakho-ke, ngokulutha, ukuzivumelanisa nokwehla komgwaqo kungenzeka kubandakanyeke ekulahlekelweni kokulawula kokudla kokudla. Lokhu kube nge-neuroadaptations kungahle kwehle ekunciphiseni kokudubula kwamaseli e-DA kwamathiseli, ukudubula okwenziwe ngamaselula kwe-DA ngokuphendula izidakamizwa noma izinkomba zokudla kanye nokuncishiswa komsebenzi wokuphatha kubandakanya ukungasebenzi kahle kokuzibamba (Umusa, i-2000;Wanat et al., 2009).

Ukuxhumana kwe-Striatocortical

Ukuxhumeka kwe-cortical deficits ye-striatal dopaminergic ngaphezu kwalokho akulindelekile. Ucwaningo lwe-anatomical ezilimini ezingezona ezabantu nokwakusetshenziselwa ukubhala ukuthi le moto, somatosensory and dorsolateral prewardal cortices projekti to dorsal striatum (I-Künzle, 1975;I-Künzle, 1977;USelemon noGoldman-Rakic, 1985;Middleton noStrick, 2002;UKelly noStrick, 2004;U-Künzle no-Akert, 1977), nokuthi iphrojekthi ye-anterior cingulate (ACC) ne-orbitof Pambal (OFC) yamaphothi we-cortices ukuze i-ventral striatum (Ilinsky et al., 1985;USelemon noGoldman-Rakic, 1985;UFerry et al., 2000;Haber et al., 2006;UPowell noLeman, 1976;I-Yeterian neVan Hoesen, 1978).

Muva nje, uDi Martino nozakwabo bakwazile ukuphindaphinda lezi zijikelezo ze-striatocortical besebenzisa amafushane (<7 min) amaseshini wokuskena we-MRI ekuphumuleni ezifundweni zabantu ezingama-35 (UDi Martino et al., 2008) futhi yeseka ukucubungula kwe-meta kwezifundo ze-PET ne-fMRI ehlonze ukuxhumana okusebenzayo phakathi kwe-anterior dorsal striatum kanye ne-insula (I-Postuma ne-Dagher, 2006). Ukubuyiselwa kokusebenza kwesimo okusebenzayo (i-RSFC) kuyazuzisa lapho kufundwa iziguli ezinokushiyeka okusebenzayo ngoba idatha iqoqwa ekuphumeni ekugwemeni izingqinamba zokusebenza (izinqubo zokuvusa umsebenzi zidinga ukubambisana kanye nogqozi lwezifundo), futhi inamandla njenge-biomarker yezifo ezithinta ubuchopho Uhlelo lwe-DA.

Ucwaningo lwakamuva lubhale ukungasebenzi kahle kokuxhumeka okusebenzayo kokubhekiswa kwezidakamizwa kanye nokukhuluphala ngokweqile. Ukuxhunyaniswa kokusebenza okuyisilinganiso esiphansi kubikwe phakathi kwe-dopaminergic midbrain nuclei (VTA ne-SN) nge-striatum kanye ne-thalamus (UGu et al., 2010;UTomasi et al., 2010), phakathi kwama-hemispheres (Kelly et al., 2011), naphakathi kwe-striatum ne-cortex (UHanlon et al., 2011) eziluthweni lwe-cocaine. Ukuxhuma okungajwayelekile kwe-striato-cortical kwaqoshwa nakubantu abaphuza ngokweqile (I-Rzepecki-Smith et al., 2010), abahlukumezi be-opioid (Ma et al., 2010;Liu et al., 2009;Ma et al., 2011;Upadhyay et al., 2010) nezihloko ezikhuluphele (I-Nummenmaa et al., 2012;Kullmann et al., 2012;I-García-García et al., 2012). Sekukonke, lezi zifundo zisikisela ukuthi ukuxhumana okungajwayelekile phakathi kwezifunda zesifunda kanye nezesifunda esingezansi kungenzeka kube ngaphansi kwezindawo ezifundayo ekuluthweni kwezidakamizwa kanye nokukhuluphala. Ukuvulelwa kwemininingwane emikhulu ye-RSFC ehlanganisa imininingo egciniwe evela ezifundweni eziningi kuthembisa amandla akhulayo wezibalo nokuzwela ukufaka uphawu lokuxhumana kwengqondo yomuntu (I-Biswal et al., 2010;UTomasi noVolkow, 2011). Lapha sivelisa futhi amaphethini we-RSFC kusuka ezinhlamvwini zangaphakathi nasendaweni ebhalwe nguDi Martino nozakwabo (UDi Martino et al., 2008) kusampula enkulu yezihloko ezinempilo. Ukuxhumanisa kwamaqoqo angajwayelekile okwabhalwa ngezifundo ezedlule ze-neuroimaging ngokudla / izidakamizwa kwaqikelelwa kulezi zinkundla zokuxhumana ngenhloso yokuhlola ukuthi zithini ukulutha nokukhuluphala. Ezinye izifunda zembewu ye-striatal (ie dorsal caudate) zazingadingeki ngoba izindlela zazo zokuxhumana zafakwa kakhulu ngaphakathi kobumbano lwamaphethini we-RralF ne-dorsal RSFC.

Amaphethini we-RSFC abalwa kusetshenziswa imininingo yolwazi emithathu enkulu (iBeijing: N = 198; eCambridge: N = 198; Oulu: N = 103) yephosta yezithombe zomphakathi "1000 Functional Connectomes Project" (http://www.nitrc.org/projects/fcon_1000/), okubandakanya inani lezifundo ezinempilo ze-499 (abesilisa abangama-188 kanye nabesifazane abangama-311; iminyaka: iminyaka ye-18-30). Sisebenzise indlela kaDi Martino et al. ukumaka amanethiwekhi we-dorsal ne-ventral striatal. Ukuhlelwa kwesithombe okujwayelekile (ukulungiswa kabusha nokuhlelwa kabusha kwendawo kube yindawo ye-MNI) kuthathwe ngephakheji lokubonisa imephu ye-parametric (SPM5; Wellcome Trust Center for Neuroimaging, London, UK). Ngemuva kwalokho, ukuhlaziywa kwe-seed-voxel uxhumano nge-Gram-Schmidt orthogonalization (Margulies et al., 2007;UDi Martino et al., 2008) isetshenziselwe ukubala ukuxhumana okusebenzayo kwendawo ebambisanayo (x = ± 28 mm, y = 1 mm, z = 3 mm) ne-ventral (x = ± 9 mm, y = 9 mm, z = -8 mm) izifunda zembewu yesibindi (ama-0.73ml ama-cubic volumes). Ngaphezu kwalokho, ukuxhumana okusebenzayo kwenzalo ephakathi nendawo ebonakalayo eyisisekelo ye-cortex (x = ± 6 mm, y = -81 mm, z = 10 mm; i-calcarine cortex, i-BA 17) yabalwa njengenethiwekhi yokulawula. Lawa mamamephu e-RSFC asheleliswa ngokwendawo (8-mm) futhi afakwa ekuhlaziyweni kwendlela eyodwa kwe-voxel-one of variance (ANOVA) SPM5, ngokuzimela kwenzalo yangemuva kanye ne-ventral striatal. Ama-Voxels ane-T-score> 3 (p-value <0.001, engalungisiwe) athathwa njengokuxhunyaniswa kakhulu nezifunda zembewu futhi afakwa njengengxenye yamanethiwekhi.

