Ka pele. Psychiki, 07 Hlakubele 2016 | http://dx.doi.org/10.3389/fpsyt.2016.00030
- Lefapha la Biology, Setsi sa Neuroscience, Univesithi ea Texas ho San Antonio, San Antonio, TX, USA
Boitšoaro bo amanang le lithethefatsi ho batho le litoeba hangata ho nahanoa hore li hlaha litsing tsa ho ithuta tse sa sebetseng hantle. Boithuto ba preclinical bo bonts'a hore ho fumana le ho itlhahisa ha mekhoa e mengata e itšetlehileng ka lithethefatsi ho kenyelletsa sebaka sa ventral tegmental (VTA), sebopeho sa midbrain se entsoeng ka dopamine, GABA, le glutamate neurons. Boiphihlelo ba lithethefatsi bo fetola tšusumetso ea thabo le thibelo ea methapo ea kutlo ea VTA dopamine neurons, ho fana ka maikutlo a karolo ea bohlokoa bakeng sa barekisi ba VTA ho ahlola litlamorao tsa lithethefatsi. Thutong ena, re hlahisa bopaki bo amang VTA ka boits'oaro bo amanang le lithethefatsi, re totobatsa phapang ea palo ea baahi ba Vonal, 'me re tšohla litlamorao tsa boitšoaro ba batho ba ikhethileng ba amanang le VTA. Liteko tsa nako e tlang lia hlokahala ho tseba hore na li amanang le VTA le hore na ke liphoofotsoana life tse fumanoang kahare ho VTA tse amanang le boits'oaro bo itseng. Lithuto tse ling hape li hlokahala bakeng sa ho tseba liphetoho tse amanang le li-synaptic tse amanang le li-dopamine le li-neuron tse seng dopamine ho VTA e latelang ea taolo ea lithethefatsi. Ts'oaetso ea li-circuits tsa neural le liphetoho tse amanang le boits'oaro bo tsamaeang le lithethefatsi li ka totobatsa liphofu tse ka ba teng tsa neural bakeng sa ts'ebetso ea ts'ebetso ea meriana le ts'usumetso e matla ea kelello ho phekola mathata a tlhekefetso ea lithethefatsi.
Selelekela
Tšebeliso e mpe ea lithethefatsi ke bothata bo bohlokoa ba lefats'e, 'me Ofisi ea Machaba a Kopaneng ea Lithethefatsi le Bokebekoa e hakanya hore batho ba limilione tse 246 lefatšeng ka bophara ba sebelisitse lithethefatsi tse seng molaong ho 2013. Bothata bo boholo ke koluoa e kholo ea mathata a ts'ebeliso ea lithethefatsi (SUDs), eo ho eona 2014 e neng e hakantsoe hore e hlokofatse batho ba limilione tse 21.5 ba Amerika, e tsamaellanang le ~ 8% ea baahi (1). Ntle le tšusumetso ea motho ka mong ea SUD, ho na le tšusumetso e kholo ea moruo ka lebaka la tlhahiso e lahlehileng, tlolo ea molao le litšenyehelo tsa tlhokomelo ea bophelo bo botle, tseo ho latela ofisi ea US ea Pholisi ea Naha ea Dawa e hakanyetsoang ho bitsa $ 180.8 bilione ka selemo ho US. a le mong.
Hona joale li-SUD li amoheloa hore li lula li le teng khafetsa moo bophahamo ba lefu lena bo amanang le palo ea mekhoa ea tlhahlobo e fumanoeng ke motho ka selemo se fetileng. Ho latela DSM-V, litekanyetso tsa SUD li oela lihlopheng tse 'ne tse kholo tsa matšoao: taolo e senyehileng (ke hore, sebelisa se fetang seo ho reriloeng), ho senyeha ha sechaba (ke hore, tšebeliso ea lithethefatsi ka litšenyehelo tsa likamano tsa motho le ts'ebetso e senyehileng ea mosebetsi), e kotsing boits'oaro (ke hore, sebelisa ntle le litlamorao tse tsejoang tse mpe), le litlamorao tsa meriana (ke hore, ho mamella le ho ntša). E 'ngoe ea lintlha tse tšosang ka ho fetisisa tsa ho phekola li-SUD ke ts'oaetso e phahameng ea ho oela hape, e etsahalang ka ~ 40-60% ea batho ka bomong (2). Ho basebelisi ba lithethefatsi, ho pepesetsoa lithahasello tse tsamaeang ka lithethefatsi ho etsa hore ba lape hore le bona ho ka khothaletsa monyetla oa ketsahalo e khutlang hape (3). Ho theola likamano lipakeng tsa lithethefatsi le litloaelo tse amanang le eona ho na le ts'episo e le mokhoa o seng oa meriana oa ho phekola li-SUDs (4). Leha ho le joalo, kutlwisiso ea rona ea li-circuits tse khethehileng tsa neural le liphetoho tsa neural tse ikarabellang bakeng sa boits'oaro bo amanang le lithethefatsi ha li ea fella.
Mefuta ea Rodent ea Lithethefatsi tse Ikemetseng
Lisebelisoa tsa mofuta oa Rodent li tloaetse ho sebelisoa ho lekola litlamorao tsa lithethefatsi tse sebelisitsoeng hampe. Thutong ena, re tla tsepamisa maikutlo ho li-psychostimulants le li-opiates, kaha lipatlisiso tsa laboratori tse pharaletseng li tsepamisitse likarolo tsena tsa lithethefatsi. Tsamaiso e se nang khang ea li-psychostimulants kapa opiates e eketsa tšebetso ea locomotor ka litoeba (5). Lits'oaetso tsa lithethefatsi tse se nang liphapang tse phetoang khafetsa li ka lebisa keketseho e tsoelang pele le ea nako e telele mosebetsing ona o khothalletsoang ke lithethefatsi, ketsahalo e bitsoang boits'oaro ba boitšoaro (5). Ente e le 'ngoe ea koae ho litekanyetso tse phahameng le eona e khona ho tsosa maikutlo (6, 7). Ntle le moo, leha ho sa sebelisoe lithethefatsi, ts'ebetso ea locomotor e phahamisoa maemong a ts'oanang moo liphoofolo li fumaneng ente e le 'ngoe ea lithethefatsi ka letsatsi le fetileng (8). Liphetho tsena li bonts'a hore setsoalle pakeng tsa sethethefatsi le moelelo oa sethethefatsi se fumanehang se ithutoa ka potlako kamora ho pepesoa.
