Tlhophiso ea mantlha ea li-neurotransmitters mabapi le boitšoaro ba ho batla meputso (2014)

Oscar Arias-Carrión,^1,2 Xanic Caraza-Santiago,¹ Sergio Salgado-Licona,¹ Mohamed Salama,³ Sergio Machado,^4,5 Antonio Egidio Nardi,⁴ Manuel Menéndez-González,⁶ 'me Eric Murillo-Rodríguez⁷

Sebaka sa "ventral tegmental" se amana haufi le tsamaiso ea moputso. Dopamine e lokolloa libakeng tse kang li-nucleus accumbens le preortal cortex ka lebaka la liphihlelo tse khotsofatsang tse kang lijo, thobalano le maikutlo a ho se nke lehlakore a amanang le tsona. Ho susumetsa ka motlakase hoa sebaka sa ho teba ha sepheo sa motlakase kapa litsela tsa sona tse tsoang ka boeona e ka ba moputso o matla. Lithethefatsi tse fapaneng tse eketsang maemo a dopamine lia putsoa ka mokhoa o hlollang. Le ha dopaminergic system e emela sekhahla sa sistimi ea moputso, li-neurotransmitters tse ling tse kang li-opioids tsa endo native, glutamate, γ-Aminobutyric acid, acetylcholine, serotonin, adenosine, endocannabinoids, orexins, galanin le histamine kaofela li ama sisteme ena ea mesolimbic dopaminergic. Ka lebaka leo, liphoofotso tsa liphatsa tsa lefutso tse amanang le methapo ea kutlo li na le tšusumetso ea ho lokisa moputso oo ka nako e 'ngoe e ka amang boitšoaro bo ikhethileng ba sechaba le ho ba le ts'oaetso ea ho lemalla. Mona, re buisana ka bopaki ba hajoale ka taolo e tloaelehileng ea li-neurotranmitters tse fapaneng mabapi le boitšoaro bo batlang meputso le phello ea eona e ka bang teng tlatsong ea lithethefatsi.

Lithethefatsi le moputso oa tlhaho

Taba e le 'ngoe e hlokang ho netefatsoa ke hore na lithethefatsi le meputso ea tlhaho li etsa hore li-neuron tse tšoanang li sebetse hantle. Leha ho na le ho na le pharalla libakeng tsa boko tse anngoeng ke moputso oa tlhaho le lithethefatsi tsa tlhekefetso [4], phallo e ts'oanang ho lipalo tsa neural tse anngoeng ke meputso ea tlhaho le lithethefatsi li ke ke tsa tiisoa [5,6]. Motheong oa data e fetileng, na re ka utloisisa ho lemalla ha lithethefatsi ka ho ithuta ka moputso oa tlhaho? Bopaki ba morao-rao bo supa hore ho pepesehela meputso e meng eo e seng ea lithethefatsi e ka "u sirelletsa" melemong ea litheko tsa lithethefatsi. Mohlala, tsoekere le saccharin li ka fokotsa ho itlhomme pele ha cocaine le heroin [7].

Liphuputso tse 'maloa li senotse hore tlhekefetso ea lithethefatsi hangata e qala ka ho eketsa phaello ea batho meputsong ea tlhaho (sensitization). Hamorao phaello ena e fokotseha ka tšebeliso e telele ea lithethefatsi (ho qobelloa). Ho makatsa hona ho lula ho sa hlalosoe ke likhopolo tsa hona joale tsa bokhoba. Khopolo ea ts'usumetso ea matla a ho susumetsa e nkoa e le mokhoa o ts'episang morerong ona, leha e sa fane ka mokhoa oa ho hlalosa ho fokotseha ha phaello ea meputso ea tlhaho ha nako ea ts'ebeliso ea lithethefatsi e ntse e eketseha. Haufinyane, Anselme o hlalositse mofuta o bitsoang mohlala oa sebopeho se lebelletsoeng (ADM) o bonts'ang karolo ea bohlokoa ea ho lebella le ho tsotella litšebelisanong tsa susumetso [8]. Ntle le ho itšetleha ka data e matla ea neuropsychopharmacological, ADM e fana ka mohopolo oa mantlha oa tšusumetso e ikhethang. Taba ena e ka lemohuoa joalo ka keketseho ea mohopolo oa tšusumetso ea maikutlo le ts'usumetso e khothalletsang hore na lithethefatsi li sebelisana joang le moputso oa tlhaho.

Khopolo-taba e 'ngoe ke hore ho qobelloa ho etsoa ka lebaka la neuroadaptations tsamaisong ea mesoporticolimbic dopamine le glutamatergic corticolimbic circry eo ho eona likhakanyo tsa dopamine li kenelletseng (Setšoantšo. 1) [9]. Sena se bululetsoe ho tsoa lipatlisisong tse mabapi le karolo ea liketsahalo tsa cellular tse tlasa Mehato ea Synaptic Plasticity ea ho ithuta le litlamorao tsa lithethefatsi [10]. Ka Synaptic plasticity re bolela liphetoho maemong a sehokelo, ka tsela e tloaelehileng e lekantsoeng ho sebelisoa mekhoa ea electrophysiological (mohlala: liphetoho ho karolelano ea AMPA / NMDA). Litlalehong tsa bokhoba ba lithethefatsi li-neural li pepesetsoa liphetoho tse tlisoang ke lingaka tse lemalloang, li lebisang moetlong oa litakatso tsa bokhoba [11]. Bopaki ba liphetoho tsena bo ka bonoa ka mefuta e mengata ea polasetiki libakeng tsa boko tse tsejoang ho ama tšusumetso, le ts'ebetso ea moputso [12-14]. Liphetoho tsena li fapana ho tloha maemong a fetohileng a neurotransmitter ho a fetohileng morphology ea sele le liphetoho mesebetsing ea bongoli [15]. Morphologically, boholo ba li-neuroadaptations tsena li fumanoe tsamaisong ea mesocorticolimbic le amygdala e atolositsoeng [13,15,16]. Kaha libaka tsena li bapala karolo tse hlahelletseng taolong ea mohono le ts'ebetsong ea meputso ea tlhaho, polasetiki e hokahane ka matla le boits'oaro bo tlatsetsang [7].

 

Setšoantšo sa 1

Dopaminergic sistimi le ts'ebetso ea moputso. Li-neuron tsa dopaminergic li fumaneha ka har'a midbrainakhiwo substantia nigra (SNc) le sebaka sa ventral tegmental (VTA). Li-axon tsa bona li kenella ho striatum (caudate nucleus, putamen le ventral striatum ho kenyelletsa ...

Tlatsetso le boteng

Boemong ba bokhoba ba lithethefatsi, ho sebelisitsoe likhopolo tse 'maloa ho hlalosa kamano lipakeng tsa polasetiki le bokhoba ba tahi. Ho ea ka polelo ea tšusumetso e boletsoeng pejana, ho pepesetsoa lithethefatsi khafetsa ho hlokomela litheko tsa tšusumetso ea lithethefatsi le litheko tse amanang le lithethefatsi. Liphetoho tsena li etsa hore sensitised nucleus accumbens (NAc) e lokolle dopamine (DA) kamora ho pepesetsoa lithethefatsi kapa litso tse amanang le eona (Setšoantšo. 1). Sena se ka bonahatsa ka boits'oaro e le takatso e matla ea lithethefatsi. Sena se ka etsoa ka mokhoa o lekiloeng ka ho lekanya boits'oaro ba ho batla lithethefatsi mabapi le mekhoa e hokahantsoeng le taolo ea lithethefatsi ho liphoofolo.17]. Hoa utloahala hore tlhokomeliso e fumaneha hohle bakeng sa moputso bobeli ba litheko le ba sa sebeliseng lithethefatsi [18].