Iphethini ye-RSFC yembewu ye-dorsal striatal imbewu (I-Fig 3) yayizimbili futhi ifaka phakathi ngaphambili i-dorsolateral preparal (BAs: 6, 8, 9, 44-46), low low (BA: 47) and frontal frontal (BAs: 8-10), temporal (BAs: 20, 22, 27, 28, 34, 36, 38, 41, 43, 2, 3, 4 I-5, i-7-39, i-parietal ephansi nephakeme (BAs: 40, 19, 23, 24, 32, 19, 30), i-occipital (BA: 11), ne-cingulate (BAs: 8, 10 ), i-occipital (BA 20) ne-limbic (BA: 21) cortices, thalamus, putamen, globus pallidus, caudate, midbrain, pons, kanye ne-cerebellum. Iphethini ye-RSFC yembewu ye-ventral striatal nayo yayixutshwe kabili futhi yafaka i-ventral orbitof Pambal (BA: 27), i-frontal ephambili (BAs: 29-34), temporal (BAs: 36, 38, 39-23, 26, 32, 30, 47, 9, 20, 27, 28, 34, 36, 38, 23, 24, 32, 30 i-parietal ephansi (BA: XNUMX), kanye ne-cingrate (BAs: XNUMX-XNUMX, XNUMX) kanye ne-limbic (BA: XNUMX) cortices, thalamus, putamen, globus pallidus, caudate, midbrain, pons, and cerebellum. Lawa maphethini we-ventral ne-dorsal afakwe ngaphakathi kwe-inferior (BA: XNUMX) ne-frontal ephakeme (BAs: XNUMX), temporal (BAs: XNUMX, XNUMX, XNUMX, XNUMX, XNUMX, XNUMX), cingulate (BAs: XNUMX, XNUMX, XNUMX) and XNUMX i-limbic (BA: XNUMX) cortices, thalamus, putamen, globus pallidus, caudate, midbrain, pons, and cerebellum. Ngakho-ke, bekukhona ukugqagqana okukhulu kanye nomehluko obalulekile phakathi kwalezi zinhlaka zamanethi naphakathi kwe-dorsal ne-ventral elungisa lezo ezivela kwaDi Martino et al (UDi Martino et al., 2008) futhi ayahambisana namaphethini abikwe ngezifundo ze-anatomical masomo (Haber et al., 2000). Iphethini ye-RSFC ye-cortex eyi-basic visual cortex (V1) nayo ibe yimbili futhi ifaka phakathi i-occipital (BAs 17-19), i-temporal (BA 37), i-parietal ephezulu (BA 7), i-auditory (BAs 22 ne-42) ne-premotor (BA 6) ama-cortices kanye ne -patrate yangaphandle yezwe ephakeme (I-Fig 3). Ngakho-ke, iphethini yokuxhuma ye-V1 yayincane (ivolumu yenethiwekhi ye-V1 = i-16% i-grey nkhaniyi) futhi yanamathela ingxenye yenethiwekhi ye-dorsal striatal network (ivolumu ye-6% grey udaba ku-BAs 6, 7, 19 ne-37) kodwa hhayi inethiwekhi ye-ventral striatal .

 

I-Fig 3

Amanethiwekhi we-RSFC avela ku-dorsal ne-ventral striatum

Ukuhlaziywa kweMeta

Kulokhu okulandelayo sibukeza izifundo ezisebenzayo ze-neuroimaging mayelana notshwala, i-cocaine, i-methamphetamine, nensangu (Amathebula 1--4), 4), kanye nokukhuluphala nezinkinga zokudla (Amathebula 5 futhi And6) 6) ashicilelwe phakathi kukaJanuwari 1, 2001 noDisemba 31, 2011; umlutha we-nicotine awufakwanga ngoba bekunezifundo ezinhlanu ze-FMRI kuphela eziluthweni lwenicotine futhi akekho owaveza umehluko wobuchopho phakathi kwababhemayo nabangabhemi. Amagama athi "activation", "uxhumano", "dopamine", "cocaine", "insangu", "cannabis", "methamphetamine", "utshwala", "PET" no "MRI" afakiwe ekusesheni kontanga. kubuyekezwe ukushicilelwa ku-PubMed (http://www.ncbi.nlm.nih.gov/pubmed/) ukukhomba izifundo ezifanele zokucabanga ngobuchopho. Izifundo kuphela ezibike ukuxhumanisa kwendawo kwamaqoqo (e-Montreal Neurological Institute (MNI) noma amafreyimu we-Talairach stereotactic of reference) abonisa umehluko omkhulu wokusebenzisa / we-metabolic phakathi kwabasebenzisi bezidakamizwa / iziguli ezikhuluphele kanye nezilawuli (P <0.05, ilungisiwe ngokuqhathanisa okuningi) zifakiwe ekuhlaziyweni.

 

Ithebula 1

Isifinyezo sezifundo ezisebenza ngokusetshenziswa kwengqondo kogwayi olusebenzayo (olwenziwe phakathi kwe2001 ne2011) ngemiphumela yokuluthwa utshwala emsebenzini wobuchopho okufakiwe Amakhiwane 4 futhi And5.5. Izifundo zihlelwe ngama-paradigms wokukhuthaza zaba ezine ezinkulu ...

 

Ithebula 4

Isifinyezo sezifundo ezisebenza ngokusetshenziswa kwengqondo ezinamandla okusebenza kwezibalo (ezenziwa phakathi kwe2001 ne-2011) ngemiphumela yokulutha kwensangu emsebenzini wobuchopho kufakiwe Amakhiwane 4 futhi And5.5. Izifundo zihlelwe yi-paradigm yokukhuthaza izigaba ezine ezinkulu. ...

 

Ithebula 5

Isifinyezo sezifundo ezisebenza ngokusetshenziswa kwengqondo ezinamandla okusebenza kwezibalo (ezenziwa phakathi kwe-2001 ne-2012) ngemiphumela yokukhuluphala emsebenzini wobuchopho kufakiwe Amakhiwane 4 futhi And6.6. Izifundo zihlelwe yi-parimigm yokukhuthaza izigaba ezimbili ezinkulu. Inani le ...

 

Ithebula 6

Isifinyezo sezifundo ezisebenza ngokusetshenziswa kwengqondo ezinamandla okusebenza kwezibalo (ezenziwa phakathi kwe-2001 ne-2011) ngemiphumela yokudla nokuphazamiseka kokudla emsebenzini wobuchopho kufakiwe Amakhiwane 4 futhi And6.6. Inani leziguli (S) nokulawulwa (C) kwezifundo nemisebenzi ...

Ukuhlaziywa kwe-meta-okusetshenziselwa ukuxhumanisa kusetshenziselwe ukuhlola ubungako besivumelwano phakathi kwezifundo. Sisebenzise indlela yokulinganisa amathuba wokusebenzisa (UWager et al., 2004) ukwakha amathuba okuthi akwazi ukuhlangana neqoqo ngalinye elibikiwe. Ngokukhethekile, i-3D Gaussian density (15-mm-wide-wide-Half-maximum) ibigxile ekuxhumaniseni kwe-MNI kweqembu ngalinye elibike ukungezwani okubalulekile maqondana nokulawulwa kwabasebenzisi bezidakamizwa, abantu abakhuluphele kanye neziguli ezinenkinga yokudla, noma ngabe kwakuyanda noma kuncipha. I-SPM5 indlela eyodwa ye-ANOVA yayisetshenziselwa ukuhlaziya ukubaluleka kwezibalo zamamephu wokuqonda (i-3-mm isotropic resolution) ehambelana nezifundo ze-44 zokulutha kwezidakamizwa (Amathebula 1--4), 4), kanye nezifundo ze-13 ezikhuluma ngokukhuluphala nezinkinga zokudla (Amathebula 5 futhi Ne -6) .6). Ukuhlaziywa kwe-meta kubonise ukuthi ama-cortices angaphandle kanye naphakathi ahlala ekhombisa ubuthi obucashile ezifundweni ze-neuroimaging zokuluthwa kwezidakamizwa, nokuthi i-putamen / posterior insula, i-hippocampus, i-cortex esezingeni eliphakeme nephansi ye-cortex kanye ne-cerebellum ivame ukubonisa ukusebenza ukungahambi kahle ezifundweni ezikhuluma ngokukhuluphala nezinkinga zokudla (P_FWE <0.05, ilungisiwe ngokuqhathanisa okuningi ebuchosheni bonke kusetshenziswa ithiyori yenkambu engahleliwe ngokulungiswa kwamaphutha okuhlakanipha komndeni; I-Fig 4; Ithebula 7). Lokhu kuhlaziywa kwe-meta kubuye kwaveza ukuthi amathuba wokutholwa okungejwayelekile kokutholakala kokufakwa kwe-putamen / posterior insula, i-hippocampus, i-parahippocampus kanye ne-cortices yesikhashana kuvame ukuphakama ezifundweni ezikhuluma ngokukhuluphala kanye nokuphazamiseka kokudla kunokuba kufakwe kwizifundo zokulutha kwezidakamizwa._FWE <0.05; I-Fig 4; Ithebula 7). Ku-ACC (BA 24 ne-32), i-PFC (BA 8), i-putamen / posterior insula, i-hippocampus (BA 20), i-cerebellum, i-tempperum ephakathi nephezulu (i-BAs 21, i-41 ne-42) ne-supramarginal gyri amandla okuxhumana okusebenzayo. iqinile kune-dorsal kune-ventral striatum naku-anterior medial frontal cortex (BAs 10 kanye 11) yayinamandla kune-ventral kune-dorsal striatum (P_FWE <0.05; Ithebula 7).