Lits'oants'o tse nang le lithethefatsi li na le tšusumetso e matla litabeng tsa boitšoaro ho batho ba nang le SUD (3). Nts'etsopele ea mokhatlo o lipakeng tsa lithethefatsi le litheko li ka hlahlojoa bathong ba laboratori (9, 10), hammoho le litoeba ka ho sebelisa mokhoa o ikhethileng oa sebaka (CPP) boits'oaro bo tšoanang (11). Ts'ebetso ena ea litoeba e kenyelletsa liente tse se nang liphapang tsa lithethefatsi ka phapusing e le ngoe le ho laola liente ka phapusing e haufi, empa e ikhethileng. Kgetho e lekanang pakeng tsa maemo a kopantsoeng ke lithethefatsi le taolo e tla hlahlojoa kamora nako ea tlhahlobo moo litoeba li ka kenang ka bolulo likamoreng tsohle tse se nang lithethefatsi (11). Ts'ebetso ea koetliso ea CPP e ka kenyelletsa mohato oa ho fela le liteko tsa ho nchafatsa (12, 13), e bontshang ho khaotsa ho sebelisa lithethefatsi le ho oela hape ho batho ba nang le bothata ba SUD. Ha li-paradigms tsa CPP li hlahloba ho ithuta ho akaretsang ho kenyelletsa sephetho sa ho tiisa, sepheo sa ho hloea sebaka (CPA) se lekola ho ithuta ho kenyelletsa sephetho se khathollang. Haholo-holo, li-paradigms tsa CPA li sebelisoa hangata ho ithuta maemo a mabe kamora ho tlosoa hoa lithethefatsi (14, 15).
Ho bonolo ho kenya ts'ebetsong maikutlong a boitšoaro le mekhoa ea CPP, empa li hloka liente tsa liteko tsa liteko. Liroto li ka koetlisoa habonolo hore li itlhahise ka lithethefatsi ka ho sebelisa catheter e kenang. Mekhoa e mengata ea ho itlhahisa ea lithethefatsi e ntlafalitsoe ho bontša matšoao a boits'oaro a batho a nang le SUD. Mohlala, litoeba tse nang le phihlello e lekanyelitsoeng (1 h) ho lithethefatsi lithupelong tsa letsatsi le leng le le leng tsa taolo li boloka tšebeliso ea lithethefatsi e tsitsitseng. Leha ho le joalo, litoeba tse nang le phihlello e atolositsoeng (6 h) ho lithethefatsi li eketsa ho jella ha tsona lithupelong tse ngata tsa koetliso, tse tšoanang le tšebeliso e mpe ea lithethefatsi e ka bonoang ho batho ba fumanoeng ba tšoeroe ke SUD (16-18). Joalo ka ha tšebeliso ea lithethefatsi e sa hlile e lebisa ho SUD, ha se mofuta o mong le o mong oa lithethefatsi o tla etsa mofuta o amanang le bokhoba. Ha litoeba li koetliselitsoe haholo ho sebelisa lithethefatsi (~ likhoeli tsa 3), sephutheloana sa litšobotsi se fumanehang ho batho ba nang le li-SUDs, joalo ka ho ts'oara lithethefatsi ho sa hlokahale, ho etsa boiteko bo boholo ho fumana ts'ebetso ea ts'ebetso ea lithethefatsi, le ho batla lithethefatsi ho sa natse litlamorao19). Liroto tse koetliselitsoeng ho itaola lithethefatsi le tsona li sebelisetsoa ho khutlela morao. Ho oela hape ho batho hangata ho laeloa ke lintlha tse tharo tse kholo: ho sebelisa lithethefatsi, ho pepesehela lits'ebetsong tse amanang le lithethefatsi kapa ho phela ka ketsahalo e sithabetsang (20-22). Lintho tsena tse ka u bakang (tšebeliso ea lithethefatsi, ho pepesehela litšebelisanong tse amanang le lithethefatsi, kapa khatello ea maikutlo) li ka khutlisetsa boits'oaro bo batlang lithethefatsi mefuteng ea taolo ea lithethefatsi le tsona (23).
Joalo ka ha ho na le batho ba nang le SUD, boits'ebetso bo itšetlehileng ka lithethefatsi litonong bo kenyelletsa karolo ea ho ithuta, ekaba ke maemo (maikutlo a boits'oaro, CPP, CPA, le ho khutlisetsoa hoa cue-induction kapa motho ea sebetsang le ona). Ha libaka tse ngata tsa boko li ameha ka ho ithuta ho ithutoang le boits'oaro bo amanang le lithethefatsi, re tla shebana le sebaka sa "ventral tegmental" (VTA) tekong ena. Re tla boela re tšohla lisupa tse kholo tsa VTA, hore na likhakanyo tsena li susumetsa ts'ebetso ea li-neuron ea VTA joang, le ho hlahisa liphumano tsa morao-rao mabapi le hore na barekisi ba VTA ba kenella joang boits'oarong bo itšetlehileng ka lithethefatsi.
Kopano ea VTA ho lithethefatsi tse tlisoang ke lithethefatsi
Li-neuron tsa dopamine tse tsoang ho VTA projeke eo ho isa bokong ba li-nucleus bokella ho kenya letsoho liketsong tse matlafatsang tsa lintho tse hlekefelitsoeng (24-26). Ha lithethefatsi tse hlekefelitsoeng li eketsa maemo a dopamine ho NAc (27, 28), lithethefatsi tse ngata tseo e seng tloaelo ea ho tloaela ha li ame ho ata ha dopamine (27). Psychostimulants e ama maemo a dopamine haholo-holo ka ho fetola tumello ea dopamine ho tloha sebakeng sa extracellular (29, 30), athe opiates e phahamisa ka kotloloho phetiso ea dopamine ka ho hatella phetisetso ea inhibitory ho li-dopamine neurons (31-33).
Potoloho ea neural e tsamaisang mokhoa ofe kapa ofe e rarahane, leha lipatlisiso tse ngata lilemong tse mashome a seng makae tse fetileng li bontša hore VTA e ameha ka mokhoa o makatsang boits'oarong bo botle bo khahlisang le bo sa thekeseleng lithethefatsi. Mohlala, VTA e hlokahala bakeng sa boitšisinyo ba boitšoaro bo susumetsoang ke amphetamine kapa mu-opioid receptor agonists, leha ho le joalo bopaki ba ho kenya letsoho ha VTA ho hohelang hoa boitšoaro ba koae ho kopantsoe (5). VTA e ameha hape ka CPP bakeng sa li-psychostimulants le li-opiates (34-39), hape le CPA e khothalelitsoe ke ts'ebetso ea kappa opioid receptor activation (15). VTA e boetse ea hlokahala bakeng sa ho khutlisetsoa khatello ea maikutlo, cue- le lithethefatsi lithethefatsing.23, 40-42kapa heroin (43-45). Le ha boitšoaro bo itšetlehileng ka VTA hangata bo kopantsoe ke li-dopamine neurons, bopaki bo ntseng bo eketseha bo bontša ho kenya letsoho ha li-neurons tsa VTA tse seng dopamine ho laola sephetho sa boitšoaro.