Khopolo e 'ngoe e ka hokahanyang polasetiki le bokhoba ke mohopolo oa mohanyetsi [19]. Ka bokhutšoanyane, e fana ka maikutlo a hore ho na le lits'ebetso tse peli tse etsahalang nakong ea liphihlelo tse phetoang: 1. vagituation kapa hedonic habituation, le 2. Ho tlosoa hoa motho e mong kapa ea mo tšehelitseng [20]. Khopolo ena e ikhethile ka mokhoa o ikhethang oa tlhekefetso ea opiate moo litlamorao tsa phello e potlakileng li emelang ts'ebetso e amanang le bolulo, ha ho sa tlosoe liponahatso tsa ho tlohela chelete ho khannelang motho ea lemaletseng ho ts'oara lithethefatsi [21].

Ho bonahala e le keketseho ea mohopolo oa mohanyetsi mohopolo ke mokhoa o sisimosang oa lits'ebetso tsa ts'ebetso ea boko [19]. Ka Allostasis re na le litsamaiso tse peli tse hanyetsanang, adaptara ea hare-system le adaptation ea system e lipakeng. Ts'ebetsong ea ts'ebetso e kahare ho ts'ebetso, lithethefatsi li etsa hore ho be le karohano e hanyetsanang le ea ts'ebetso eo ho eona lithethefatsi li fumanang tšebetso ea lona ea mantlha le e sa phetheng, ha li ntse li le lipakeng tsa ts'ebetso ea mahareng, lits'ebetso tse fapaneng tsa neurobiological tse motho a li qalileng pele ke moriana. lia ngolisoa. Haufinyane, George et al., O bontšitse thahasello ea ho fetoha ho dopaminergic le corticotropin ho lokolla lintho tse tsamaisang lintho joalo ka hare ho sistimi le lipakeng tsa system-neuroadaptations ka tatellano, e tlatselletsang ts'ebetsong ea mohanyetsi ho lithethefatsi tsa tlhekefetso [22]. Ba kholoa hore ts'ebetso e khaotsoeng khafetsa ts'ebetsong ea dopaminergic le ts'ebetso e tsitsitseng ea sistimi ea CRF-CRF1R e nang le liketsahalo tsa ho tlohela e ka lebisa ho moroalo o matla haholo o tlatsetsang haholo phetohong ea bokhoba ba lithethefatsi. Ho itokolla ka matla lithethefatsong tsa tlhekefetso ho hlahisa mohanyetsi joalo ka liphetoho lipakeng tsa li-neurotransmitters tsa meputso maemong a ikhethang a meputso e amanang le tsamaiso ea mesolimbic dopaminergic le ho hira lits'ebetso tsa khatello ea maikutlo tsa amygdala le CRF tse hanyetsanang ka matla le litlamorao tse matla tsa lithethefatsi tsa tlhekefetso. Liphetoho tse joalo ho dopamine le CRF lits'ebetso tsena tsa boko tse amanang le nts'etsopele ea likarolo tsa susumetso tsa ho tlohela li khothalletsoa hore e be mohloli o moholo oa liphetoho tsa neuroadaptive tse tsamaisang le ho boloka ho lemalla. Ts'ebetso ea dopaminergic e fokotsehileng bokong ba nyutlelie le karolo e atolositsoeng e ka nka karolo ho rarolleng tšebetso, ke hore, kapa ho matlafatsoa hoa ts'ebetso ea meputso ea tlhaho le lithethefatsi tsa tlhekefetso, athe ho hira sisteme ea CRF-CRF1 mohlomong le dynorphin / κ opioid Sisteme ho CeA, BNST, le VTA nakong ea ho tlohela e ka nka karolo ho hlaha ha ts'ebetso ea b, ke hore, kapa boemo bo bobe ba maikutlo bo susumetsang takatso ea ho batla lithethefatsi. Leha bopaki bo bong bo fana ka maikutlo a hore dopaminergic le CRF lits'ebetso li ka sebelisana haufi-ufi, lipatlisiso sebakeng sena li haella. Ha ho tsejoe hore na ts'ebetso ea pele ea ts'ebetso ea dopaminergic ho VTA (a-process) e hlokahala bakeng sa keketseho ea ho lokolloa ha CRF lithutong tse atolositsoeng tsa amygdala le VTA (b-process) lithutong tse itšetlehileng ka lithethefatsi le tse tlosoang tse lebisang ho lithethefatsi tse qobelloang. ho batla le ho eketsa takatso ea lithethefatsi. Ha ho le joalo, liketsahalo tsa khafetsa tsa ho tlohela le ts'ebetso e tsitsitseng ea CRF-CRF1R e ka lebisa ho moroalo o matla haholo o tlatsetsang haholo phetohong ea bokhoba ba lithethefatsi.

Khopolo ea boraro ea ho hlalosa karolo ea ts'ebetso ea neuroplasticity ho lemalla ke ho hira methapo ea kutlo e thehiloeng lemating nakong eohle ea ho pepesetsoa lithethefatsi [14]. Ts'ebetsong ea koae ea cocaine ho liphoofolo, ho na le liphetoho phetohong ea tsoekere ea glucose le li-receptor tsa dopamine D2 le transporter ea pele e amang ventral striatum, liphetoho tsena hamorao li atoloha ho ama dorsal striatum [23]. Tsoelo-pele ena ea polasetiki ho tloha ho ventral ho ea ho dorsal striatum e ka ikarabella bakeng sa phetoho ho tloha ho sepheo-ho ea ho ho ithuta ho thehiloeng mesebetsing ea khale [24].

Khopolo-taba e 'ngoe e reng libaka tsa neural tse tšehetsang ts'ebetso ea motlakase ea boko (ESSB), li theha tsamaiso ea kelello ea maikutlo, sena ke seo re ka se bitsang Sistimi ea HO BONAHALA / HO BONAHALA. Sistimi ena e fetola maikutlo a motho ka mong tikolohong, 'me e etsa hore tebello e lebelle tsela bakeng sa meputso ea kamoso [25]. Ho bonahala ho khahlisa mabapi le khopolo ena ke hore ts'ebetso ea HO BONAHALA e fumanoa ke likokoana-hloko e le moputso ka mokhoa o le mong, e lebisang ho ESSB ntle le tlhoko ea mofuta o fe kapa o fe oa moetlo oa ho khatholla le meputso e hlakileng ea maikutlo [25,26]. Ho ipapisitsoe le tsamaiso ea lithethefatsi, li-microinjections le lithuto tsa lesion, mokhoa oa ML-DA o bonahala e le karolo ea bohlokoa ea litlamorao tsa phello e khothatsang ea MFB [27]. Le maemong a opioids (a nang le litlamorao tse khahlisang), liphoofolo li tloaetse ho iphelisa ka likhomphutha tse eketsang maemo a DA libakeng tsa ML [25,26,28]. Le ha lithuto tsa intracranial tsa tsamaiso e senola karolo ea li-neurochemicals tse ling tse fapaneng le DA ho moputso oa boko le mesebetsi ea ho atamela [28], Sisteme ea ML-DA e lula e le "neurochemical" e ka sehloohong e kenyang letsoho la ts'ebetso ea SEEKING. Haufinyane, Alcaro le Panksepp ba khothalelitse hore bokhoba ba motho ka kakaretso bo khetholloe ka polelo e sa tloaelehang ea SEEKING [29]. Haeba khatello ea maikutlo e tšoauoa ka phokotso e akaretsang ea HO RATA litlhoko, bokhoba bo ka hlalosoa e le mokhatlo o hlophisitsoeng oa maikutlo a fokolang a fokolang joalo ka mesebetsi e ikhethileng e atisang ho ba teng tikolohong. Ka pono ea bona e amanang le neuroethogical, bokhoba ke litholoana tsa "ho hloka kelello", ka lebaka la ho lula le lecha la ho batla ho nkeha maikutlo ka litakatso tsa meputso le litabatabelo tse tlisoang ke ho kokobetsa dysphoria e tlohang ho tloheng lithethefatsi [30].