 

I-Fig 4

Ukudidiyelwa kwemeta-based meta-ukuhlaziywa kwezifundo ze-neuroimaging ngokulutha kwezidakamizwa, ukukhuluphala nezinkinga zokudla

 

Ithebula 7

Ukudidiyelwa kwe-meta-ukuhlaziywa kwezifundo ze-neuroimaging ekulutha kwezidakamizwa, ukukhuluphala kanye nokuphazamiseka kokudla okushicilelwe phakathi kwe-2001 ne-2011 (Amathebula 2-7). I-MNI ixhumanisa (x, y, z) kanye nokubaluleka kwezibalo (T-amaphuzu) kumaqoqo abonise ukubaluleka ...

Utshwala

Ocwaningweni ophuzo oludakayo, izifundo ezenziwa nge-postmortem kanye nezifundo zobuchopho babike ukwehliswa kwe-D2R e-striatum, kufaka phakathi i-NAc (UFreund noBallinger, 1989). Izifundo ze-fMRI zabantu abaphuze utshwala zibike izimpendulo ezingejwayelekile zokuphinda zisebenze, inkumbulo yokusebenza, inhibition, kanye nemizamo yokuthinteka kwengqondo ezifundeni zobuchopho ze-cortical and subcortical (Ithebula 1). Ngesikhathi sokusebenza kabusha kwe-cue-reseacity noma ukuvezwa kotshwala, ngaphezu kwe-67% yama-activation cluster ahlukanise utshwala kusuka kuzilawuli kufakiwe kumanethiwekhi we-striatal (I-Fig 5). Isibonelo, i-ethanolous ethanol ikhuphule ukusebenza ku-ventral striatum nakwezinye izindawo ezinamandla kubaphuzo bezenhlalo kepha hhayi kwabaphuza kakhulu (UGilman et al., 2012) nezindawo zokunambitha utshwala ezivuselelekile ze-PFC, i-striatum kanye ne-midbrain kubaphuza kakhulu (Filbey et al., 2008). Ukuphuza utshwala kukhuphule ukusebenza kwe-fMRI ku-dorsolateral PFC (DLPFC) kanye ne-anthior thalamus lapho izidakwa zivezwa ezinkingeni zotshwala (UGeorge et al., 2001). I-Alcoholics ibuye ikhombise ukusebenza kwe-fMRI ephezulu kuneziqondisi ku-putamen, ACC ne-PFC yokwehla futhi yehla ku-ventral striatum kanye ne-PFC uma ubuka izintambo zokuphuza utshwala / zokulawula (U-Grüsser et al., 2004;I-Vollstädt-Klein et al., 2010b). Amaqembu abika ukungasebenzi okuphathelene notshwala ngesikhathi semisebenzi yokwenziwa kabusha kwecala kwakuvame ukutholakala kwinethiwekhi "yokugqagqana" echazwe ukuhlangana kwendawo yangaphakathi nenethiwekhi (I-Fig 3, magenta; I-21% yevolumu ndaba ye-grey udaba) kunezifundeni ezixhunywe ku-V1 kahle, noma ngabe zinamaki (aphuzi) noma cha (luhlaza) ngamanethiwekhi wesitri. Le mininingwane iphakamisa ukuthi ukuchayeka ezinhlotsheni ezihlobene notshwala kufaka phakathi ukuhlangana kwezindawo zokuxhumana zangaphakathi nezangaphakathi lapho kuvumelana nokutholwe yiPET okubonisa ukungahambi kahle ezindlini nasezinhlwini ze-dorsal striatal D2R nasekusayineni kwe-DA ekunxiliseni izidakwa (I-Volkow et al., I-2007).

 

I-Fig 5

Inani elihlobene lamaqembu angajwayelekile kwinethiwekhi ngayinye: Ukuluthwa kwezidakamizwa

Amanethiwekhi e-striatal afaka nengxenye enkulu yokutholwa okuhlobene notshwala kokusebenza kwememori yokufaka inkumbulo. Ukuhlola umphumela wokudakwa kotshwala ekusebenzeni kwengqondo, uGundersen et al. kuhlolwe ukusebenza kwe-fMRI ngesikhathi sememori yokusebenza n lapho emuva lapho izifundo ziphuza utshwala uma baphuza iziphuzo ezibandayo. Bathole ukuthi ukuphuza kakhulu utshwala kwehle kakhulu kwehle ukusebenza kwe-dorsal ACC ne-cerebellum, nokuthi lokhu kuncipha kuhluka ngomthwalo wokuqonda kanye nokugxila kotshwala begazi (UGundersen et al., 2008). Ama-Alcoholics ahlolwe ngomsebenzi wememori esebenza ngokubonakalayo abonisa ukungabi namandla kokusebenziseka kwe-fMRI ezifundeni zeparahippocampal, esekela umbono wokuthi i-hemisphere efanele isengozini enkulu yokulimala okuhlobene notshwala kunokwesobunxele (Yoon et al., 2009), nokwanda ukusebenza kwe-ACC kuqhathaniswa nezilawuli (I-Vollstädt-Klein et al., 2010a). Ngaphezulu kwe-90% yokutholwa okuhlobene notshwala okutholakele kumanethiwekhi e-striatal. Lokhu okutholakele kuyakusekela ngokuqinile ubudlelwano obuphakathi kokukhubazeka kokusebenza ngesikhathi sokusebenza inkumbulo kanye nokungasebenzi kahle kwezidakwa.