Baahi ba fapaneng ba Neuronal ka har'a VTA
VTA hammoho le "compacta" ea boahelani e hlahisang li-nuclei bokong (46). Lirekoto tsa elektronike tsa pejana li bonts'a hore VTA e ne e entsoe ka mefuta e 'meli e ikhethileng ea neuronal, e nkoang e le dopamine neurons le li-interneurons tsa GABA tsa lehae.31, 47). Leha ho le joalo, sephutheloana sa li-neurons tsa VTA se bonts'itse karabelo e ikhethileng ea electrophysiological ho serotonin le opioid receptor agonists, e fana ka bopaki ba ho ba teng ha palo e eketsehileng ea li-neuronal ho VTA (48). Bopaki bo felletseng lilemong tse leshome tse fetileng bo bonts'itse ho rarahana ha VTA ka bobeli mabapi le sebopeho sa neuronal le sepheo sa projeke.
Dopamine neurons e na le palo e kholo ea li-neuronal ka har'a VTA, ha tyrosine hydroxylase (TH), enzymme e fokotsang litekanyetso tsa dopamine synthesis, e fumanoa ho ~ 60% ea li-neurons tsa VTA (46, 49). VTA dopamine neurons ka tloaelo e nka sebaka se le seng feela sa sepheo, ka lipalo tse fapaneng tse kenang ho li-nuclei tse ngata tsa boko, ho kenyeletsoa NAc, dorsal striatum, cortex, amygdala, globus pallidus, le lateren habenula (LHb) (46, 50, 51). Leha ho le joalo, bopaki ba morao-rao bo bonts'a hore li-neurop tsa dopamine tse kenang ho NAc ea mori le tsona li romella li-dhamana ka ntle ho striatum (50). Ka setso, dopamine neurons le tsona li fumanoe li ipapisitse le thepa ea motlakase, ho kenyelletsa le ho ba teng ha nako e telele ea ts'ebetso ea triphasic, sekhahla se tlase sa ho thunya, ho thunya ho hoholo le ho ba teng ha Ih hona joale (52, 53). Leha ho le joalo, nako e ka 'nang ea nka nako e ka' na ea se lekane ho khetholla se ka hare ho neurotransmitter ea li-neurons tsa VTA (49, 54). Ho feta moo, li-neuron tse ngata tse kahare ho likarolo tsa bophelo tsa VTA li bile Ih empa o se na TH. Ha ts'ebetso e ka nka nako le Ih ha se kamehla e bonts'ang dopamine, thepa ena ea motlakase e ka amana le hore na projeke ea "VTA neurons" e sebetsa joang.55-57).
Baahi ba bobeli bo boholo ba li-neuronal ho VTA ba na le GABA neurons (~ 25%) tse tsebahalang hangata ka ho ba teng ha glutamic acid decarboxylase (GAD) (58, 59). Leha ho ne ho nahanoa hore e sebetsa hantle joalo ka li-interneurons tsa lehae (31), Li-neurons tsa VTA GABA li susumetsa ka ho toba tšebetso ea li-neurons tsa VTA dopamine (60, 61) hape le projeke ea "ventral pallidum" (VP), lateral hypothalamus (LH), le LHb, ka merero e nyane ho amygdala, preortal cortex (PFC), le NAc (62-64). Haufinyane, li-neuron tsa dopamine li ile tsa bonoa e le mohloli o eketsehileng oa GABA ho VTA, kaha li-neuron tsena li ka kopanya GABA ka tsela ea aldehyde dehydrogenase-Mediated (65). VTA le substantia nigra dopamine neurons sephutheloana sa GABA ka har'a li-vesicular ka transporter ea vesicular bakeng sa dopamine, e bonts'a hore GABA e ka kenngoa kelellong ka dopamine ho fumana litlamorao tsa li-electrophysiological ho li-neuron tse bohareng tsa spiny ka bobeli ba NAc le dorsal striatum (66, 67).
Ntle le li-neuron tsa dopamine le GABA, liperesente tse nyane tsa li-neuron tsa VTA li na le vesicular glutamate transporter 2 (VGluT2), letšoao bakeng sa li-neuron tsa glutamate. Li-neuron tsena li lula litulong tsa VTA ka mokhoa o hlophisehileng 'me li sebetsa ho "ventral striatum", PFC, VP, amygdala, le LHb, hape li oela ka har'a methapo ea methapo ea methapo (dopamine neurons).57, 64, 68-72). Subset ea li-neuron tse ntle tsa VGluT2 ho VTA le eona e bua ka TH mme e ka proa ho PFC le ventral striatum (70). Li-neuron tsena li lokolla dopamine ka bobeli le duramine (glutamate)73-77) leha li sa lokolloe sebakeng se le seng kapa ho li-vesicles tse tšoanang tsa synaptic (78). Le ha VTA ho ne ho nahanoa hore e entsoe feela ka li-neuron tsa dopamine le GABA, liphuputso tsa morao tjena li bonts'a hore VTA e entsoe ka li-neurone tsa dopamine tse ka rothisang Gaba, li-dopamine neurons tsa mantlha tse kang glutamate, li-neurons tsa GABA le li-glutamate neurons.
Optogenetic modulation ea li-neurons tsa VTA e ka etsa hore ho be le litakatso kapa liphello tsa boits'oaro bo fapaneng ho latela bongata ba neuronal bo shebiloeng. Ts'ebetso ea li-neurop tsa dopamine e tiisa haholo ebile e lekane ho theha CPP, athe ho khutsisa li-neurop tsa dopamine ke ntho e khopisang 'me e etsa hore CPA e sebetse.60, 79, 80). Ho matlafatsa li-neurons tsa VTA dopamine hape ho ntlafatsa boitšoaro bo matlafatsang mesebetsing e sebetsang (81-84). Ho fapana le moo, ts'ebetso ea li-neurons tsa VTA GABA e khethang e khelosa, e phahamisa CPA, 'me e fokotsa tšebeliso ea moputso ka ho thibela ts'ebetso ea li-neuron tsa VTA tsa dopamine tsa lehae.60, 61). Ho khahlisang, ho kenya tšebetsong li-neurons tsa VTA GABA tse arohanang le mahlakore a machinuric ho NAc ho matlafatsa khethollo pakeng tsa sepheo se seng molaong le se phephetsang (63). Ts'ebetso ea Optogenetic ea li-neurons tse nang le VGluT2 ho VTA e boetse e lekane ho theha CPP, e leng phello e kopantsoeng le ho kenya tšebetsong methapo ea kutlo ea VTA ea dopamine ea lehae (72). Ka kopanelo, lithuto tsena li fana ka maikutlo a hore litlamorao tsa boitšoaro bo kopaneng le VTA, ho kenyelletsa le boits'oaro bo itšetlehileng ka lithethefatsi, li ka kenyelletsa puisano e rarahaneng pakeng tsa palo e fapaneng ea li-neuronal ho VTA.