Ho na le lintlha tse 'maloa tse tšehetsang qeto ea hore ts'ebetso ea ts'ebetso ea mesencephalic dopamine e kenelletse ho hohelang le ho arabela meputsong ea mefuta e mengata e fapaneng. Leha ho le joalo, karolo e tobileng ea dopamine ts'ebetsong ea moputso e ntse e le taba ea lipatlisiso [26,31,32]. Pele ho ne ho lumeloa hore dopamine e jara thabo kapa letšoao la hedonic, le bontšang boleng ba moputso oa lintho tse nang le boiphihlelo [32,33]. Tlhaloso ena e ipakile e le bonolo haholo. Ho amohela meputso ho ka etsa hore mosebetsi oa dopaminergic o eketsehe, empa ho na le maemo a mangata a sena. Ho hlahisitsoe maikutlo a 'maloa bakeng sa ho nkela sebaka sa hedonia hypothesis [27,33]. Tlhahlobo ena e shebisisa mohopolo oa hore ts'ebetso e fetoha ho dopamine neurons e kenyelletsa phoso ponelopele ea nako le palo ea meputso ea hang-hang le ea nako e tlang (polelo ea polelo e boletsoeng esale pele). Ho eketseha ha tšebetso ea dopaminergic ho etsoa ka sepheo sa ho bontša hore tebello ea moputso ea kapele kapa ea nako e tlang e ne e le betere ho feta kamoo e neng e lebelletsoe pele, ha ts'ebetso ea dopaminergic e tšoaea moqoqo [34]. Letshwao lena le kanna la sebelisoa ho ithuta ho bolela meputso le ho tataisa liqeto tse reretsoeng ho fumana meputso [27,35].

Dopaminergic sistimi le ts'ebetso ea moputso

Bokong ba batho ba baholo, li-neuron tsa dopaminergic (DA) ke sehlopha sa lisele tse nang le tšusumetso e sebetsang ebile li sebetsa hantle sebakeng sa mesencephalon, diencephalon le bulb ea olgicory [32,36]. Leha ho le joalo, hoo e batlang e le lisele tsohle tsa DA li lula karolong ea moea ea mesencephalon (Setšoantšo 1). Mesodiencephalic DA neurons e theha sehlopha se itseng sa neuronal se kenyelletsang karolo ea substantia nigra pars compacta (SNc), sebaka sa ventral tegmental (VTA) le tšimo ea retrorubral (RRF). Mohlomong, e tsejoang ka ho fetisisa ke sistimi ea nigrostriatal, e simolohang ho SNc mme e fetisetsa likhoele tsa eona ka har'a caudate-putamen nucleus mme e bapala karolo ea bohlokoa taolong ea motsamao oa boithaopo [37,38]. More medial tseleng ena ke sistimi ea mesolimbic le mesocortical DA, e hlahang ho li-neurons tsa DA tse teng VV mme e nka karolo boits'ebetsong bo amanang le maikutlo bo kenyeletsang khothatso le moputso [33,39,40]. Tsamaiso ea mesolimbic DA e kenyelletsa lisele tsa DA tsa VTA tse sebetsang haholo ho li-nucleus accumbens, ho li-tubercle tsa boiketso empa li boetse li boloka septum, amygdala le hippocampus. Ts'ebetsong ea mesocortical DA, VTA e atolosa likhoele tsa eona ka pele, cingrate le perirhinal cortex. Ka lebaka la ho hloka kutloano lipakeng tsa lits'ebetso tsena tse peli khafetsa li bitsoa tsamaiso ea mesocorticolimbic (Setšoantšo. 1) [41,42].

Bathong, ho na le li-neuron tse 'maloa ho SN le VTA, tse baloang ka tlase ho 400,000 ho SN le 5,000 e batlang e le VTA [36,43]. Le ha palo ea methapo ea kutlo e le nyane, likhakanyo tsa li-neuron ka bomong li pharalletse haholo ka hona li na le litlamorao tse matla tšebetsong ea boko. Ho nahanoa hore methapo ea methapo ea kutlo e tloaelehileng ea methapo ea kutlo ea DA e na le bolelele bo felletseng ba li-axonal (ho kenyeletsoa le li-collater) tse ka bang 74 cm [36]. Khokahano ea Synaptic le eona e pharalletse ka ho lekana, 'me li-terminals tsa 500,000 li tloaelehile ho neuron e le' ngoe [36]. Boemong ba striatum, moo li-terminals tsa DA li leng matla ka ho fetisisa, li ikarabella ka 20% ea li-synapses tsohle tse ka hara sebopeho [44,45].

Ho tloha ho li-nuclei tsa bona tse fapaneng, li-axon tsa DA li tsoella ka mokhoa oo li ikopanyetsang le ho kenella ka bongata ba methapo ea methapo ea methapo (MFB) ho ea kapuleteng e ka hare [36]. Ho tloha kapuleteng e kahare, lekala la axons le qala ho etsa li-synapses libakeng tsa bona tse [36]. Substantia nigra neurons e felisa haholo-holo ka har'a caudate le putamen nuclei (striatum), e theha tsamaiso ea nigrostriatal. Ma-axons a DA a hlahang ho VTA a khaola haholo karolong e kenellang ea striatum; sebaka se bitsoang nucleus accumbens (NAc), mme ke likarolo tsa mantlha tsa mesolimbic system [36].

Mekhoa e fapaneng ea 'mele ea DA e aroloa ke bonyane li-protein tse hlano tse fapaneng tsa protheine ea G e kopaneng.46,47]. Banyalani ba babeli ba D1-like receptor subtypes (D1A-1D le D5) ho G protein Gs le activate adenylyl cyclase [46,47]. Li-subtypes tse ling tsa receptor ke tsa D2-like subfamily (D2, D3, le D4) 'me ke prototypic ea G protein-coupled receptor e thibelang li-cyclase tsa adenylyl le li-activation tsa K + [46,47].

Li-receptor tsa DA li na le mokhoa o ts'oanang le oa ho ajoa ha li-neuron tsa projeke [32,48]. Khokahano e lekanang ea li-receptor tse tšoanang le D1 e bapisoang le D2 receptor e phahame kahara cortex ea pele, athe mahloriso a li-receptor tse kang D2 a phahamisoa ka har'a li-caudate nucleus, putamen, le li-bokellase tsa nucleus tsa batho [46,49]. Le ha li-receptor tsa D1 le D2 li na le litlamorao tse fapaneng mokhatlong oa limolek'hule, hangata li sebetsa ka mokhoa o kopaneng ha ho hlahisoa tse ling tse thata [50,51].

DA e sebetsa ka li-receptor tse kopantsoeng le G-protein ka mokhoa o tloaelehileng oa neuromodulatory [52]. Liwebsaete tsa tokollo tsa DA li beoa hang-hang kantle ho synftic cleft [53,54]. Ha e qeta ho lokolloa, DA e fapana ka har'a mokelikeli oa kantle ho oona o tsoa butle butle ka lebaka la ho khutla hape ka metabolism [55]. DA ha e ame ka kotloloho boits'oaro ba li-membran tse amohelang empa e fetola karabelo ea bona ho kenyelletso e amanang le eona [56,57]. Lintlha tsena tse tharo (tokollo ea extrasynaptic, phetiso ea lets'oao la receptor ea G-protein-coupled le mochini oa modulatory) li kenya letsoho ho karolo ea mantlha ea phetisetso ea DA, ke hore, tieho e telele e etsahalang lipakeng tsa ts'ebetso e tlamang (ho thunya ho phatlohileng) le liphetoho tse sebetsang ho likarolo tse amohelang. Ho hakanngoa hore, kamora ho ts'oara ha motlakase li-neuron tsa DA, phetoho ea ts'ebetso e tlalehiloe ho li-neuron tsa pelehi kamora ho lieha ho ka bang 300 ms [58]. Leha ho phatloha ho thunya ha li-neuron tsa DA ho etsahala ka lebaka la susumetso e loketseng [59], ha ho na monyetla oa hore matšoao ana a DA a phasic a susumetse, ho isa tekanyong efe kapa efe ea bohlokoa, karabelo ea boitšoaro (e kopantsoeng le ho fetisoa ka tsela e potlakileng) ho khothaletso e ts'oanang e ba bakileng [60,61]. Kahoo, pono e hlakileng ea karolo ea DA ho araba e kenyelletsa DA e le mokhoa oa ho lieha oa ho araba, o amang tšusumetso ea boits'oaro e latelang e ileng ea etsa hore e lokolloe [60,61].