Amanethiwekhi e-striatal afaka nengxenye ebalulekile yokutholwa okuhlobene notshwala ezifundweni ezithinta ukulawula imizwa nokulawula ukungavinjelwa. Ngesikhathi silindela ukuthola imali okwenziwe ukuthi i-alcohol idakwe i-activation ku-activral kodwa kukhombise ukusebenza okuphezulu ngesikhathi sokuvezwa kotshwala, obekuhambisana nokulangazelela notshwala kepha hhayi ezilawulweni (Sula et al., 2007). Ucwaningo olwenzelwe abasebasha abasengcupheni yokuluthwa utshwala (izingane zotshwala, noma i-COA) lubike ukusebenza okuphezulu ku-PFC engazweli kanye nokwenza kusebenze kancane ezikhungweni ezisengozini yokuphuza utshwala kunakulawulo olukumelana notshwala (Heitzeg et al., 2008). Ucwaningo ngokuphoqelelwa lubike ukusebenza kwe-fMRI enkulu kwi-DLPFC kanye ne-ACC ngesikhathi sokuhlolwa kokuphazamiseka kwe-Stroop (I-Silveri et al., 2011), kanye nokusebenza kwangaphansi kokusebenza kwe-ventral striatum, i-ventC PFC, kanye ne-OFC ngesikhathi somsebenzi wokuvimbela / ukungahambi (Heitzeg et al., 2010) ye-COA kunokulawula intsha. Ukusabalala okuphezulu kokutholakele ngaphakathi kwamanethiwekhi we-striatal ngesikhathi salezi zifundo (> 83%) kusikisela ngokuqinile ukuthi ukuba sengozini kotshwala nokukhubazeka okuhlobene namandla okuvimbela kanye nezindlela zokulawula kuhlotshaniswa nokungasebenzi kahle kwe-striatal. Ngempela, sibhale ukutholakala okuphezulu kunokujwayelekile kwe-D2R ku-dorsal kanye ne-ventral striatum ehlotshaniswa nomsebenzi ojwayelekile ezifundeni zobuchopho zangaphambili (OFC, ACC, DLPFC) kanye nokufakwa ngaphakathi kwe-COA ababengezona izidakwa njengabantu abadala (Umfanekiso we-2) (I-Volkow et al., I-2006). Sabeka ukuthi ukwenyuka kwesibindi kwe-D2R kubenze bakwazi ukugcina umsebenzi ojwayelekile ezifundeni zobuchopho bangaphambili, babavikele kunotshwala.

cocaine

Amanethiwekhi e-striatal athumbe i-83% yama-activation angajwayelekile ezifundweni ze-cocaine eziphakamisa ukungasebenzi kahle kwe-cortico-striatal ekulutha kwe-cocaine. Izidakamizwa zokuthi izidakamizwa (amagama) zikhombise ukwenziwa kwe-fMRI ephansi ku-ACC ye-rostral ventral ne-caudal dorsal ACC kunamagama angathathi hlangothi kumilutha ye-cocaine (UGoldstein et al., 2007b) obekhombisa ukusebenza okuphansi kunokulawula kulezi zifunda ze-ACC (UGoldstein et al., 2009a) kodwa ukusebenza okuphezulu kumabhuzu wasebusuku (UGoldstein et al., 2009b). Ukuphathwa kwe-DA ethuthukisa umuthi i-methylphenidate (20 mg ngomlomo) kwenziwe kusebenze i-hypo ACC kusebenze kumilutha ye-cocaine (UGoldstein et al., 2010). Ngesikhathi sevidiyo ye-cocaine-cue, kusebenze ubuchopho ku-DLPFC kanye ne -ortciniport cortex yamandla kunamandla kwizifundo ze-cocaine kunakulawula kahle (UWilcox et al., 2011). Kodwa-ke, i-glucose metabolism ku-insula yesobunxele, i-OFC ne-NAc, ne-parahippocampus yangakwesokudla yayiphansi ngenkathi izifundo ze-cocaine zibuka ividiyo ye-cocaine-cue kunangesikhathi zibuka ividiyo engathathi hlangothi cue kanye ne-methylphenidate (20 mg, ngomlomo) yanciphisa impendulo engajwayelekile ku-cocaine -AmakhasiI-Volkow et al., 2010b). Lapho beyalwa ukuba bavimbele ukunxanela kwabo ngaphambi kokuchayeka ku-cocaine-cues, abahlukumezi be-cocaine bakwazi ukunciphisa umzimba we-OFC ne-NAc (ngokuqhathaniswa nesimo lapho bengahlose ukulawula ukuthanda kwabo), umphumela obikezelwa ekuqaleni komzimba oyisisekelo kwikholithi engaphansi ephansi ye-cortex (BA 44) (I-Volkow et al., 2010a). Kwabesifazane abayimilutha ye-cocaine, kodwa hhayi emadodeni, ukuchayeka kuma-cocaine-cues (ividiyo futhi kulinganiswe ne-PET ne-FDG) kuhlotshaniswa nokwehliswa okukhulu kwe-metabolism ezifundeni zobuchopho ze-cortical ezitholakala ngaphakathi kwamanethiwekhi we-striatal futhi ibuye ibe yingxenye yokulawula amanethiwekhi (I-Volkow et al., 2011a). Njengoba i-DA ilingisa amanethiwekhi ngokulawula izindlela ezitholakalayo lezi zinto ezitholakele zisekela ukubandakanyeka kwamanethiwekhi okulawula umlutha. Lapho kuvezwa umuthi ovuselelayo uqobo (i-intravenous methylphenidate, abahlukumezi be-cocaine ababikele ukuthi babe nemiphumela efanayo naleyo ye-intravenous cocaine) abahlukumezi be-cocaine bakhombise ukwanda kokusebenza kwe-metabolic ku-OFC kanye ne-cralate cingate ngenkathi izifundo zokulawula zinciphise umsebenzi we-metabolic kulezi zifunda (I-Volkow et al., I-1995).

Amanethiwekhi e-striatal abuye futhi afake i-71% yama-activation a-cocaine ahlobene nokuhlangana ngesikhathi sokusebenza kwememori nemisebenzi yokubuka kanye nezindawo zokulawula (ezisebenza ngokuxhumeka ku-V1) ezimboze inethiwekhi yasesiteshini se-dorsal striatal network (I-Fig 3, ophuzi) wayenethuba eliphakeme kakhulu lokuhlukunyezwa kunalelo elingazange lidlule amanethiwekhi we-striatal (luhlaza). Ngesikhathi sokukhuluma ngemibhalo ye-cocaine ye-nbal emuva esebenza ngomlomo ibonise ukusebenza kwe-thalamus ne-midbrain, i-dorsal striatum, i-ACC, kanye nezindawo ezinamalungu (amygdala kanye ne-parahippocampus) kanye ne-activation ye-hyper e-PFC kanye ne-cortices ye-parietal (UTomasi et al., 2007a). Ezinye zalezi zinto ezingahambi kahle zazifakwa kubanukubezi be-cocaine ngemisipha emihle ye-cocaine ngesikhathi sokufunda bebonisa ukuthi kungenzeka ukuthi ukuntuleka kokuthile kungabonakali ngaphambi kokushiya i-cocaine ekuqaleni (UTomasi et al., 2007a). Ngempela, ngesikhathi sokudambisa ukwelashwa kwabantu abafuna usizo i-cocaine abasafuna usizo ekuqaleni babonise ukusebenza kwe-hypo ku-striatum, ACC, PFC ephansi, i-precentral gyrus, ne-thalamus kuqhathaniswa nezilawuli (Moeller et al., 2010). Ezinye izifundo kwimemori yokusebenza ziveze ukuthi izintambo ze-cocaine zingakhuphula ukusebenza kobuchopho ku-occipital cortex (UHester noGaravan, 2009). Ngesikhathi sokubonwa kwemisebenzi yokunaka, abahlukumezi be-cocaine babene-activation ephansi ye-thalamic kanye ne-occipital cortex ephakeme kanye nokwenza kusebenze kwe-PFC kunokulawulwa (UTomasi et al., 2007b). Ubudlelwano obuphakathi kwe-cortico-striatal dysfunction kanye ne-fMRI engajwayelekile ngesikhathi sokukhumbula nemisebenzi yokunaka kwenzeka ikakhulu ekuhlanganeni kwendawo yokuxhumana yangaphakathi ne-ventral, eyayinezivikelo eziphakeme ze-3 izikhathi eziphakeme (inani elilinganayo leqoqo elisetshenziswe njengevolumu yenethiwekhi) kunezifunda ezingekho ngokuxhumeka i-striatum (I-Fig 5).