Taolo e fapaneng ea VTA
VTA e na le litlatsetso tse ngata tse fapaneng, tse ngata li hokahane. Liphatlalatso tse kholo ho VTA li kenyelletsa "rostromedial tegmental nucleus" (RMTg), VP, nucleus ea "stria terminalis" (BNST), LH, "patunculopontine tegmental nucleus" (PPT), laterodorsal tegmental nucleus (DR), NAc , PFC, le Amygdala (50, 85-87). Ha li-dopamine tsa VTA le li-Gaba li hlahisoa ke likarolo tse ngata tsa boko bo tšoanang (50), ha ho tsejoe hanyane ka maiketsetso a tsoang ho VGluT2 neurons e ntle ho VTA. Ka tlase, re tla tšohla hore na lintho tse hlahelletseng tse tsoang ho VTA li ka susumetsa tšebetso ea li-neurons tsa VTA joang, hore na lintho tse kenang li ka susumetsa boitšoaro bo itšetlehileng ba VTA joang, le liphumano tsa moraorao ho barekisi ba VTA ba amehang ka boits'oaro bo its'ebetsong ba lithethefatsi.
Rostromedial Tegmental Nucleus
RMTg (eo hape e bitsoang mohatla oa VTA) ke khubu e kopantsoeng le li-neurons tsa GABA tse sebetsang joaloka kamano ea khokahano pakeng tsa LHb le VTA (86, 88-92). Lesions tsa RMTg li bonts'a karolo e mahlonoko bakeng sa tikoloho ena ea boko ho fetoheng boits'oaro bo fapaneng (86). Ntle le moo, li-neurons tse ho RMTg li kentsoe tšebetsong ke maikutlo a ho hlohlona hape a thibeloang ke meputso (86). RMTg e susumetsa haholo ho thunya ha li-neuron tsa VTA, kaha RMTg inactivation e eketsa dopamine neuron ho thunya (93), athe e hlasimollang RMTg e kenyelletsa dopamine neuron ho thunya (93-95).
RMTg e ntse e hlomphuoa le ho feta e le senotlolo sa bohlokoa sa ho loants'a litlamorao tsa lithethefatsi tse hlekefelitsoeng. Matla a matlafatsang a "opiates" qalong a ne a nahanoa hore a ka hlaha ts'ebetsong ea li-mu-opioid receptors tsa li-Intneurons tsa VTA GABA (31), leha bopaki bo bokellang bo bonts'a sepheo sa mantlha sa li-opiate ho fapana le liphatlalatso tsa RMTg ho VTA (33, 96, 97). Tsamaiso ea morphine e fokotsa ho thunya ha sele ka RMTg, e fokotsang thibelo ho li-neuron tsa VTA dopamine, e hlahise ho phahamisa dopamine neuron ho thunya (94-96). Kannete, ts'ebetso e khethiloeng ea li-mu-opioid receptors ho li-neuron tsa RMTg tse hlahisang VTA e lekane bakeng sa ho khetha khetho ea sebaka sa nnete (98). Ka mor'a ho khaotsa opiate, ho thibela li-neurons tsa RMTg ha ho sa phahamisa ho thunya ha VTA dopamine neuron. Ho sitoa hona ha RMTg ho disinhibit dopamine neurons ho aroloa ka karolo ka ho fetoloa hoa molumo oa lentsoe la VTA glutamatergic (93). Ha morero oa RMTg ho VTA o buisana le litlamorao tse matlafatsang tsa opiates (33, 96, 98), mekhoa e meng e amanang le VTA e kopanetsoeng le mamello ea dopamine neuron ho li-opiates tse latelang ho tlosoa (93).
Psychostimulants e boetse e susumetsa ts'ebetso ea li-neuron tsa RMTg (94). Ts'ebetso ea cocaine e seng e nang le khang e phahamisa maemo a Fos, e leng taba e amanang le ts'ebetso e amanang le "neuronal" ho "RMTg neurons"99, 100). Ho khahlisang, maemo a Fos ho RMTg neurons e hlahisang VTA a phahamisoa kamora ho felisoa ha likhoto tse ipehelang cocaine (101). RMTg e boetse ea hlokahala bakeng sa boits'oaro bo amanang le koae bo bonoang hang ha bo hlahisa litholoana tse ntle tsa koae.102). Teko e tsoelang pele e hlokahala ho netefatsa hore na morero oa RMTg ho VTA o ameha ka boits'oaro bo matla le bo matlafatsang bo hlahisoang ke koae.
Ventral Pallidum
VP e kenella ho sebetsana le boits'oaro bo khotsofatsang le boits'oaro bo susumetsang (103). Li-neurons tsa GABA ho VP li fana ka mohloli o moholo oa tlhahiso ea thibelo ho VTA (87, 104). Ho kenya tšebetsong li-terminals tsa VP ho etsa hore maqhubu a thibelo ea methapo ea methapo ea methapo ea methapo ea methapo ea methapo e fete.105). Phello e sebetsang ea ho nolofalletsa VP e baka keketseho ea mosebetsi oa baahi ho li-neuroprophene tsa dopamine tse behang (106) leha ho le joalo phello ea li-neurons tse se nang dopamine VTA ha e tsejoe. Ho na le bopaki bo bongata bo amang VP ka boits'ebetso ba ts'ebeliso ea lithethefatsi. Li-neuron tsa VP tse kenang ho dopamine le li-neuron tse se nang dopamine li thibetsoe haholo ke opiates (105). Ntle le moo, marang-rang a VP kapa manong a thehiloeng ho lithethefatsi ho VP a ka thibela maikutlo a susumetsang a morphine (107, 108), CPP e susumetsoang ke lithethefatsi (35, 109, 110)111), le ho khutlisoa (40, 41, 112). Li-neuron tsa VP tse phatlalalitsoeng ho VTA ke li-Fos tse buoang kamora ho khutlisetsoa hoa cueine ka ho feteletseng bakeng sa cocaine (101) le ho khutsisa methapo ena ho lekane ho thibela ho khutlisetsoa hoa moea oa cue-ikiwa (113). Ha li-neuron tsa VP li qala ho sebetsa ka bobeli ho dopamine le li-neuron tse se nang dopamine ho VTA (105), ha ho hlake hore na ke li-neuronal (s) life tsa VTA tse susumetsoang ke lipehelo tsa VP nakong ea boits'oaro bo its'epahallang ke lithethefatsi.