Lithethefatsi tse ikemetseng li ama tsamaiso ea dopaminergic

Mokhoa o ikhethileng oa ho khetholla lits'ebetso tsa DA ts'ebetsong ea moputso o qalile ka lipatlisiso ka thepa e matlafatsang ea lithethefatsi tsa tlhekefetso. Boholo ba liphetho bo tšehetsa sephetho sa hore lithethefatsi tse lemalloang li arolelana thepa e tloaelehileng ea ho ntlafatsa ts'ebetso ea ts'ebetso ea ma-midbrain DA, haholoholo maemong a bona a "lietsahala"62,63]. Cocaine ke mofuta oa "blockerine" oa tlhahlobo ea litonki e kopanyang batho ba tsamaeang le dopamine. Bo-rakhoebo ba nkang marang-rang a tšehetsang DA, ke mochine o ka sehloohong oa ho tlosa dopamine ho li-synapses. Ho thibella ha batsamaisi, ka hona, ho ntlafatsa katleho ea ts'ebetso ea DA. Ke tšusumetso ena eo ho lumeloang hore ke sesosa sa bokhoba ba koae [64]. Amphetamines e sebetsa ka mokhoa o tšoanang. Ntle le ho thiba bapalami ba nkang bahahlauli ba DA, li-amphetamines le tsona li nkuoa ke baeti, 'me ka lits'ebetso tse kahare ho ts'usumetso e khutlisetsang ts'ebetso ea transporter [65,66].

Sephetho ke tokollo e kholo ea DA ke baeti ba nkang thepa, ka hona, mosebetsi oa DA o ile oa eketseha. Lithethefatsi tse ling tsa tlhekefetso li na le litlamorao tse sa tobang mosebetsing oa DA [67,68]. Joala bo lumeloa hore bo ama ts'ebetso ea boko haholo ka ho ntlafatsa ts'ebetso ea li-receptor tsa GABA, li-receptor tsa "inhibitory" tsa mantlha bokong [69]. Ethanol e tsejoa ho fokotsa sekhahla sa ho thunya ha li-neuron ka har'a substantia nigra pars reticulata [70], tseo ho lumeloang hore li fokotsa ho thunngoa ha li-neuron tsa DA [70,71]. Ka ho thibela li-neuron tsena, joala bo baka keketseho e kholo ea ho thunya ha sele ea DA, le ho eketsa tokollo ea DA maemong a striatum le nucleus [72,73]. Li-Opiates li baka tokollo e ts'oanang ea DA ho striatum [74], ka bobeli ka disinhibition ho VTA le ka litlamorao tse tobileng litsing tsa DA [74,75]. Ho feta moo, ho thibela li-opioid receptor ho VTA kapa li-nucleus kukusanya ho fokotsa taolo ea heroin [76]. Boitlhokomelo ba nicotine bo boetse bo thijoe ke ho kenella ha li-dopamine receptor antagonists kapa ke lesion ea dopamine neurons ka har'a li-nucleus accumbens [77]. Ka hona mokhoa oa DA o kopantsoe hore o kenye letsoho ka mokhoa o nyarosang litabeng tsa bokhoba ba nikotine hape [78]. Tlhahiso ea hore tsamaiso ea DA e kanna ea ba karolo ea tsela ea ho qetela e tloaelehileng ea litlamorao tsa lithethefatsi tsa tlhekefetso e khahla haholo mme e lumellana hantle le lingoliloeng tse mabapi le ho itlhakisa ha boko [79]. Ntle le moo, ho pepesehela lithethefatsi tsa tlhekefetso ho baka ho fetoha ha nako e telele mohlaleng oa cAMP, tlhahiso ea tyrosine hydroxylase, polelo ea DA, li-coupept tsa receptor le liprotheine tsa G, le sekhahla sa ho chesoa ha methapo ea methapo ea VTA-DA [80,81]. Mechine ena ho nahanoa hore e hohela bokhoba ebile e kenya letsoho ho oeleng hape ts'ebelisong ea lithethefatsi nakoana tse latelang.17,82,83].

Bokhoba ba lithethefatsi ha bo bobebe joalo ka sehokelo sa tsamaiso ea DA se ka re. Lits'ila tse tsoang ntle le batsamaisi ba DA, e leng karoloana ea litlamorao tsa koae tsamaisong ea DA, li ntse li khona ho nts'etsa lithethefatsi tsa koae [84,85]. Se sibolotsoeng se bontšitse hore litlamorao tsa koae ho ba tsamaisi ba serotonergic le noradrenanergic le tsona li ka phetha karolo ea bohlokoa tlhekefetsong ea lithethefatsi [86]. Mohopolo ona o tšehetsoa hape ke taba ea hore ts'ebetso e ntlafalitsoeng ea serotonergic e fokotsa taolo ea tahi [87,88]. Leha ho le joalo, leha mekhoa ea ts'ebeliso e mpe ea lithethefatsi le tšebeliso ea lithethefatsi e sa hlake, ho fumanoe hore dopamine e bapala karolo ea bohlokoa liketsahalong tsena ka bobeli, ka ho etsa joalo, e tiisa khokahano lipakeng tsa lits'ebetso tsa dopamine tsa boko le ts'ebetso ea moputso (Setšoantšo. 2).

 

Setšoantšo sa 2

Neurotransmitter molaoana oa boits'oaro ba ho batla moputso. Tsela e tloaelehileng ea boitšoaro bo batlang moputso bokong ke mesolimbic dopamine tsela. Tsela ena e fetoloa ke lintho tse ngata tse hlahang ka bokong hore li hlahise lintho tse tloaelehileng ...

