Dev Cogn Neurosci. 2014 Nov 22. pii: S1878-9293 (14) 00082-6. Doi: 10.1016 / j.dcn.2014.11.001.
Sim1, Moghaddam B2.
inahaneloang
Ho nahanisisa ka ho hloka taolo ea moputso oa bongoana ho kenyelletsa ho etsa liqeto tse fosahetseng le ho eketsa monyetla oa ho holofala le ho ba le mathata a kelello. Hanyane hanyane hoa tsebahala; leha ho le joalo, mabapi le hore na boko ba bongoana bo sebetsana joang le moputso. Likhopolo tsa hona joale tsa ho lokisa moputso li nkiloe mefuteng ea batho ba baholo. Mona re lekola lipatlisiso tsa morao-rao tse shebileng kutloisiso ea hore na boko ba bongoana bo arabela joang moputsong le liketsahalong tse amanang le moputso. Karolo e bohloko ea mosebetsi ona ke hore liphapang tse amanang le lilemo li bonahala ka ts'ebetso ea neuronal ea liketsahalo tse amanang le moputso libakeng tsohle tse ngata tsa boko le ha likhoto tsa bocha li bontša boitšoaro bo tšoanang le ba batho ba baholo. Tsena li kenyelletsa phapang ea ho lokisa moputso pakeng tsa likhoto tsa bocha le tsa batho ba baholo ho cortex ea orbitofrontal le dorsal striatum. Ho makatsang ke hore phapang e nyane e amanang le lilemo e bonoa ho li-ventral striatum, e 'nileng ea e-ba ntlha ea lithuto tsa nts'etsopele. Re tsoela pele ho buisana ka litlamorao tsa phapang ena bakeng sa mekhoa ea boits'oaro e amehang bohlankaneng, joalo ka ho ts'oaroa, ho ipeha kotsing le ho fetoha ha boitšoaro. Ka kopanelo, mosebetsi ona o fana ka maikutlo a hore tšebetso e hlahisoang ke moputso e fapana e le mosebetsi oa lilemo le hore libaka tse joalo ka dorsal striatum tse sa amaneng le ts'ebetso e ntle ho batho ba baholo e ka ba tsa bohlokoa bakeng sa ts'ebetso ea moputso le ho ba kotsing ea kelello ho batjha.
LINKOTSO:
Bacha; Dopamine; Electrophysiology; Rat; Moputso; Striatum
Lintlha-khōlō Tsa
Boko ba bongoana bo etsa moputso o fapaneng le oa batho ba baholo.
Phapang ena e etsahala leha boitšoaro bo tšoana pakeng tsa lihlopha tsa lilemo.
DS e ne e le sebaka sa phapang e kholo ea nts'etsopele ea mosebetsi oa moputso.
Ho makatsang ke hore liphapang li ne li sa phatlalatsoe joalo ka VS
Liphapang tsena li ka ba le phello ea ho ba kotsing ea kelello ea bongoaneng.
1. Selelekela
Patlisiso ea hajoale mabapi le mafu a mafu a kelello e kentse khatellong e matla ea ho lemoha le kalafo ea pele. Matšoao a mangata a lefu la kelello: ho ferekana kelellong le bokhoba ba tahi li bonahala pele nakong ea bohlankana.Adriani le Laviola, 2004, Casey et al., 2008, Schramm-Sapyta et al., 2009 'me Mitchell le Potenza, 2014). Ka hona, ho bohlokoa ho hlakisa maemo a kotsi le a tlhaho le tikoloho a fang bacha ba kotsing e kholo ea mafu ana. Tsebo e joalo ea mochini e hlokahala bakeng sa nts'etsopele ea mehato ea ho thibela kapa ho fumana ho hlaha ha lefu.
Patlisiso ea pejana ea nts'etsopele ea bokong le lefu e hlile e lekotse liphetoho tsa morphological kapa liphetoho maemong a li-receptor. Lithuto tsena li fane ka leseli le tebileng mabapi le biology ea bocha le boitšoaro. Ha ho tsejoehale, leha ho le joalo ka matla a nnete a ketsahalo ea neuronal nakong ea boitšoaro. Tlhahisoleseling ena e bohlokoa haholo ka lebaka la likhopolo tsa morao-rao tse bontšang hore ts'ebetso ea ts'ebetso ea methapo ea methapo e sebetsa hantle molemong oa thuto ea lefu lena (Uhlhaas le Sebini, 2012 'me Moghaddam le Wood, 2014). Ho utloisisa ka botlalo hore ts'ebetso ea ts'ebetso ea methapo ea kutlo e fetoloang ka mokhoa o nepahetseng e fetoloa joang ho batho ba tlokotsing, re tlameha ho utloisisa pele hore na li-neurons le li-enuralles tsa encordles li kenella joang liketsahalong tsa bocha le ba baholo.