Ngesikhathi sokuthatha izinqumo nomsebenzi wokuhlukumeza i-Iowa ubonise ukugeleza kwegazi okuphezulu kwenhliziyo (i-rCBF; kulinganiswe ne- ¹⁵I-O-water PET) kwesokudla kwe-OFC ne-rCBF ephansi ku-DLPFC ne-PFC yokwelapha kuqhathaniswa nezilawuli (I-Bolla et al., 2003). Ngesikhathi sokukhethwa kokuphoqelelwa ngaphansi kwezimo ezintathu zenani lemali izifundo ze-cocaine zikhombisa izimpendulo eziphansi ze-fMRI zokuthola umvuzo wezezimali e-OFC, PFC kanye ne-occipital cortex, midbrain, thalamus, insula kanye ne-cerebellum (UGoldstein et al., 2007a). Ukutholakala okuphansi kunokwejwayelekile kwe-D2R ku-dorsal striatum kwakuhlotshaniswa nezimpendulo ze-thalamic activation ezishintshile kanti ku-ventral striatum kwakuhlotshaniswa nokwanda kokunyanzelwa kwe-PFC kubantu abayimilutha ye-cocaine (Asensio et al., 2010). Ngokufanayo nemisebenzi yokuqonda, okutholakele ekuxhumaneni kwendawo yangaphakathi neyangaphandle kubonise amathuba aphezulu kunalawo ezifundeni ezixhumene ne-striatum.

Amashumi ayisikhombisa nane nane amaqoqo obuchopho abikwe izifundo ze-fMRI ngemisebenzi yokuvimbela ukufakwa afakiwe ezinkundleni zokuxhumana. Ngesikhathi umlutha we-cocaine we-go / no-go ovinjelwe ukhombise ukusebenza okungaphansi kunolawulo e-OFC, indawo eyengeziwe yezimoto kanye ne-ACC, izifunda ezingaba síxaka kokulawulwa kokuqonda (UHester noGaravan, 2004). Abasebenzisi abasebenzisa i-cocaine abafushane nabakhela isikhathi eside bakhombisa ukusebenza okwahlukile ku-PFC, i-cortex yesikhashana, i-cingulum, i-thalamus kanye ne-cerebellum (ICololly et al., 2011). Ngesikhathi semisebenzi ehlukene yokuvimbela (ukungenelela kwe-Stroop) imilutha ye-cocaine ikhombise i-rCBF ephansi ngakwesokunxele ku-ACC nakwa-PFC yangakwesokudla, kanye ne-rCBF ephakeme kwi-ACC yangakwesokudla kunezivumelwano (I-Bolla et al., 2004). Ukuxhumana okusebenzayo kwe-Striatal kuhlulekile ukuchaza umehluko wokusebenza kobuchopho ocwaningweni olusebenzise imisebenzi yokumisa uphawu (Li et al., 2008). Lezi zifundo zikhombise ukusebenza kancane kwe-ACC, parietal kanye ne-occipital cortices kubahlukumezi be-cocaine. Izifundo ze-PET ezilinganisa ama-mu opioid receptors (kusetshenziswa [¹¹I-C] carfentanil) ikhombise ukubopha okuphezulu okucacisiwe kokubanjwa okuthembekile kosuku lwe-cocaine kosuku olulodwa kunokulawulwa, futhi lokhu kukhubazeka kwehla ngokungazilandeli futhi kuxhumene nokusetshenziswa kwe-cocaine (UGoinin et al., 2005;I-Ghitza et al., 2010).

I-Methamphetamine

Kuqhathaniswa nokulawula izifundo, abahlukumezi be-metamphetamine abahlolwa ngesikhathi sokudalwa kwangaphambi kokukhipha izidumbu bakhombise ukwehla kwe-glucose metabolism ku-striatum ne-thalamus ngenkathi bekhombisa umsebenzi owandayo ku-cortex ye-parietal (I-Volkow et al., 2001a). Lokhu kuphakamise ukuthi bobabili iDA kanye nezindawo ezingezona ze-DA ezilungiswe ubuchopho zithinteke ngokusetshenziswa okungapheli kwe-metamphetamine (I-Volkow et al., 2001a). Ngaphezu kwalokho sehlile umsebenzi we-DA we-striatal uhlotshaniswa nethuba elingcono lokuphinda ubuye ngesikhathi sokwelashwa (U-Wang et al., 2011b), ukulahla obekuhanjiswa kwakuhambisana nokubuyiselwa okuyingxenye kwe-DAT ye-striatal (I-Volkow et al., 2001b) kanye nokusebenza kwengqondo kwesifunda (U-Wang et al., I-2004), kanye nokunciphiswa kwe-striatal D2R nakho kuhlotshaniswa nokuncipha kokusebenza kwe-metabolism e-OFC muva nje abahlukumezi be-methamphetamine abasanda kuxoshwa (I-Volkow et al., 2001a).

Ingxenyana enkulu (70%) yokutholwa kwe-fMRI ehlobene ne-methamphetamine yayihlanganiswa ngamanethiwekhi we-striatal (I-Fig 5). Uma kuqhathaniswa nezilawuli, abantu abathembele ku-methamphetamine babonisa ukusebenza kwe-ACC ephakeme ngesikhathi sokuvinjwa kokuhamba / ukungahambi-kokuhamba (Leland et al., 2008), kanye nokwehliswa kwesokudla kwe-PFC ngesikhathi sokuphazanyiswa kweStroop (Salo et al., 2009). Iningi lalawa ma-activation ama-activation angajwayelekile (i-88%) avela kunethiwekhi ye-dorsal (kufaka phakathi ukunqwabelana kwayo nenethiwekhi ye-ventral). Ngesikhathi sokuthatha izinqumo, noma kunjalo, ingxenye engezansi (i-64%) yamaqoqo yayihlanganiswa ngamanethiwekhi wesikhungo. Besebenzisa umsebenzi wokuqagela ukukhetha okubili, uPaulus nozakwabo bathola ukuthi ukusebenza kwe-fMRI kwakuphansi ku-PFC (UPaulus et al., 2002), I-OFC, i-ACC ne-cortex ye-parietal yezifundo ezixhomekeke emethamphetamine kunezilawuli (UPaulus et al., 2003). Ngaphezu kwalokho, inhlanganisela yezimpendulo zokwenza kusebenze kulezi zifunda yabikezela kangcono isikhathi sokuphinda ibuye futhi yakhombisa amaphethini e-activation ahlukahlukene njengomsebenzi wezinga lephutha ku-insula yangakwesokunxele ne-DLPFC (UPaulus et al., 2005).