Ho bonahala eka mokhoa oa neural oa "moputso" oa setso o ka atolosoa ho kenyelletsa mekhoa e 'meli e arohaneng, empa e hokahanyang, tsamaiso ea limbic ho ts'usumetso ea ts'usumetso ea lithethefatsi, le prefrontal cortex (PFC) ho laola taolo ea thibelo mabapi le tšebeliso ea lithethefatsi. Boithuto ba pele-pele bo fana ka bopaki bo sa fetoheng ba kamano pakeng tsa taolo ea lithethefatsi ea nako e telele, neuroadaptations ea PFC (haholo-holo meketjana e meraro ea li-PFC-striatothalamic, DLPFC, OFC le ACC), le phehello ea boitšoaro bo batlang lithethefatsi. Liphuputso tse entsoeng ka Neuroimaging li senotse hore bofokoli ba boitšoaro ba cocaine bo kenyellelitsoeng bo amanang le tlhekefetso e hlophisitsoeng ho OFC le ACC, le hypoacaction ea libaka tsa pele tsa cortical, haholo ACC le PFC. Bokhoba ba Opiate bo fokotsa bokhoni ba ho etsa liqeto. Maemong ana, lithuto tsa neuroimaging li bonts'itse likarabo tse sa tloaelehang tsa neural ho PFC; ba senotse tšebetso e kenelletseng ho ACC, ka likarabo tse fetotsoeng ka hara DLPFC le OFC. Ho se sebetse hantle libakeng tsena tse ka pele ho fumanoeng ho amahanngoa le khaello ea ts'ebetso ea botsamaisi le bokhoni ba ho etsa liqeto ho batho ba itšetlehileng ka opiate. Ho etsa liqeto tse nang le phoso ho tla beha motho ea lemaletseng ho etsa liqeto tse sa lokelang maemong a fapaneng. Ho itšetleha ka joala ho ne ho amahanngoa le litekanyetso tse fokotsehileng tsa taolo ea boiqapelo, boits'oaro bo sa potlakang le bokhoni ba ho etsa liqeto tse kotsi. Boithuto bo bobe ba batho ba itšetlehileng ka joala bo pepesitse taba e fokotseng bophahamo ba boko ba DLPFC, bo tšehelitsoeng ke lithuto tse sebetsang tsa neuroimaging, tse fumanoeng hore liphetoho lipakeng tsa taolo ea tšusumetso li kopantsoe ke hypoactivity ea DLPFC. Kahoo, ho bonahala eka batho ba lemaletseng joala ba na le ts'oaetso ea ho ba le litšila tse sa sebetseng tsa ts'ebetso ea ts'ebetso ea ho thibela maemo a kotsi. Sena le sona se ka fokotsa monyetla oa ho lula o sa tsitsa mme se ka thusa ho hlalosa litheko tse phahameng tsa ho oela hape har'a batho ba ts'ebelisanang le joala [89].

Lingoliloeng tse ngata li bonts'a hore polokeho ea khetla ea NAc e na le karolo ea bohlokoa ho e lokisa sepheo sa mantlha sa susumetso sa meputso e khahlisang le e khelohang [90]. Psychostimulants ka ho khetheha e susumetsa ho lokolloa ha dopamine ka khetla [91], 'me liphoofolo li tla tsamaisa lingaka tsa dopamine ka kotloloho sebakeng sena [92]. Pharmacologic inhibition ea khetla e eketsa boitšoaro bo susumetsang le likarabo tsa hedonic ho latsoa tšusumetso [93]. E lumellana le liphumano tsena, Wheeler le basebetsi-mmoho, 2011 e hlokometse - ka ho hlahloba ka potlako li-cyclic cytlic ho lekola ho lokolloa ha dopamine ea nako ea nnete litšoantšong tse nang le tatso e monate ea tatso e boletseng ho lieha ho fumaneha ha koae le nakong ea boipuso- hore tokollo ea dopamine sebakeng sena, empa eseng subregion ea mantlha, e phahamisoa ka potlako ka linako tse ratehang, mme e fokotseha ka ho sa rateheng, tatso ea tatso [94]. Ntle le moo, ba bonts'itse hore liphetoho tse potlakileng tse lokollotsoeng li ka fetoloa ke ho latela mekhatlo e ithutileng, haholo kamano ea boitlamo le ea nakoana ea tatso ea tatso le ho fumaneha ha koae. Leha ho le joalo, tokollo ea dopamine e potlakileng e ile ea bonoa nakong ea ho itlhokofatsa ka koae le bakeng sa mekhoa ea ho tsamaisa koae ka potlako (ekaba tastants kapa audiovisual).

Sisteme ea Dynorphin le dopamine

Li-peptide tse kang dynorphin li bonahala li kopantsoe tsamaisong ea moputso oa boko. Liphuputso tse fetileng li bontša hore ho hlohlelletsa li-receptor tsa kappa-opioid ho lebisa boemong bo bobe ba maikutlo ka ho thibela ho lokolloa ha dopamine ho striatum. Bahanyetsi ba Kappa-Opioid receptor ba na le litlamorao tse matla tse kang li-antidepressant [95], ho feta moo, ho fanoe ka maikutlo a hore tšebeliso e sa foleng ea lithethefatsi e etsa hore li-neuroadaptations li tsamaee ka har'a ts'ebetso ea dynorphin ea kelello e thibelang lithethefatsi tse khothalletsang tokollo ea dopamine. Leha, tlhahiso e ntseng e eketseha ea li-peptide tse kang dynorphin e kanna ea loants'a litlamorao tsa lithethefatsi tsa tlhekefetso, liphetoho tse tšoanang li ka ba le litlamorao tse mpe ha ts'ebetso ea lithethefatsi e khaotsa ho tlohela pina bakeng sa li-neuroadaptations tse sa hlahisoang ke li-dynorphin. Ho loketse ho hlokomela hore kappa-opioid receptor agonists e ka fumana matšoao a tlisoang ke lithethefatsi ka ho fokotseha ha glutamatergic, GABAergic, kapa phetisetso ea Oradrenergic bokong [96]. Joalokaha ho ka bonoa tšebeliso ea lithethefatsi e ka baka liphetoho tsamaisong ea dynorphin haholo ho caudate putamen, globus pallidus, le ventral pallidum [97]. Mesebetsi ea morao-rao e bontšitse hore libaka tsena li bapala karolo ea bohlokoahali ho laoleng mebuso ea maikutlo ntle le karolo ea tsona e tsebahalang ea ho laola mesebetsi ea makoloi. Lintlha tsena li ka hlahisa li-dynorphin e le li-palyers tsa bohlokoa tsamaisong ea moputso le ho kenella, ho etsa lipatlisiso mabapi le karolo ea bona ho ka thusa ho hlakisa mekhoa ea phekolo e eketsehileng bakeng sa tlhekefetso ea lithethefatsi.

Phapang ea motho ka mong

Lilemong tsa ho qetela, ho se tšoane ka liphatsa tsa lefutso ho palo ea batho e bile sehlooho se bohlokoa lipatlisisong tsa bongaka [98]. Ho 'nile ha nahanoa hore mefuta e tloaelehileng ea liphatsa tsa lefutso e ka kenya letsoho kotsing ea liphatsa tsa lefutso le hore e ka susumetsa karabo ea thuto ea tlhekefetso ea lithethefatsi. Haufinyane, ho bontšitsoe hore phapang e pakeng tsa batho ka bomong e bonahala lebaleng la moputso oa lithethefatsi [99]. Ho 1999, Volkow et al a., e hokahanya botebo ba euphoria le palo ea tokollo ea dopamine kamora ho hlohlelletsa D2 [100]. Liphumano tsena li bonts'itse phapang lipakeng tsa lithuto tse lekiloeng. Tlalehong e 'ngoe, khokahano pakeng tsa ho lokolloa ha dopamine ho araba amphetamine le boits'oaro ba lithethefatsi bo lahliloe [101]. Boithuto ba fMRI bo ikopantseng le ho ipapisa le joala ka ts'ebetso ea bootsoa; hona ho ka bontša hore ts'ebetso ea ts'ebetso ea ts'ebetso ea ts'ebetso ea ts'ebetso ea ts'ebetso e ka ama maikutlo a tlase le moputso oa lithethefatsi. Patlisiso tse ngata tse bonts'ang ho fokotseha ha phihlello ea D2 ea receptor, e hloka tokafatso e eketsehileng hore na sena ke phello ea ts'ebeliso e mpe ea lithethefatsi, kapa sebopeho se ikhethileng se ikemiselitseng ho lemalla.102].