Liphetoho tse amang, tse susumetsang, le tšebetso e susumetsang nakong ea bocha ke tse ling tsa mekhoa ea pele e boletsoeng esale pele ea lefu la kelello le mafu a mang a kelello a leng kotsing e kholo ea batho ba (Ernst et al., 2006, Gladwin et al., 2011 'me Juckel et al., 2012). Ho utloisisa nts'etsopele ea matšoao nakong ena ea tlhabollo ea tlokotsi, ho bohlokoa ho hlakola mekhoa ea mantlha e tlatselletsang ts'ebetsong ea meputso ea bohlankana. Lintlha tsa morao-rao tse bokelletsoeng ka har'a lab ea rona re sebelisa likhoto tsa bocha li fana ka maikutlo a phapang e kholo e amanang le lilemo ketsahalong e hlahisang moputso. Liphapang tsena li bonahala esita le ha (1) boitšoaro bo lekantsoeng bo lekana lipakeng tsa lithuto tsa bocha le ba baholo, mme (2) maemo a motheo a ts'ebetso ea neuronal a lekana pakeng tsa lihlopha tsa lilemo. Kahoo, ts'ebetso ea "neuronal" e hlahisitsoeng ke moputso ka linako tse ling e ka sebetsa haholo ho feta mehato ea boitšoaro ea tšusumetso kapa ts'ebetso ea mantlha e le letšoao la ho ba kotsing ea lefu habonolo. Thutong ena, re akaretsa lintlha tsa ho ntlafatsa moputso oa bongoana tse fumanoeng mohlaleng oa likhakanyo libakeng tsohle tse ngata tsa boko, 'me re tšohla moelelo oa liphapang tsena bakeng sa boitšoaro ba bongoana le ho ba kotsing ea mafu.
2. Ts'ebetso ea moputso oa bongoaneng e fapana le ea batho ba baholo ho pholletsa le libaka tse ngata
Mokhoa o tsepamisitsoeng maikutlo tekong ena ke ho rekota ha motho a le mong moo likarolo tsa tšebetso ea methapo ea methapo e mengata li ka lekanngoang ka nako ea sebele le liphoofolo.Sturman le Moghaddam, 2011b). Bakeng sa mokhoa ona, lisebelisoa tsa li-electrode tsa multwire li kentsoe libakeng tse itseng tsa boko mme lipontšo tsa motlakase lia ntlafatsoa ebe li fetisoa ka mokhoa o phahameng ho khetholla tšebetso e phahameng ea maqhubu a neuronal, joalo ka ts'ebetso ea ts'ebetso kapa ts'ebetso ea tšimo ea lehae.Buzsaki, 2004, Sturman le Moghaddam, 2011b 'me Wood et al., 2012). Ho lekanya tšebetso ea maqhubu a ho etsa likhoto tsa bocha ketso e thata ke mosebetsi o boima, hobane fensetere ea lilemong tsa bocha e fela pakeng tsa matsatsi a ka mor'a ho beleha 28-55 (Spear, 2000). Kamora ho etsa accounting bakeng sa nako e hlokahalang bakeng sa ts'ebetso ea ho kenella ea electrode, ho hlaphoheloa le ho lula, nakoana e setseng ea fensetere e thibela ts'ebeliso ea likarolo tse rarahaneng tsa boits'oaro bo nang le electrophysiology. Ka hona, mesebetsi ea boits'oaro e sa hlokeng linako tse telele tsa ho ikoetlisa e tlameha ho sebelisoa ho lekanya ts'ebetso ea moputso mefuteng ea bocha. Leke ea rona e sebelisa mosebetsi oa bohlokoa o thusitsoeng moo likhoto li ithutang ho kenella koung ea litere ho amohela pellet e le 'ngoe ea tsoekere, ha mesebetsi ea neural e tlalehiloe ho tloha ho li-elektrode tse kentsoeng libakeng tse itseng tsa boko (Feie. 1). Habohlokoa, mosebetsi o bonolo ka ho lekana hore ho ithuta le ho etsa lintho tsa mantlha tsa mosebetsi ho ka bapisoa pakeng tsa batho ba baholo le bacha (Sturman et al., 2010), ka hona phapang efe kapa efe mesebetsing ea neuronal ke sesupo sa phapang ea ho lokisa moputso, ho fapana le sehlahisoa sa boits'oaro bo botle lipakeng tsa lihlopha. E 'ngoe le e' ngoe ea liketsahalo tsena ea boits'oaro e ka hokahanngoa le mehato ea ts'ebetso ea neural ka tharollo ea nakoana ea nakoana ea nakoana, ho lumella tlhahlobo ea liketsahalo tsa neural tse amanang le mekhoa e amanang le moputso, liketso tse lebisitsoeng ho sepheo, le tebello ea ho lebella le ho fana. Re sebelisa lintho tse fapa-fapaneng tsa mosebetsi ona, re ne re rekota ho tloha orbitofrontal cortex, dorsal le ventral striatum, le sebaka sa likarolo tse ka hare ho likhoto tsa batho ba baholo le ba lilemong tsa bocha. Joale re buisana ka hore na liphapang tsena mabapi le ho lokisa moputso li ka amana joang le litšobotsi tse lemohileng tse amanang le moputso tse bonoang nakong ea bocha, ho kenyelletsa ho se ts'oenyehe, ho ipeha kotsing le ho fetoha habonolo ha boitšoaro.