I-Marijuana

Ukubandakanyeka kokudonswa kanzima kokuluthwa kwensangu akucaci kahle ngoba akusikho ukukhishwa okuyisisekelo kwe-D2R noma ukukhishwa kwe-striatal kwe-DA (ngemuva kwenselelo ye-amphetamine) eziwombeni eziphuze kakhulu zePET ezenziwa [¹¹C] i-raclopride (Urban et al., 2012;Stoke et al., 2012). Ucwaningo lwe-FDG luveze ukuthi ngenkathi benikezwa abahlukumezi be-tacrahydrocannabinol (i-THC) abahlukumezi bezokwelapha bakhombisa ukwanda kwe-OFC kanye ne-PFC yezokwelapha kanye nase-striatum kanti izilawuli azizange, kodwa zanda umfutho we-cerebellar kubo bobabili abahlukumezayo nezilawuli eziphakamisa ukuthi amanethiwekhi we-striatal abandakanyeka ekuluzweni kwensangu (I-Volkow et al., 1996b). Imikhondo ehlobene nentsangu eyejwayelekile ebhekise eqenjini lokutholwa necala lokungathathi hlangothi iboniswe ukwandisa ukusebenza kwe-fMRI eVTA, thalamus, ACC, insula, nase-amygdala, ukusekela ukubandakanyeka kwamanethiwekhi ezokuvakasha, kanye nakwezinye izigameko zangaphambili, zeparietal kanye ne-occipital kanye ne-cerebellum kwensangu esanda kubanjiswa abasebenzisi (Filbey et al., 2009). Ngesikhathi somsebenzi wokunaka okubukwayo, abahlukumezi bensangu babenamandla okusebenza okuphansi kwe-FMRI ku-PFC yangakwesokudla, i-cortex ye-parietal ne-cerebellum (ejwayelekile nesikhathi sokungalawulwa) kanye nokusebenza okuphezulu endaweni engaphambili, e-parietal kanye ne-occipital cortices kunezilawuli (Chang et al., 2006). Ngesikhathi sokusebenza inkumbulo yokusebenza, noma kunjalo, abahlukumezi bensangu babonisile ukunciphisa ukusebenza ku-lobition yesikhashana, i-ACC, i-parahippocampus ne-thalamus ngokunyusa ukusebenza komsebenzi, ukusebenza kokusebenzisana kweqembu = okuphambene nokulawula (UPadula et al., 2007). Ngesikhathi sokuvinjezelwa kokuya / ukungahambi, intsha ebinemilando yokusetshenziswa kwensangu ikhombise ukwenziwa okuphezulu kwe-fMRI eDLPFC, parietal kanye ne-occipital cortices, nokufaka ukucabanga kwentsha ngaphandle kwemilando yokusetshenziswa kwensangu (I-Tapert et al., 2007). Ngesikhathi sokuhlanganiswa kwe-visuomotor nomsebenzi wokubona womunwe ohambisanayo obonwe yinqola ebheka ukukhanya, abasebenzisi bensangu babene-activation ephezulu ye-PFC kanye nokuqaliswa kwangaphansi kwe-cortex okubukwayo kunokulawula (King et al., 2011). Amashumi ayisithupha nesishiyagalolunye amaphesenti ama-activation ama-activation angajwayelekile ezifundweni ngemiphumela yensangu ekusebenzeni kobuchopho ayatholakala ezifundeni ezixhumene ne-striatum.

Ukukhuluphala

Ukuziphatha okuphoqelela okufana nokuphoqelela kumagundane agcwele kuhlotshaniswa nokucishwa phansi kwe-striatal D2R (UJohnson noKenny, 2010) kanye nokukhuluphala kuxhunyaniswe ne-striatal D2R ebantwini (U-Wang et al., I-2001), kuphakamisa ukuthi i-neuroadaptations ejwayelekile emendweni oyisisekelo we-DA ingadonsa ukukhuluphala ngokweqile nokulutha kwezidakamizwa. Izifundo ze-PET zesisekelo se-brain glucose metabolism kubantu abagcwele ngokweqile zibike ukwehla komsebenzi we-metabolic e-OFC naku-ACC ehlotshaniswa nokutholakala kwe-D2R ye-striatal ephansi kunokujwayelekileI-Volkow et al., 2008b).

Ukusebenza kobuchopho ezindlini ezi-dorsal and ventral striatum, insula, hippocampus, OFC, amygdala, PFC medial kanye ne-ACC kuphakanyiswe ukudalulwa okubukwayo ekudleni okuphezulu kwe-caloric kwakuphakama kwabakhuluphele kunokulawula abesifazane (URothemund et al., 2007;Stoeckel et al., 2008). Ngokufanayo, izinkomba zokudla ezibonakalayo zivezile izimpendulo ezandisiwe ze-fMRI ezindaweni zangaphambili, zesikhashana, nezomzimba kubantu abadala abakhuluphele kunokulawula (I-Dimitropoulos et al., 2012), nokwenza kusebenze kwe-hippocampal kubonise ukuhlangana ngamazinga we-plasma wokuzila we-insulin kanye nokwehlukaniswa okhalweni kwentsha (UWallner-Liebmann et al., 2010). Ukusebenza kwe-striatal ukuphendula i-chocolate milkshake etholakele kuhlotshaniswa nokutholwa kwesisindo somzimba kanye nokuba khona kwe-A1 ye-alqine ye-TaqIA limited discature polymorphism, ehlotshaniswa nohlobo lwe-D2R ekubopheleni ukusayina kwe-striatal kanye nokusayina umkhondo we-striatal DA ukusayina (UStice et al., 2008). Intsha enobungozi obukhulu bokukhuluphala ikhombise ukwenziwa okuphezulu ku-caudate ne-operculum ekuphenduleni umthamo we-chocolate milkshake kunalawo asengozini ephansi yokukhuluphala (UStice et al., 2011). Ngesikhathi sokudonswa esiswini, njengoba kwenzeka ngesikhathi sokudla, izifundo ezikhuluphele zikhuphule ukusebenza kwe-fMRI kunezifundo ezijwayelekile zesisindo ku-cerebellum nase-posterior insula futhi kwehlisa ukusebenza ku-amygdala, midbrain, hypothalamus, thalamus, pons, and anterior insula (UTomasi et al., 2009b). Amaphesenti ayisishiyagalombili nambili% we-activation group kusuka kulezi zifundo ekusetshenzisweni kwe-cue-reacuction kwenzeka ezifundeni ezixhumene ne-striatum (I-Fig 6). Kuhambisana nalezi zifundo ze-PET zokuphendula ezilinganisa i-D2R nge [¹⁸I-F] i-fallypride ngesihloko se-feta isikhombisa ukuvumelana okungafani phakathi kwe-ghrelin ne-D2R endaweni ephakathi nendawo ne-ventral striatum nakwi-cortex yesikhashana yesikhashana, isigxobo sesikhashana, insula ne-amygdala (UDunn et al., 2012).

 

I-Fig 6

Inani elihlobene lamaqembu angajwayelekile kwinethiwekhi ngayinye: Ukukhuluphala nezinkinga zokudla

Ukuqonda kokudla nokulawulwa kokudla

Ezimweni ezijwayelekile ukunatha kokudla kucatshangwa ukuthi kuzonqunywa yizo zombili izinto zasekhaya (ukulinganisela kwamandla nezakhi emzimbeni) kanye nezinto ezingeyona i-homeostatic (injabulo yokudla), kanti ubuchopho be-DA buhlotshaniswa nezindlela zokudla (I-Volkow et al., 2003b). Izifundo ze-FMRI ze-Pharmacological zikhombisile ukuthi i-hypothalamic activation ibikezela ukudliwa kokudla lapho inani le-plasma okuhlushwa nge-PYY, i-peptide hormone enikeza isibonakaliso satiety somzimba esisuselwa ebuchosheni, siphansi futhi ukuthi ukusebenza ku-OFC striatum, VTA, SN, cerebellum, I-PFC, i-insula ne-cingulum ingabikezela ukusebenza kokuphakelwa uma izinga lokuphazamiseka kweplasma le-PYY liphezulu (IBatterham et al., 2007).