Nucleus ea bethe ea Stria terminalis
BNST e kenella ho loantšeng tšabo le matšoenyeho (114-120) mme e nkuoa e le khubu ea tumellano lipakeng tsa khatello le litsela tsa moputso (121, 122). Sebopeho sa neuronal sa BNST se fapane, se na le lipalo tse sebetsang tsa GABA le li-glutamate neurons hammoho le li-interneurons tsa GABA le cholinergic tsa lehae (122, 123). Li-neurons tsa BNST li boetse li bontša assortment ea li-neuropeptides tse kenyeletsang neuropeptide Y, corticotropin-releasing factor, enkephalin, dynorphin, le ntho P (124). Ho susumetsa ka motlakase ha BNST ho ba le tšusumetso e thabisang ho li-neuron tsa dbamine tsa midbrain (122, 125, 126) hape e phahamisa tokollo ea dopamine ho NAc (127). Boithuto ba morao-rao bo bonts'a hore phello ena e thabisang ho li-dopamine neurons e kenella ka ho khetheha ka li-neurons tsa GABA BNST tse thibelang li-neurons tsa VTA GABA, e hlahisang liphello tsa ho tšoenyeha le ho putsa boitšoaro (128-130). Ho khahla, "glutamate neurons" ho BNST ho boetse ho na le li-neurons tsa VTA GABA, 'me ts'ebetso ea methapo ena ea kutlo e thusa ho etsa lintho tse khahlisang le tse tšoenyang (129). Tlas'a maemo a boitšoaro bo itšetlehileng ka ts'ebeliso ea lithethefatsi, mananeo a meriana ea lehae a bontša karolo ea bohlokoa ea BNST ho khutlisetsong ho matla hoa khatello ea lithethefatsi (41, 131, 132). Ntle le moo, liphuputso tsa morao tjena li ama tsela ea BNST-VTA litlamorao tsa ts'ebetso ea cocaine (133) le ka polelo ea cocaine CPP (134), leha ho kenyelletsoa ho kenyelletsa tsela ena mekhoeng e meng e amanang le lithethefatsi.
Hamorao Hypothalamus
LH e bohlokoa bakeng sa ho hlahisa mekhoa e susumetsang e kenyelletsang ho fepa le ho batla lithethefatsi (135). LH e fana ka litlatsetso tsa glutamate le tsa GABA ho VTA (85, 136). Ntle le moo, li-neuron tsa LH tse hlahisang VTA le tsona li na le li-neuropeptide tse kang neurotensin le orexin / hypocretin (137, 138). Ts'usumetso ea motlakase ea LH e eketsa ts'ebetso ea li-neuropropathy tsa dopamine mme e thibela ts'ebetso ea li-neurons tsa GABA tse tlatsitsoeng ho VTA (139). Bopaki bo bongata bo bonts'a hore ts'ebetso ea tsela ena ea LH-VTA ea matlafatsa. Liroto li tla itlhahisa habonolo bakeng sa ts'ebetso ea motlakase ea LH, empa ts'ebetso ena ea boitšoaro e thibeloa ke dopamine receptor antagonism (140) kapa inactivation ea VTA (141). Ho feta moo, ts'ebetso ea optogenetic ea ts'ebetso ea LH ho VTA e boetse e ts'ehetsa ho intlafatsa ka mochine o itšetlehileng ka neurotensin (142).
Bopaki bo felletseng ka lilemo tse leshome tse fetileng bo totobatsa bohlokoa ba li-neuron tse nang le orexin ho fepa, potoloho ea ho robala / ho tsosa, le boitšoaro bo amanang le lithethefatsi (143). Li-neuron tse hlahisang Orexin li fumaneha feela kahare ho hypothalamus mme li sebetsa hohle ho bokong (144), leha ho le joalo e le projeketso ea VTA e amanang haholo le boits'oaro bo its'epahalletseng lithethefatsi. Li-entra-VTA tsa Intra-VTA tsa li-antillonists tsa receptor li fumana morphine CPP (145, 146), e lumellanang le ts'oaetso e fokotsehileng ea morphine e bonoang ho litoeba tse haellang tsa orexin (147). Ka lehlakoreng le leng, tsamaiso ea intra-VTA ea orexin e khutlisetsa morphine CPP (12). Bohanyetsi ba Orexin bo shebileng VTA le bona bo fokotsa ts'ebetso ea maikutlo ho cocaine (148), khokahano ea koae149), le ho khutlisetsoa hoa morao-rao (150). Ho khahlisang, li-neuron tsa orexin ho LH le tsona li na le dynorphin, e thibelang tšebetso ea VTA dopamine neurons. Phuputso ea morao-rao e bonts'a hore orexin ho VTA e nolofalletsa boits'oaro bo amanang le lithethefatsi ka karolo ka ho fumana litlamorao tsa dynorphin (149). Leha li-neuron tse nang le orexin ho LH li amohetse tlhokomelo e kholo maemong a bokhoba, litla-morao tsa li-neuronal tse tseleng ea LH-VTA le tsona li ka ba le seabo litloaelisong tse amanang le lithethefatsi, joalo ka li-neuron tse sa hlahiseng orexin ho LH. Fos e kenelletseng kamora ho khutlisetsoa kahare hoa cue101).
Nyutlelie ea Motsoallo oa Metsoalle ea Afterodorsal le Nunus ea Tebo ea Pedunculopontine
LDT le PPT li kentse letsoho ho fetoleng boits'oaro bo khahlisang le bo tsamaisang moputso (92, 151-154). Li-nuclei tsena li entsoe ka mefuta e fapaneng ea acetylcholine, GABA, le li-glutamate neurons tse sebelisang tsamaiso ea methapo ea methapo ea mmele (midbrain dopamine system)155, 156). Boithuto ba anatomical bo bonts'a hore VTA ka kotloloho e amohela tlhahiso ho tsoa ho LDT (87, 155, 157). Ka vivo liteko tsa electrophysiological li bontša hore tšusumetso ea motlakase ea LDT e baka ho thunya ho hoholo ha VTA dopamine neurons (158). Ts'ebetso e khethiloeng ea ho kenella ha LDT ho VTA e etsa hore maqhubu a thabisang a VTA dopamine neurons a qale ho NAc ea morao-rao (92). Ho susumetsa tsela ena ea LDT-VTA Ka vivo e fumanela CPP le ho matlafatsa ts'ebetso e arabelang (92, 154). Bopaki bo ntseng bo eketseha bo bontša hore LDT e boetse e kenella mekhoeng e amanang le lithethefatsi. Haholo-holo, manokotsoana a k'hemik'hale a moo a bonts'ang LDT e bohlokoa bakeng sa ho fumana le ho hlahisa cocaine CPP (159), hammoho le ho khutlisetsoa hoa koae ka lebaka la koae160). Ho khahlisang, li-neuron tsa cholinergic tsa LDT li ameha le karabelo ea boitšoaro ho lithahasello tse tsamaeang le koae (161). Ho hlokahala lithuto tse ling ho netefatsa hore na boits'oaro bo itšetlehileng ka lithethefatsi bo boetse bo kenyelletsa GABA le lipakanyo tsa glutamate ho tloha LDT ho ea VTA.