Mokhoa oa Hypocretin / Orexin le sistimi ea moputso

Hypocretin / orexin (Hcrt) neuron e fumaneha feela ho hypothalamus, haholo-holo likarolo tsa eona tsa pherekano, dorsomedial le hamorao [103,104]. Li-fibre tsa Hcrt li pharalla hohle bokong 'me ka kakaretso li na le phello e thabisang liseleng tsa bona tsa postynaptic [105-107]. Li-neurons tsa Hcrt li laola lintho tse khahlisang 'me li bontšitsoe hore li na le tšusumetso ho moputso oa lijo le boits'oaro ba ho batla lithethefatsi [105]. Anatomically, orexin neurons e boemong bo botle ba ho fetola moputso o sebetsang [103,104]. Morero oa li-neurons tsa Hcrt oa ho putsa libaka tse amanang le boko, ho kenyeletsa le nucleus accumbens (NAc) le VTA, mme Hcrt e kenya ka ho toba li-neurons tsa VTA-DA ka Hcrt-1 receptor [108]. Sena se bonts'a karolo e ka etsoang ke Hcrt mosebetsing oa moputso le tšusumetso, e lumellanang le lithuto tse fetileng tse hatisang Hcrt ho fepa. Ebile, ts'ebetso ea li-neurons tsa Hcrt e bontšitsoe hore e hokahane ka matla le litakatso tsa mekhoa e amanang le meputso ea lithethefatsi le lijo [109]. Dopaminergic neurons tse simolohang ho VTA mme li kenella bokamosong, haholo-holo NAc, li khethollotsoe e le 'tsela ea moputso' [32]. Lithethefatsi tsa tlhekefetso li hlohlelletsa tsela ena. ICV kapa infusions ea VTA ea sebakeng seo ea Hcrt e bontšitsoe ho khutlisa mokhoa o batlang lithethefatsi kapa boitšoaro bo batlang lijo ka litoeba [109,110]. Ka lehlakoreng le leng, "subcutaneous morphine" (μ-opioid receptor agonist) - e reretsoeng ho rata libaka le hyperlocomotion tse bonoang litoeba tsa mofuta oa hlaha li felisitsoe ka litoeba tse neng li sena mofuta oa li-prero-Hcrt [111], le liente tsa antagonist ea Hcrt-1 receptor ho VTA e thibela nts'etsopele ea khetho ea sebaka se nang le boemo ba morphine [111]. Ka ente ea vivo e nang le PKC inhibitor chelerythrine chloride e khethileng kapa 2-3-1-methyl-1H-indol-3-ylmaleimide HCl (Ro-32-0432) sebakeng sa ventral tegmental (VTA) e hatelletse haholo khetho ea sebaka le maemo a eketsehileng. ea dopamine ka nucleus accumbens (NAcc) e bakoang ke ente ea intra-VTA ea Hcrt [112]. Liphetho tsena li ts'ehetsa ka matla mohopolo oa hore ts'ebetso ea ts'ebetso ea neuron e nang le orexin ho VTA e lebisa ts'ebetsong e tobileng ea li-neuron tsa mesolimbic dopamine ka ts'ebetso ea ts'ebetso ea PLC / PKC tsela ka alpha kapa Gbetagamma-subunit activation, e ka ba e amanang le nts'etsopele ea phello ea eona e khotsofatsang.

Mosebetsi oa morao-rao o fane ka leseli tse khahlisang ka har'a methapo ea methapo ea cellular le ea limolek'hule tse hlahisang litlamorao tsena ka ho bonts'a hore Hcrt-1 ea kenya letsoho ho VTA potentiates NMDAR (N-methyl-d-aspartate receptor) -mediated neurotransmission ka protheine kinase C e thehiloeng ho li-NMDAR ho VTA dopamine neuron synapses litokisetsong tsa slice [113,114] Ho feta moo, ts'ebetsong ea vivo ea "Hcrt-1 receptor antagonist blockag sensitization sensomization" ea cocaine le occlude e khothalletsoang ke cocaine e susumetsoang ke maqhubu a thabo a VTA dopamine neurons [113,114]. Liphetho tsena li bonts'a karolo ea bohlokoa bakeng sa ho saena hoa Hcrt ho VTA ka polasetiki ea neural e amanang le moputso, mme e bonts'a hore Hcrt e boetse e kenya letsoho ho hlokofatseng kelello ea kelello ea k'hok'heine le ho batla moputso. Liphumano tsena li bonts'a karolo ea bohlokoa ea orexin mekhoeng ea moputso le bokhoba ba lithethefatsi. Nako le nako, litoeba tsa prero-Hcrt-knockout ha li na tšenyo ea letho ho feta liphoofolo tsa mofuta o hlaha ho latela morapo oa morphine, joalo ka ha li arabeloa ka karabelo ea ho ikhula ea mmele.115]. Ho khahlisang ke hore bakuli ba bang ba narcolepsy ba robalang motšehare ba neng ba alafshoa ka li-amphetamine tse kang li-amphetamine le / kapa sodium oxybate (xy-hydroxybutyrate, eo hape a tsejoang e le GHB) ka nako e telele e neng e sa qale ka tlhekefetso ea lithethefatsi [116]. Lipatlisiso tsena li bonts'a tšebelisano e matla ea tšebelisano lipakeng tsa tsela ea Hcrt le sistimi ea DA [117].

Lithutong tsa litekanyetso, ho latela ho felisoa ha koetliso ea koae ho ile ha khutlisoa ka ho pepesetsoa lits'ebetso tse amanang le lithethefatsi. Leha ho le joalo, puseletso ena e bakiloeng ke cueine e bakiloeng ke ho batla koae kapa e kentsoeng molemong oa ho batla k'hok'heine [118] e ne e koetsoe ke tsamaiso ea tsamaiso ea 20 kapa 30 mg / kg SB (ORX-1 blocker) [119]. Liphetho tse tšoanang, leha ho le joalo, ha lia ka tsa fumanoa ho sebelisa OxR2 antagonist 4pyridyl methyl (S) -tert-leucyl 6,7-dimethoxy-1,2,3,4-tetrahydrosisquinoline (4PT), e bonts'a karolo e ikhethang ea orexin signaling ka kotloloho ho OxR1 ho coca119]. Ho feta moo, SB e bontšitsoe hore e fokotsa haholo ho itaola ha ethanol, nicotine, lijo tse nang le mafura a mangata, le sucrose [120], hammoho le ethanol ho ja litheolelo tse tsoang kantle ho joala [121]. Joalokaha ho ka bonoa sistimi ea Orexin e bapala karolo ea bohlokoa ho etseng moputso.

Orexin le letsoalo

Ts'ebetso ea orexin system e bonahala e amana le sebaka sa bona sa marang-rang. Kahoo, mesebetsi ea ho batla moputso e amana haholo le lisele tsa orexin ho LH, athe lits'ebetso tse khahlisang- le khatello ea maikutlo li hokahane le li-neuron tsa orexin ho DMH le PeF [122]. Lithuto tse 'maloa li tšehetsa maikutlo ana. Mohlala, li-neuron tsa PeF le DMH orexin li bonts'a ts'ebetso ea Fos nakong ea ho tsoha ha e bapisoa le boroko [123]. Ka lehlakoreng le leng, li-neuroleptics li etsa hore activate ea LH orexin neurons [124]; Tšebeliso e sa feleng ea ethanol e ile ea eketsa sebaka sa polelo ea orexin mRNA ka LH empa eseng DMH / PeF. Ts'ebetso tsena tse arohanang tsa li-neurone tsa orexin li bonts'a marang-rang a fapaneng a amanang le marang-rang kapa moputso. Kahoo, projeke ea LH orexin cell projeke ho VTA kapa medial prefrontal cortex (mPFC) [124]. Ha a ntse a, li-neuron tsa PeF / DMH orexin li kenelletse ka hare ho libaka tse ling tsa hypothalamic [117].