Ucwaningo oluhlobene nomcimbi oluqhathanisa izimpendulo zobuchopho nokunambitheka kwe-sucrose namanzi angenamsoco kukhombisile ukuthi indlala ihlotshaniswa nokusebenza kwe-fMRI insula, thalamus, cerebellum, cingulum, SN kanye nasezifundeni zobuchopho be-cortical kanti ubuciko be-satiety buhlotshaniswa nokuzenzakalela ku-parahippocampus, hippocampus, amygdala ne I-ACC (I-Haase et al., 2009). Kulolu cwaningo imiphumela ehlukile yendlala eqhathaniswa nokuqina kokusebenza kwengqondo ukunambitheka (okunosawoti, omuncu, obabayo, obumnandi) bekuqine kwabesilisa kunakulabesifazane, ikakhulukazi ku-dorsal striatum, amygdala, parahippocampus nase posterior cingulum (I-Haase et al., 2011). Izifundo ze-PET zokulawulwa kwe-inhibitory ezimweni zokulamba ezisebenzisa ukuvuselelwa kokudla kwangempela kwaveza ukuthi inhibition enenhloso yesifiso sokudla yancipha i-glucose metabolism e-amygdala, hippocampus, insula, striatum ne-OFC emadodeni kodwa hhayi kwabesifazane (U-Wang et al., I-2009). Ingxenyenamba enkulu (> 31%) yamaqoqo wokuqalisa yenzeke ezifundeni ezisebenza ngokuxhuma kuzo zombili i-dorsal ne-ventral striatum (I-Fig 6, magenta).

kwemikhuba yokudla

Ucwaningo lwe-Chemacological luveze ukuthi ukuphazamiseka kokusayinelwa yi-DA kwi-striatum kungavimbela ukondla okujwayelekile kumagundane (I-Sotak et al., I-2005;I-Cannon et al., 2004) nokuthi i-DA ekhombisa imodigates ibuye isebenze ezindlekweni zokudla kubantu (I-Volkow et al., I-2002). Izifundo ze-PET zeziguli ezihlushwa i-anorexia (ukulawula okungaphezulu kokudla) zikhombisa ukuthi kuphakeme kakhulu kunokutholakala kwe-striatal D2R (UFrank et al., 2005). Ngokuphambene nalokho, ucwaningo lwakamuva ezigulini ezingama-non-feta ezinamafutha okudla okudla kakhulu lukhombisa ukuthi ngenkathi zingafani ekutholakaleni kwe-D2R kusuka ezilawulweni zikhombisile ukukhululwa kwe-striatal DA ngesikhathi sokuvuselelwa kokudla (U-Wang et al., 2011a). Ucwaningo lwe-fMRI luveze ukuthi uma zivezwa ezithombeni zokudla ezimnandi iziguli ezinenkinga yokudla okunezifo ezinamandla zinezimpendulo eziqinile ze-OFC ezilawula ngenkathi iziguli ezine-bulimia amanosa zine-ACC enamandla nezimpendulo zokufakelwa ukwedlula izilawuli (I-Schienle et al., 2009). Ngesikhathi sokuvinjezelwa kokuya / ukungahambi njalo, intsha edla ukudla / yokuhlanza intsha yabesifazane ibonise ukusebenza okuphezulu ku-cortex yesikhashana, i-PFC ne-ACC kunokulawula, kanye neziguli ze-anorexia manthaosa zikhombisa ukwenziwa okuphezulu kwe-hypothalamus kanye ne-PFC yamuva (Khiya u-al., 2011). Njengoba munye kuphela walawa maqoqo owatholakala ngaphandle kwamanethiwekhi ezwekazi, le mininingwane ibuye futhi ifundise indima yamanethiwekhi we-cortico-striatal ezinkingeni zokudla.

Izifunda ezingaphambi

I-cortex yangaphambi kokuqala kanye ne-striatum zixazululwa ngamanethiwekhi we-cortico-striatal alungiswe yi-DA (I-Haber, i-2003). I-cortex yangaphambili idlala indima eyinkimbinkimbi ekuqondeni, kufaka phakathi ukulawulwa kokuvinjezelwa, ukwenza izinqumo, ukumiswa kwemizwa, inhloso, isisusa kanye nokuqondiswa kobuntu phakathi kwabanye. Kubhekwe phezulu ukuthi ukungasebenzi ezindaweni ezihamba phambili kungalimaza ukulawulwa kokudla izidakamizwa okuphoqelekile (I-Kalivas, i-2004;I-Volkow ne-Fowler, i-2000), nokuthi ukuphazamiseka kwangaphakathi kwe-cortex kungaba nemiphumela emibi ekuluthweni kwezidakamizwa (I-Volkow et al., I-2006).

Ukuhlukunyezwa kwangaphandle okuvezwa ukuhlaziywa kwethu kwe-meta kuyahambisana nokuhlobana phakathi kokuncishiswa kwe-striatal D2R futhi kunciphise umsebenzi we-metabolic ku-ACC, OFC ne-DLPFC ngaphambili kwabikwa ukuthi kune-cocaine nabahlukumezi be-methamphetamine nabaphuza utshwala (I-Volkow et al., I-1993;I-Volkow et al., 2001a;I-Volkow et al., I-2007). Kusukela i-ACC, i-lateral ye-OFC ne-DLPFC zibandakanyeka ekulawuleni inhibitory kanye nokwenza izinqumo (I-Goldstein noVolkow, i-2002;I-Phan et al., I-2002), le nhlangano iphakamisa ukuthi ukulahleka kokulawula ukusetshenziswa kwezidakamizwa (I-Volkow et al., 1996a) kungakhombisa ukumelwa okungalungile kwe-DA kulezi zifunda zangaphambili. Le hypothesis isekelwa izifundo ezihambisana nokuncishiswa kwe-striatal D2R kanye nezimpawu zokuphoqelela kubahlukumezi be-methamphetamine (U-Lee et al., 2009) namagundane (Everitt et al., 2008) kanye nalezo ezixhumanise ukungasebenzi kwe-ACC nokuziphatha okuphoqayo nokuziphoqelela (I-Rolls, i-2000). Kodwa-ke, okunye okungenzeka ukuthi ukuhlukunyezwa kokuqala ezindaweni ezingaphambili kubangela ukusetshenziswa kwezidakamizwa okuphindaphindiwe nama-neuroadaptations anciphisa i-striatal D2R. Isibonelo, abantu abangaphuzi utshwala abanomlando womndeni wotshwala babephezulu kune-DriNUMXR evamile ye-striatal ehambisana nokusebenza okuvamile kwe-ACC, OFC ne-DLPFC, bephakamisa ukuthi umsebenzi ojwayelekile ezifundeni ezingaphambi kokuthuthukisa ukulawulwa kokuvinjezelwa nokulawulwa okungokomzwelo kungaba yindlela obevikela lezi zifundo ekusetshenzisweni kabi kotshwala (I-Volkow et al., I-2006). Kuyathakazelisa ukuthi ucwaningo lwakamuva oluqhathanisa ukungathandani kwengane yakho ngokulutha okuvuselelayo kukhombise umehluko obonakalayo kwivolumu ye-medC ye-medC (Ersche et al., 2012), kuphakamisa ukuthi lo mehluko ubonise ukuvezwa komuthi kunokuba sengozini yofuzo (IVolkow kanye noMthengisi, i-2012).