Le ha VTA e khethiloe ka mokhoa o khethiloeng ke LDT, PPT eona e shebile haholo nigra (87, 155). Le ha bopaki ba anatomical bo bontša hore ho na le projeke e nyane ea PPT ho VTA (87, 155), lithuto tsa electrophysiological Ka vivo 'me a vitro fana ka maikutlo a kamano e sebetsang pakeng tsa PPT le VTA (106, 162, 163). Phapang pakeng tsa lithuto tsa anatomical le electrophysiological ha e hlake, empa litlhaloso tse reriloeng li kenyelletsa monyetla oa hore neurone e le 'ngoe ea PPT e be le li-neurons tse ngata tsa VTA kapa hore ts'usumetso ea motlakase e thabisa likhoele tsa likarolo kapa libaka tse haufi, joalo ka LDT (87). Ho sa tsotelehe, lits'usumetso tsa motlakase tse lebisang ho PPT li eketsa ho thunya ha "li-dopamine neurons tsa maiketsetso tsa"106), ha ts'ebetso ea PPT e fokotsa ts'ebetso ea dopamine neuron ho thunya ho sesapo sa maikutlo (162). PPT e ameha hape boits'oarong bo itšetlehileng ka lithethefatsi, ha lesapo attenuate amphetamine- le morphine-ikiwa le ts'ebetso ea locomotor (164), le ts'ebeliso ea PPT e fokotsa ho ts'oaroa hoa lithethefatsi tsa koae (cocaine)160). Lapo ea PPT e fokotsa ho itaola le ho itaola ka morphine CPP (165, 166). Leha ho le joalo, li-neurons tsa PPT cholinergic ha li amehe ka ho iphelisa ka koae, ho itaola ka heroin, CPP ea cocaine, le heroin CPP (167), ho fana ka maikutlo a ho nka karolo ha li-PPT glutamate le / kapa li-neurons tsa GABA litloaelong tsena tse amanang le lithethefatsi.
Dorsal Raphe
DR ke mohloli o ka sehloohong oa serotonin bokong, empa hape e na le glutamate (85), GABA (168), le li-neurone tsa dopamine (169). Le ha DR hangata e ithutoa maemong a ho laola maemo a amanang le ona (170), e boetse e kenella ho matlafatseng boits'oaro ba sebopeho (171). Serotonin e fana ka likarabo tse fapaneng tsa electrophysiological ho li-neurons tsa VTA. E hlahelletseng a vitro Karabelo ho "putop dopamine neurons" e ea thabisa, leha karolo e nyane ea li-dopamine neurons e thibetsoe ke serotonin (172). Ka lehlakoreng le leng, lipalo tse lekanang tsa li-neurons tsa GABA tse behang li thabile ebile li thibetsoe ke serotonin (172). Litlamorao tsa likarabo tsena tsa motlakase li bonahala li thabisa, joalo ka Ka vivo Tsamaiso ea intra-VTA ea serotonin e phahamisa maemo a dopamine ho NAc (173).
Serotonin e susumetsa boitšoaro bo amanang le lithethefatsi (174), e ka kenyelletsang serotonin neurons ea projeke ea VTA. Leha ho le joalo, monahano oa DR ho VTA ha e le hantle o entsoe ka li-neuron tse glutamate tse ka hare li etsang dopamine neurons (85, 87, 175). Ts'ebetso ea li-neurons tsa "gl glamamate" e tsosa maqhubu a thabisang ho VTA dopamine neuron le elicits dopamine e lokolloe ho NAc (175). Ts'ebetso e ikhethileng ea tsela eo e seng ea serotonergic DR-VTA e tiisa boitšoaro ba ts'ebetso mme e lekane ho eletsa CPP (175, 176). Ka lehlakoreng le leng, ts'ebetso ea serotonergic DR neurons e phatlalalitsoeng ho VTA e matlafatsa feela ka mokhoa o fokolang (176). Liphumano tsena tsa boithuto ba boithuto le boits'oaro li fana ka maikutlo a hore VTA ha se mohlomong e leng sebaka sa mantlha moo serotonin e sebetsang ho susumetsa boitšoaro bo amanang le lithethefatsi. Sebakeng seo, li-neurons tsa non-serotonergic DR tse hlahisang VTA li boemong bo botle ba ho buisana le boits'oaro bo itšetlehileng ka lithethefatsi, leha sena ha se so ka se hlahlojoa ka liteko.
Nucleus Accumbens
Li-neurons tsa GABA morerong oa NAc ho VTA 'me ho nahanoa hore li thusa ho fana ka karabo ea "long-loop" ho laola mosebetsi oa dopamine neuron (177). Mu-opioid receptor agonists e thibela lithibelo tsa GABA ho tloha NAc ho ea ho VTA (33, 178). Phetiso ea inhibitory e tsoang liphatsong tsa NAc ho li-neurons tsa VTA GABA e matlafatsoa kamora liente tse pheta-phetoang tsa koae, eo ka lehlakoreng le leng e thibelang li-neurons tsa VTA dopamine (179). Ntle le ho susumetsoa ke opiates le psychostimulants, li-AUc tse amanang le VTA li Foselitsoe nakong ea ho khutlisetsoa hoa koae (ea koae).101). Leha liphetho tsena li bontša tsela ea NAc-VTA e kentse letsoho lits'ebetsong tse amanang le lithethefatsi, ha ho liteko tse etsoang ho fihlela joale tse ileng tsa hlahloba sephetho sa boitšoaro ba ho senya tsela ena ka boomo.