Corticotropin-releasing factor (CRF) le orexin / hypocretin

Haufinyane ho khothalelitsoe hore N / OFQ (nociceptin / orphanin FQ) le li-neuropeptide Orx / Hct neuropeptide li sebelisana le tsamaiso ea CRF. N / OFQ e thibela tšebetso ea li-neuron tsa Orx / Hcrt [125]. Phello ena e tla lebisa ho khopolo-taba ea hore N / OFQ e boetse e hlophisa mesebetsi ea Orx / Hcrt, ho kenyelletsa le karabelo ea boitšoaro ho khatello ea maikutlo, ho tšoenyeha, moputso le ho lemalla. Patlisiso ea litšebelisano tsena e tla ba sepheo sa bohlokoa sa lipatlisiso tsa nako e tlang mabapi le litsamaiso tsa taolo ea khatello ea methapo [126].

Sistimi ea histori le ea moputso

Le ha dopaminergic system e emela lejoe la sekhutlo ho le meputso, lits'ebetso tse ling tsa li-neurotransmitter tse kang li-opioids tsa endo native, glutamate, GABA, acetylcholine, serotonin, adenosine, endocannabinoids, orexins, galanin le histamine li fumanoe ho fetola litlamorao le litlamorao tsa lithethefatsi tse lemalloang [127]. Boithuto bo 'maloa bo senotse hore tsamaiso ea histaminergic e fetolela phetisetso ea mesolimbic dopamine. Ho feta moo, e bonahala e ntlafatsa thepa e tlisang lithethefatsi. Ho tšehetsa khopolo ena ke ho fumana hore li-dopamine anatagonists li ile tsa sitoa ho paka katleho ea tleliniki ho phekola tlhekefetso ea lithethefatsi. Sena se netefalitsoe ke ho fumana hore H inverse agonist BF2.649 (Tiprolisant) e ntlafalitse tšebetso ea histamine ea histamine mme ea fokotsa tšebetso ea methamphetamine-induction locomotor [128].

Sisteme ea bongo histamineergic

Nuberomamillary nucleus (TM) e na le li-neuron tse fokolang, e leng mohloli o ka sehloohong oa histamine bokong. Leha ho le joalo, li-neuramin tsa histaminergic li na le marang-rang a mangata a lipalo a ka fihlellang libakeng tse ngata tsa boko. Empa ho na le libaka tse fapaneng pakeng tsa libaka mabapi le botebo ba likhakanyo tsena ka letsoalo le phahameng haholo ho "hypothalamic nuclei". Li-receptors tsa H ke li-receptor tsa G protein-coupled (GPCRs): the. Likarolo tse tharo ho tse 'ne tsa li-X receptors 1-3 li phatlalalitsoe haholo ka har'a tsamaiso ea methapo ea methapo ea mali. Li-receptor tsa H li fumaneha haholo lits'ebetsong tsa postynaptically le Mediate liketso tsa thabo ho ts'ebetso ea boko bohle. H1 receptor e kopantsoe le G q / 11 e lebisang ts'ebetsong ea phospholipase C, le man messengersosa a mabeli a bobeli, DAG le IP (3). H2, ka lehlakoreng le leng, e kopantsoe le Gs le ho kenya ts'ebetso ea cymbase ea adenylyl, PKA le karolo ea cAMP-reaction element binding (CREB). Ho fapana le moo, li-receptor tsa H3 li kopantsoe le G i / o e nang le cyclase ea inhibit adenylyl. Sena se etsa hore e be li-receptors tsa inhibitory. Li ka thibela ho kopana le ho lokolloa ha li-neurotransmitter tse fapaneng ho kenyeletsa DA, noradrenalin, GABA le acetylcholine [129].

Ho hokela histaminergic le dopaminergic system

Matsoalo a phahameng a li-receptor tsa H2 le H3 li fumanoa li-striatum (ho kenyeletsoa NAc) ho litoeba, likhoto, litšoene le batho [130]. Ho feta moo, li-interneuron tsa striatal cholinergic li na H1 [131]. Leha ho bile le ngangisano e kholo, litlaleho tse 'maloa li fumane hore H1 e hanyetsang e ka baka litlamorao tse joalo ka liphoofolo le bathong ka ho ntlafatsa ho lokolloa ha dopamine. Leha ho le joalo, kamano pakeng tsa lits'ebetso tsena tse peli ha e bobebe joalo ka ha histamine e ka sebetsa lits'ebetsong tse fapaneng tsa neuronal ho sitisa kapa ho kenya tšebetsong ts'ebetso ea midbrain dopamine. Ka li-receptor tsa H1 mohlomong tse fumanehang ho li-interneuron tsa striatal cholinergic, histamine e ka kenya tšebetsong tsamaiso ea mesolimbic. Ka lehlakoreng le leng, histamine e ka fokotsa phetisetso ea dopamine ka li-receptor tsa H 3 tse fumanehang ka ho khetheha litsing tsa dopamine kapa li-postynaptically ho li-neurons tsa GABAergic ho striatum [132].

Sistimi ea Ghrelin e bohareng le moputso

Sisteme ea Ghrelin e na le khokahano ea bohlokoa taolong ea taolo ea lijo le botsitso ba matla [133]. Sisteme ea Ghrelin e kenyelletsa litsela tse anngoeng ke ts'usumetso ea ghrelin receptor, GHS-R1A (kholo ea lekunutu la siriagogue receptor 1A). GHS-R1A e atile haholo bokong; ho kenyelletsa hypothalamus, brainstem, tegmentum le hippocampus. "Sistimi e kholo ea" ghrelin signaling "ke polelo e hlalosang hantle litheknoloji ea sethethefatsi sena, hobane e bonts'a tšebetso ha ho sa na ghrelin ligand [134]. Khopolo ea pele ea GHS-R1A e ne e le li-1980s ha peptide e ne e bitsoa kholo ea li-hormone ea ho holisa peptide 6 (GHRP6), e ileng ea fumanoa e le e tsosang takatso ea kholo ea "hypothalamo-pituitary" axis [135]. Hamorao, ligand ea bona GHS-R1A, e ile ea hlalosoa ke Merck & Co. Group. Ho sibolloa, hore lisele tsa hypothalamic tse sebelisitsoeng ke GHRP-6 e bile ketsahalo e 'ngoe ea bohlokoa ts'ebetsong ea ts'ebetso ena. Mekhoa e nepahetseng ea ghrelin e amang meputso e ntse e hloka lipatlisiso tse ling. Leha ho le joalo, ho bonahala e amana le sistimi ea moputso oa cholinergic-dopaminergic. GHS-R1A e hlahiswa pele le kamora synaptic ho VTA [136] hammoho le li-neuron tsa cholinergic ho LDTg [137]. Dickson le al. [137] o khothalelitse hore mokhoa o bohareng oa lumello ea ghrelin o sebetse e le ntlafatso ea limpho tsa moputso ka ho fetola ntlha e behiloeng ea li-neuron tsa dopaminergic ho VTA. Ho khahlisang le ho feta ke ho fumana hore GHSR1A e bonts'a ts'ebetso ntle le ligand. Sena se ka botsa hore na ke ghrelin ka boeona e fanang ka leseli ho ntlafatsa mochini oa moputso. Ka sebele GHS-R1A e fumanoe e laoloa ka mokhoa o ikemetseng oa ghrelin ka heterodimerization ho dopamine D1-like receptor [138]. Ha e e-so tsejoe hore na dopamine D1 receptor e susumetsa moea o bohareng oa ghrelin ho supa joang le bohlokoa ba 'mele ba phello ea boemo bona bo ntse bo lokela ho tsejoa. Ho feta moo, sistimi ea ghrelin e hokahane le moputso oa joala [139,140], cocaine, amphetamine [141], le lijo tse fumanehang habonolo / tse khotsofatsang [142]. Ka kopanelo lithuto tsena li fana ka maikutlo a hore ho supa li-ghrelin tse bohareng, ho kenyelletsa GHS-R1A e ka theha sepheo sa ho hlahisa maano a kalafo bakeng sa boits'oaro bo bobebe [139].