Izifunda zesikhashana

I-striatum ibuye ixhumeke nezinhlaka ze-medial temporal lobe (hippocampus parahippocampal gyrus) ezibalulekile kwimemori ebonakalayo kepha futhi nokufaka isimo (Haber et al., 2006). Izifundo zokuvuselelwa kobuchopho ezifundweni ezisuselwa emvuzweni zibhale ngokubandakanyeka kwezakhiwo ze-loal temporal lobe ekwandiseni inkumbulo elandelayo (I-Adcock et al., 2006;UWittmann et al., 2005). Ngakho-ke izintambo zezidakamizwa zingadala ukunxanela kokumemezela okusebenzayo kokufunda amasekhethi ku-medial temporal cortex, futhi lokhu kusebenze okuthuthukile kokujikeleza kwememori kungasiza ukunqoba ukulawulwa kokuvinjwa okukhishwe yi-preortalal cortex ekudleni nasekulutshweni kwezidakamizwa (I-Volkow et al., 2003a). Ukuhlaziywa kwethu kwe-meta kuveze ukuthi umlutha wezidakamizwa, ukukhuluphala kanye nokuphazamiseka kokudla kubonakaliswa yizinto ezejwayelekile zokuhlukunyezwa kobuchopho kuma-medial temporal cortex (hippocampus, parahippocampal gyrus kanye ne-amygdala), ophakeme futhi ophansi we-cortices wesikhashana kanye ne-posterior insula (P_FWE<0.05). Iphethini yokuphazamiseka kokusebenza kobuchopho ngokwengxenye igqagqanise i-dorsal (40%), i-ventral (10%) namanethiwekhi agqagqene (48%); Kuphela yi-2% yokungajwayelekile engakhombisanga ukugqagqana namanethiwekhi we-striatal. Ukuhlaziywa kwethu kwe-meta kuveze nokungajwayelekile okunamandla ezakhiweni zesikhashana se-lobe ekukhulupheni nasekudleni kokudla uma kuqhathaniswa nokulutha kwezidakamizwa (I-Fig 4). Lokhu kuphakamisa ukuthi lezi zindawo zesikhashana zibandakanyeka kumthethonqubo wokuziphatha kokudla ngezinga elikhulu kakhulu kunakulawulo lokudla izidakamizwa. Ukudla okwenziwe ngokuqondile kulawulwa yizo zombili izindlela zasekhaya nezinomvuzo futhi ngenkathi uhlelo lwe-homeostatic liguqula indlela yomvuzo, luphinde futhi lusebenzise ezinye izifunda zobuchopho ngamahomoni ahlukahlukene we-peripheral kanye nama-neuropeptides alawula indlala nokuqina. Izifunda ze-medial temporal (i-hippocampus, parahippocampus) ziveza ama-leptin receptors (U-Williams et al., I-1999) nama-insulin-receptors receptors (UWilczak et al., 2000) kanye ne-mRNA yohlobo lwe-ghrelin receptor gene (UGuan et al., 1997). Ukuzibandakanya okukhulu kakhulu kwe-medial temport cortices ekukhuluphele kunokuba umlutha kuhambisane nokubandakanyeka kwama-hormone nama-neuropeptides alawula ukungena kokudla ngendlela yomgwaqo we-homeostatic.

Umvuzo nemikhuba

Kuzo zombili izinqubo zokudla izidakamizwa nokudla ezidingweni zangaphakathi ekuqaleni zishayela isisusa sokuphinda lokhu kuziphatha. Noma kunjalo ngezimpendulo eziphindaphindwe kaningi zokubonakaliswa kanye nezinhlangano ezifundile ziguqula isisusa sokukhuthazeka kulukhuthazo olunesimo esibikezela umvuzo. Lokhu kuguqulwa, kanye nesisusa esithuthukisiwe sokwenza izimfanelo ezidingekayo zokusebenzisa umvuzo (izidakamizwa noma ukudla), kudinga ukubandakanyeka kwe-dorsal striatum (UBelin no-Everitt, i-2008). Ngaphezu kwalokho, ukuvezwa okuphindaphindiwe kokumatanisa okuhambisana nakho kwimikhuba engaqhubeka nokusebenza kokushayela (kufaka phakathi ukudla noma ukudla izidakamizwa noma utshwala) kubandakanya nezindawo ezi-dorsal striatal. Kodwa-ke, lapho kubuyekezwa ukugqama okukhulu phakathi kokuxhumana phakathi kwe-ventral kanye ne-dorsal striatal, akumangazi ukuthi izifundo zibonisa ukusebenza kwe-ventral kanye ne-dorsal striatum bobabili ngomvuzo nangemodi. Ngokufanayo ngenkathi i-dorsal striatum ihlotshaniswa kakhulu nemikhuba yokwakheka kwayo kungadinga ukuthuthuka kusuka kumaphakathi kuya ezifundeni ze-dorsal striatal (Everitt et al., 2008).

Amanethiwekhi asengozini yokulutha nokukhuluphala

Ukuthola okubalulekile kulolu cwaningo ukuthi ukusebenza okulimazayo kokudla noma umlutha wezidakamizwa kuvame ukwenzeka ezifundeni zobuchopho ngokuxhumeka kuzo zombili i-dorsal kanye ne-ventral striatum. Lezi zifunda ezisengcupheni zibalulekile ukuze kulawulwe ingqondo (i-anterior cingulum nendawo eyengeziwe yezimoto), umvuzo kanye nogqozi (i-striatum kanye ne-medial OFC) nokufunda okukhuthaza umvuzo (i-hippocampus ne-parahippocampal gyrus). Ukugqagqana kwamaphethini wokuxhuma kwe-striatal kuveza ukuthi ukushintshwa kwe-dopaminergic kusuka ku-dorsal kanye ne-ventral striatum kubalulekile kulezi zindawo, futhi ukuba sengozini kwabo okuphezulu kusikisela ukuthi ukudla umlutha wokudla / izidakamizwa kungashintsha ibhalansi yokushintshashintsha kokushintshashintshwa kwesimo nokusebenza kobuchopho kulezi zifunda.

Ukulinganiselwa

Ukuhlaziywa kwethu kwe-meta kufaka phakathi izifundo ngemiphumela emibi yezidakamizwa nokudla (izintambo), kanye nezifundo zokuqondisisa (inkumbulo, ukunakwa, ukuvimbela, ukwenza izinqumo) nomzwelo lapho izidakamizwa noma ukudla kungatholakali. Njengoba imiphumela eqondile neyesikhathi eside yokuluthwa kokudla / izidakamizwa yehlukene abahlanganyeli ocwaningweni lwangaphambili bangase noma bangabi sengozini enkulu kakhulu ekushintsheni kobuchopho. Lokhu kungakhuphula ukwahluka, kunciphisa ukuhunyushwa kwemiphumela. Ukuvezwa okungaphezulu kokukhubazeka ngokweqile kwe-medial temporal lobeatness ngokweqile kanye nezinkinga zokudla ngokuqhathaniswa nalezo eziluthelwa izidakamizwa kungabonisa ubulukhuni benkinga njengoba kungekho lula ukulinganisa ubukhulu, ubude besikhathi noma iminyaka yokuqalwa kwaloluphazamiso.

Ngokufingqa lokhu kuhlaziya kwezifundo zamuva nje zokucabanga kobuchopho ezinhlotsheni ezahlukahlukene zokuluthwa izidakamizwa nokuphazamiseka okubonakaliswa yi-dyscontrol yokuziphatha ngaphezulu kwezindlela zokuziphatha ezivuzayo (ukudla) kukhombisa ukuthi kunesimo esibonisa ukwenziwa kwezinto okungajwayelekile (kokubili ezinkambeni kanye nangemisebenzi yokuqonda) evame ukwenzeka njalo ezindaweni lapho kukhona ukugcwala phakathi kwendlela yangaphakathi neye-dorsal striatal. Lokhu kulungisa kubantu ukuthi zombili iziqubu zangaphakathi (ezihlotshaniswa kakhulu nokucutshungulwa kwemivuzo) kanye ne-dorsal striatum (ehambisana kakhulu nemikhuba namasiko ngokulutha) ziphazamiseka ekuphazamisekeni okuluthayo (Ohlakaniphile, 2009) nokuthi lezi zinto ezingahambi kahle zithonya ukucubungula imivuzo (izidakamizwa nokudla) isisusa esihlobene nomvuzo (izintambo) nezinqubo zokuqonda ezidingekayo ukuze ukwazi ukuzilawula (umsebenzi wokuphatha). Kodwa-ke, izifunda zedical tempical cortical cortical eziyingxenye yomgwaqo oshubisayo okhombisa ubungozi obukhulu ekukhuluphelweni nasekuphatheni kokudla kunokuba umlutha wezidakamizwa (I-Fig 4), okubonisa ukuthi kunezindlela ezihlukile zokungahambi kahle phakathi kwalezi zinto zokuphazamiseka.

Lo msebenzi wenziwa ngokwesekwa yi-National Institutes of Alcohol Abuse and Alcoholism (2RO1AA09481).

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