Pele Cortex
PFC ea meea e buisana ka mefuta e mengata ea mesebetsi e lemohileng (180, e nka karolo ho khutlisetsong hoa boitšoaro ba ho batla lithethefatsi (23), hape e bonts'a ts'ebetso ea Fos e latelang taolo e mpe ea amphetamine (181). VTA e amohela moroalo oa "glutamate" o tsoang ho morusu PFC (85), ka li-neuron tsa "pyramidal" li kenella ho li-neurone tsa dopamine le tse e seng tsa dopamine (62, 182). Ho hlohlelletsa PFC ka motlakase ho ka thibela kapa ho thabisa li-neurop tsa dopamine tse kahare ho VTA (183, 184). Le ha ho hlohlona ho le joalo kapa ho ts'oaroa ha PFC hangata ho thibela boholo ba li-neuron tsa VTA dopamine (183-185), Ho phatloha ho phatlohileng ha PFC hoa thabisa> 90% ea VTA dopamine neurons (184). Mochine o tšehetsang dopamine neuron e sa hlahiseng letho ha o hlake, hobane li-neurons tsa VTA dopamine li fumana ho kenella habonolo ho tsoa ho PFC (87, 186), e nang le <15% ea VTA dopamine neurons e khahliloeng ke ho khetha ho kenya letsoho lipakeng tsa lipallo tsa PFC50). Liphumano tsena ka kopanelo li bonts'a sepheo sa li-neuron tsa VTA GABA ka mokhoa o ikhethileng, leha ho le bohlokoa ba tsela ea PFC-VTA boits'ebetsong bo ts'episang lithethefatsi ha e so ka e hlahlojoa.
Amygdala
Amygdala ke sehlopha se kopaneng sa li-neri tse amehang ka ho hlahisa boleng ba maikutlo ho li-cue (187, 188). VTA e fumana mohopolo oa amygdala o hlahang ho khubu e bohareng ea karohano ea karolo ea amygdala (CeA) (87, 189). CeA e na le li-neurons tsa GABA boholo mme e ameha maemong a tšabo (187, 188, 190), hammoho le ho hanana le tšusumetso e akaretsang ea tšusumetso e khahlisang (191, 192). Boemong ba boits'oaro bo itšetlehileng ka lithethefatsi, CeA e thusa ho fana ka karabelo ea maemo a tlase (193) hape e ameha ka ho khutlisetsa khatello e matlafatsong e bakang khatello ea maikutlo ea ho batla lithethefatsi (194, 195). Le ha merero ea CeA ho VTA, hajoale ha ho tsejoe hore na tsela ena e susumetsa ts'ebetso ea neuron ea VTA le hore na e bohlokoa bakeng sa boits'oaro bo lemaletseng lithethefatsi.
Tlhatlhobo e hlahisoang ke lithethefatsi ea Synaptic Plasticity ho VTA Neurons
Phetoho ea motho ea tsoang litulong tse se nang kelello tsa lithethefatsi kapa ea lithethefatsi tse tloaelehileng ho ea ho li-SUD e kenyelletsa liphetoho mosebetsing oa li-circuits tse khethehileng tsa neural (196). Ha ho fanoa ka bohlokoa ba VTA ka boits'oaro bo amanang le lithethefatsi, liphetoho tse feto-fetohang ho VTA dopamine neuron li ithutile haholo le ho hlahlojoa libakeng tse ling (197-201). Boithuto bo bongata bo tsoang litsing tse fapaneng tsa laboratori bo bontšitse keketseho ea matla a nyakallo a synaptic ho li-neurons tsa VTA dopamine ka mor'a Ka vivo ho pepesehela lithethefatsi tse hlekefelitsoeng (202-208). Bongata ba lithuto tsena li ile tsa hlahloba phello ea lithethefatsi ho karolelano ea li-receptor tsa AMPA hona joale li-receptor ea NMDA ea hona joale (AMPA / NMDA) ho li-neurons tsa VTA, tse lumellang ho bapisa matla a nyarosang a synaptic pakeng tsa lihlopha tse fapaneng tsa liphoofolo (ke hore, lithethefatsi tse phekoloang vs. . laola). Ka vivo ho pepesehela lithethefatsi tsa tlhekefetso ho eketsa AMPA / NMDA (202-204, 206, 207), e kopantsoeng ka ho kenyelletsa li-receptor tsa calcium AMA tse amoheloang le ho tlosoa hoa li-receptor tsa NMDA ho li-dopamine neurons tsa VTA (205, 208).
Ntle le liphetoho tse thabisang tsa synaptic ho VTA dopamine neurons, Ka vivo ho pepesehela lithethefatsi ho boetse ho fetola meetso ea inhibitory synaptic ho VTA. Mohlala, liente tse pheta-phetoang tsa "cocaine potentiate" ea NAc inhibitory ea ho kenya li-neurons tsa VTA GABA, e hlahisang tlhahiso ea li-dopamine neurons (179). Disinhibition ena e boetse e nolofalletsa bokhoni ba ho tsosa bokhoni ba nako e telele ba boipheliso (LTP) ho VTA dopamine neurons (209). VTA dopamine neurons le eona e khona ho feta ka inhibitory ea PDP. Ntle le moo, LTP ena ea thibelo e thibetsoe ho latela an Ka vivo ho pepesetsoa opiates (210, 211). Ho tlalehiloe tse ling tse ngata tsa liphetoho tse fetotsoeng ke lithethefatsi, leha ho le bohlokoa ho tseba hore tlatsetso e felletseng ea liphetoho tsa motlakase le hore na nako ea liphetoho tsena ho li-neurons tsa VTA ho latela moriana, tekanyetso ea lithethefatsi, le mokhoa oo motsoako o laeloang ka oona (202-204, 206, 207, 212). Ke lithuto tse fokolang ho fihlela joale tse hlahlobisitseng hore na liphetoho tsena tse fetotsoeng ke lithethefatsi li etsahala ka tsela e ikhethileng (179, 212). Ka 'nete, Ka vivo ho pepesehela litlelase tse fapaneng tsa lithethefatsi tse hlekefelitsoeng ho fella ka liphetoho lipakeng tsa litlatsetso tse hlakileng tsa VTA dopamine neurons (212). Leha ho ithutile ho hongata mabapi le liphetoho tse feto-fetohang ho VTA kamora liente tse se nang khang tsa lithethefatsi tse hlekefelitsoeng, lithuto tse ling li hlokahala ho netefatsa ho tšoana le ho se tšoane ha liphetoho tse fetotsoeng ke lihlopha tse fapaneng tsa lithethefatsi tse hlekefelitsoeng (psychostimulants, opiates, joala, nikotini, jj.). Ntle le moo, ho boetse ho hlokahala lithuto tsa electrophysiological ho tseba hore na ke li-VTA life tse amanang le tsona le hore na ke lipalo life tsa methapo ea methapo ea methapo ea methapo tse tsamaeang le phetoho ea lithethefatsi tse latelang.