Galanin le tsamaiso ea moputso

Gut peptide galanin e ile ea fumanoa ho 80s [143]. Ho sibolotsoeng hona ho ile ha lateloa ke ba bang ho bolelang hore galainis e boetse e ajoa bokong bohle. Li-ligand tsena li ipakile li hokahana le mesebetsi e mengata ea bohlokoa e kenyelletsang boitsoaro ba ho fepa, ho feto-fetoha ha maikutlo, ho tsitsa, ho ithuta le ho hopola [144]. Ho na le li-receptor tse tharo tsa galanin: GalR1, GalR2 le GalR3 [145]. Li na le liprotheine tse G ebile li khona ho kenya protheine ea Gi le Go [146]. Ntle le ho kenya liprotheine tsa Gi le Go joalo ka GalR1-3, GalR2 e boetse e kenya liprotheine tsa Gq [146hape e ka eketsa ho tšoaetsoa ha calcium le ketsahalo ea li-phello tse tlase tse kang PKC [147]. Sena se ka bolela mesebetsi e thata ea li-subtypes tse fapaneng tsa galanin.

Galanins le sisteme ea dopamine

Galanin e fokotsa tlhahiso ea dopamine e hlohlelletsang litheko tsa litekanyetso ka mochini o kenyang liprotheine tsa Gi [148]. Sena se lumellana le bokhoni ba galanin ba ho fokotsa glutamate, empa ha se GABA lokolloe ka likhechana striatal. Ho feta moo, tsamaiso ea methapo ea kutlo ea galanin e ka phahamisa ho bokellana ha DOPA ka striatum, NAc le li-tubercles tsa boiketso mme ea fokotsa ts'ebetso ea locomotor ho likhoto [149]. Kaha litlamorao tse hlahisang boitšoaro ke ts'ebetso ea maikutlo, lingoli li khothaletsa hore keketseho ea ho bokelloa ha DOPA e tsoa ho phallo e theohileng ea dopamine, ho imolla thibelo ea methapo ea kutlo ea "patoreceptor". Kameho ea galanin ho bokelleng ba DOPA e boetse e etsahala ha galanin e kenngoa ka har'a VTA, empa eseng NAc, e fana ka maikutlo a hore VTA ke sebaka sa mantlha sa ts'ebetso bakeng sa litlamorao tsa galain ho sistimi ea mesolimbic [149]. Tumellanong le khopolo ena, galanin e fokotsa tšebetso ea locomotor ho likhoto ha e kenngoe ka har'a ventricle, VTA kapa hypothalamus [150]. Ha li kopantsoe hammoho, liphetho tsena li fana ka maikutlo a hore litlamorao tsa galain ho VTA li ka fokotsa ts'ebetso ea sisteme ea mesolimbic.

Le ha galanin e sena ts'ebetso ho palo ea li-neuron tsa TH tse sa sebetseng ka boeona, kalafo e nang le dibutyryl cAMP e eketsa palo ea li-neurons tsa TH, mme phello ena e fokotsehile ke galanin. Litloaelo tsena li bontša GalR1, GalR2, le, ho isa tekanyong e tlase, GalR3 receptor mRNA, empa kalafo e nang le dibutyryl cAMP ka ho khetheha e eketsa maemo a GalR1mRNA. Ka hona, galanin e ka sitisa ts'ebetso ea midbrain dopamine ka ho fokotsa mosebetsi oa TH o arolelanoang ka ts'ebetso ea li-receptors tsa GalR1. Ha litoeba tsa GalR1 tsa ho kokota le litoeba tsa mofuta o hlaha li sa fapana ka taba ea mantlha [151].

Galanin e etsisa mekhoa e amanang le bokhoba ba tahi

Tumellanong le bokhoni ba galanin ba ho fetola tšebetso ea midbrain dopamine, liphuputso tse 'maloa li bontšitse hore tsamaiso ea galain e etsisa mekhoa e amanang le lithethefatsi. Mohlala, tsamaiso ea galanin ho li-ventricle tse kenang hamorao e bonts'a ntlafatso ea sebaka se ratoang ke morphine ho litoeba [152]. Tumellanong le ho fumana hona, litoeba tsa "kokonate" tse sieo peptide ea galanin, ho fapana le mofuta oa litoeba tse hlaha, li tsotella thepa ea morphine ebile e bonts'a mokhoa o ikhethileng oa morphine [153]. Likamano tse ling tse 'maloa pakeng tsa sistimi ea galanin le bokhoba ba opioid li tlalehiloe. Ramatiki e sa foleng, e rarahaneng ea morphine ho likhoto fatše e laola polelo ea galanin ka amygdala e atolositsoeng ka tsela e itšetlehileng ka mo-opioid receptor-154], athe GalR mRNA e ntse e eketseha ho LC nakong ea ho khaotsa opiate [155]. Ho feta moo, li-polymorphism tsa nucleotide e le 'ngoe ka gene ea galain ea motho li amahanngoa le temallo ea heroin [156]. Galanin e boetse e bontšitsoe ho fetolela karabelo ea boitšoaro ho li-psychostimulants. Lits'ila tse haelloang ke galanin peptide li ameha haholo ho litlamorao tse khahlisang tsa koae ka ha e lekantsoe ke thatohatsi ea sebaka [157]. E lumellana le phello ena, litoeba tsa transgenic tse hlalosang galanin ha li na taba le litlamorao tsa amphetamine ha li bapisoa le litoeba tsa mofuta oa naha.158]. Ha li kopantsoe hammoho, lintlha tsena li fana ka maikutlo a hore phello e akaretsang ea galanin bokong ke ho fokotsa likarabo tsa boitšoaro ho morphine le psychostimulants.

Ho fapana le morphine le psychostimulants, galain e ka eketsa tšebeliso ea joala tlasa maemo a 'maloa a liteko. Tsamaiso ea galanin ekaba ka har'a ventricle ea boraro kapa ho PVN ea hypothalamus e ntlafatsa ts'ebetso ea joala ea boithaopo liphoofolong tse tloaelehileng, e leng tšusumetso e bileng e fumanoang boteng ba lijo le likhoto tse khethiloeng bakeng sa joala bo bongata [159]. Litlamorao tse fapaneng tsa galanin ho morphine, amphetamine le cocaine locomotion le moputso ha li bapisoa le tahi ea joala li fana ka maikutlo a hore libaka tse fapaneng tsa boko li nka likaroloana tsena tse peli tsa likarabo. Ho lekanya ho hakanya hore litlamorao tsa galinin ho li-circuits tsa hypothalamic tse kenyelletsoang ho fepa li bohlokoa bakeng sa litlamorao ho ts'ebeliso ea lino tse tahang, athe lits'oants'o tsa methapo e fetolehang ho mesolimbic dopamine systemmay e bohlokoa bakeng sa litlamorao tsa eona litloaelong tse amanang le psychotimulant-and opiate. Bokhoni ba galanin ba ho fetola norepinephrine, serotonin, acetylcholine le ho lokolloa ha glutamate li ka feto-fetola ka tsela e sa tobang ts'ebetso ea li-dopamine neurons, tsa lebisa ho feto-fetohang ha boitšoaro bo amanang le lithethefatsi. Ha ho kopantsoe 'moho, bopaki bo bongata bo feto-fetohang bo fana ka bopaki ba hore mofuta oa galanin oa tlhaho o fana ka thibelo ea tonic ho li-system tsa neurotransmitter tse ka sebelisang ho itaola le lithethefatsi le ho khaotsa ho tsuba. Boithuto ba nako e tlang bo shebaneng le bokhoni ba galanin ba ho fetola tsela ea mesolimbic ho vivo le vitro ho tla hlokahala ho fumana kutloisiso e ntle ea hore na li-pharmacological tse lebisang tsamaiso ea galain li ka sebelisoa joang ho phekola ho lemalla lithethefatsi lithutong tsa batho [160].

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