fihlela qeto e
Tekanyo e phahameng ea ho khutlela morao e bontša tlhokahalo ea ho khetholla mekhoa e mecha ea kalafo bakeng sa kalafo ea li-SUD. Phekolo ea ho itšetleha ka opioid e thatafatsoa ke matšoao a matla a tlisoang ke batho ha a khaotsa ho sebelisa lithethefatsi. Likhetho tsa hona joale tsa kalafo bakeng sa li-SUD tsa opioid hangata li shebana le tlhokomelo ea opioid ka methadone kapa buprenorphine le detoxification le alpha-2 receptor agonists. Leha ho le joalo, likhetho tsa hajoale tsa kalafo hangata li baka ho khutla hape (213). Hajoale ha ho na pharmacotherapy e amoheloang ke FDA bakeng sa kalafo ea li-SUD tsa koae N-acetylcysteine ke lithethefatsi tse tšepisang le tse mamelloang hantle tse fokotsang ho batla koae ka koae le ho lakatsa ho batho ba itšebelitseng koae.214-217). Lilemong tse leshome tse fetileng, lipatlisiso mabapi le kalafo e sebetsang ea meriana bakeng sa li-SUD tsa joala li sibollotse liphofu tse ngata tse ka bang teng, ho kenyelletsa le li-receptor tsa opioid (218), li-receptor tsa dopamine (219), li-receptors tsa glutamate (220), Li-receptors tsa GABA (221), le li-receptor tsa adrenergic (222). Patlisiso e entsoeng esale pele e totobalitse sistimi ea cannabinoid e le sepheo se ts'episang sa li-SUD tse ngata (223, 224). Leha ho le joalo, tlhahlobo ea bongaka ea pelo le pelo e hlahlobang ho sebetsa ha rimanobant, mohanyetsi oa "cannabinoid receptor antagonist"225) hape e fokolitse cheseho ea ho shebella sisteme ea endocannabinoid bakeng sa ho phekola li-SUDs. Ka bomalimabe, ha ho na pharmacotherapy e le 'ngoe eo hona joale e leng teng bakeng sa ho phekola bongata ba li-SUD.
Tsela e 'ngoe ea kalafo e ka sebelisoang bakeng sa kalafo ea li-SUD e kenyelletsa tšebeliso ea matla a tebileng a tšusumetso ea kelello (DBS), e sebelisitsoeng hangata bakeng sa kalafo ea mathata a ho sisinyeha ha mmele. Lithutong tsa preclinical, DBS e shebaneng le NAc e fokotsitse kutlo ea boitšoaro ea koae (226), morphine CPP (227), ho khutlisetsoa morao hoa ho batla ha heroine (228) le ho khutlisetsoa ha motho ea batlang koae (ea koae)229-231). Ntle le moo, DBS e shebaneng le LHb e fokotsa taolo ea koae le ho khutlisetsoa ha motho ea batlang koae (koae)232). E lumellana le liteko tsa litheko tsa DBS tse ngata, lithuto tsa tleliniki li bontša ho hlakoloa ka botlalo kapa tšilafalo e telele ea tšebeliso ea heroin kamora DBS ho NAc bathong (233, 234). Tlokotsi e kholo ea ho kenella tšebetsong DBS ho batho ke mofuta o hlaselang oa ho kenya projeke ena. Leha ho le joalo, litlaleho tse 'maloa tsa morao-rao li bontša hore tšusumetso e sa hlaseleng ea matla a motlakase a PFC e sebetsa hantle ho fokotsa tšebeliso ea lithethefatsi le litakatso (235, 236). Le ha ho ntse ho na le mekhoa e mecha e khothatsang ea kalafo bakeng sa ho phekola li-SUDs, sepheo sa mantlha sa ts'ebetso efe kapa efe ke ho sebetsa hantle le ka kotloloho kamoo ho ka khonehang ho fokotsa litla-morao. Kahoo, ho hlokahala lipatlisiso tsa mahlale a mantlha tsa mahlale bakeng sa ho khetholla mekotla e ikhethileng ea neural le liphetoho tse ikarabellang bakeng sa nts'etsopele ea boitšoaro bo its'etsahallang ke lithethefatsi.
Ts'ebetso ea mekhoa ea optogenetic le chemogenetic litekong tsa boits'oaro e netefalitse le ho khetholla li-circuits tse khethehileng tsa maqhubu tse arolelanang mefuta e fapaneng ea boits'oaro le boits'oaro bo khahlisang. Boholo ba lithuto tsena li fetotsoe ka bokong ba SUDs (237), leha e le ba fokolang ba fetotseng lipotoloho tsa neural ka har'a maemo a boitšoaro bo its'etsahallang ke lithethefatsi (98, 113, 133). Ha ts'ebetso kahare ho VTA e le karolo ea boits'oaro bo bongata ba ho latela lithethefatsi, lipotso tse ngata li ntse li le teng. Liteko tsa nako e tlang li hlokahala ho (i) ho khetholla hore na li amanang le VTA le hore na ho na le liphoofotsoana life tsa VTA tse buisanang le boitšoaro bo itseng bo amanang le lithethefatsi le (ii) ho hlakisa liphetoho tse amanang le li-synaptic tse amanang le tsona ka bobeli mabapi le dopamine le li-neuron tse se nang dopamine kahare ho VTA. Ho khetholla mekotla ea neural le li-adaptation tse ikarabellang bakeng sa boits'ebetso bo itšetlehileng ka lithethefatsi ka litoeba li ka totobatsa li-circuits tse khethehileng tsa neural bakeng sa ts'ebetso e kenelletseng ea kalafo ea meriana le DBS ho phekola batho ba nang le bothata ba SUD.
Menehelo ea Mongoli
MW le IO ba kentse letsoho ho ngoloa ha sengoloa sena sa tlhahlobo.
Tlhōlisano ea Tlhaloso
Bangoli ba bolela hore lipatlisiso li ne li etsoa ka ho se be le likamano leha e le life tsa khoebo kapa tsa lichelete tse ka nkoang e le khohlano e ka 'nang ea e-ba le thahasello.
dithuso tsa ditjhelete
Mosebetsi ona o tšehelitsoe ke National Institutes of Health Grant DA033386 (MW).
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Mantsoe a bohlokoa: VTA, mathata a ts'ebeliso ea lithethefatsi, lithethefatsi, dopamine, polasetiki
Caring: Oliva I le Wanat MJ (2016) Barekisi ba Sebaka sa Phallo ea Likolo tsa Ventral le Baetsi ba Lithethefatsi. Ka pele. Psychiki 7: 30. Doi: 10.3389 / fpsyt.2016.00030
E amohetse: 15 December 2015; E amohetsoe: 23 February 2016;
E hatisitsoe: 07 March 2016
E hlophisitsoeng ke:
Mark Walton, Univesithi ea Oxford, UK
Hlahloba by:
Giovanni Martinotti, Univesithi G. d'Annunzio, Italy
Miriam Melis, Univesithi ea Cagliari, Italy
Elyssa Margolis, Univesithi ea California San Francisco, USA