Setšoantšo sa 1.
(A) Electrophysiology ea yuniti e le 'ngoe e entsoe ka likotsi tsa bocheng tsa bocha le batho ba baholo nakong ea boits'oaro bo amanang le moputso. Likhoto li ile tsa kenngoa ka liphutheloana tsa microwire 'me tsa kenngoa kahara phaposi e sebetsang e nang le koung ea nko, ka sejo sa lijo se hlahisang meputso ea tsoekere, le leseli la cue le sebelisitsoeng ho bonts'a phallo ea moputso. Ho ke ho hlokomeloe hore boitsebiso ba seemahale e ne e le leseli, molumo kapa lefokisi le kopantsoeng la bobeli. (B) Mesebetsi ea bohlokoa e sebelisitsoeng e qalile ka ho bonesa ke khanya ea leseli, nakong eo ts'ebetso ea nosetso e nkang (ketso) e ileng ea baka phumants'o ea pellet. Kamora hore pokello e bokelle moputso, ho ile ha qaloa nako ea ho fetela tekong, ebe teko e latelang ea qala. (C) Morero ona oa mocheso o bonts'a dampuli tsa mohlala tse bonts'a karabo e tloaelehileng ea li-neuron ka ketsahalo e amanang le moputso. Mokhoa o ka tlase oa li-neuron o bonts'a sekhahla se eketsehileng sa ho thunya ho potileng ketsahalo (ka tlase), ba bang ba bonts'a sekhahla sa ho thunya nakong ea ketsahalo (holimo), mme ba bang ha ba arabele (bohareng).
2.1. Pele ho cortex
Pele-pele cortex (PFC) e tsoela pele kholisong e kholo lilemong tsa bocha, 'me nts'etsopele ena e bile le tšekamelo litloaelong tsa bocha ba bocha, haholo bokhoni ba ho laola le ho thibela boits'oaro bo susumetsang (Brenhouse et al., 2010, Geier et al., 2010, Sturman le Moghaddam, 2011a 'me Ernst, 2014). PFC e arotsoe ka likarolo tse fapaneng tse fapaneng tse fapaneng le litlamorao tse fapaneng bakeng sa boits'oaro ba bocha le ho ba kotsing ea mafu. Orbitof Pambal cortex (OFC) ke sebaka sa morao-rao se tlang pele se nkang maikutlo ho tsoa libakeng tsa maikutlo ebile se hokahane haholo le libaka tsa maoto (Theko, 2007 'me Rolls le Grabenhorst, 2008). Ka hona, OFC e loketse ho hokahanya likarolo tsa 'mele tsa melemo ea ho fana ka litholoana tse thabisang le tse khahlisang le tlhaiso-leseling ea maikutlo, ebe u sebelisa tlhaiso-leseling ena ho tataisa boitšoaro. Ts'ebetso ea Neuronal ho OFC e amahantsoe le boemeli ba liphetho tse ntle (van Duuren et al., 2007, Balleine et al., 2011 'me Schoenbaum et al., 2011), mme e bile le tšusumetso ho likarolo tse ngata tsa boits'oaro bo sa susumetseng (Berlin et al., 2004, Winstanley et al., 2010 'me Zeeb et al., 2010), e phahamisoang bathong le likhoto nakong ea bohlankana.Green le al., 1994, Adriani le Laviola, 2003, Burton le Fletcher, 2012, Doremus-Fitzwater et al., 2012 'me Mitchell le Potenza, 2014). Hobane OFC (hammoho le libaka tse ling tse tlang pele) li bontšitsoe li se li holile bongoaneng ba batho (Sowell et al., 1999 'me Galvan et al., 2006), OFC ke sepheo sa mantlha sa ho hlahloba liphapang tse amanang le lilemo ho ts'ebetsong ea moputso.
Ho rekota motho a le mong ho ile ha sebelisetsoa ho lekanya tšebetso e hlahisoang ke methapo ea kutlo. Ho batho ba baholo, ts'ebetso ea methapo ea bongata ea OFC e fokotsehile nakong ea ho khutla ha moputso (Feie. 1B). Ka lehlakoreng le leng, ts'ebetso ea baahi ea bongoaneng ea OFC e ile ea eketseha nakong ea ho khutla (Sturman le Moghaddam, 2011b). Phapang ena e matla ea ts'ebetso e etsahetse leha ho bile le litekanyetso tse tšoanang tsa motheo tsa ho thunya lipakeng tsa lihlopha, le tšitiso e ts'oanang ea neuronal nakong ea karolo ea ts'ebetso ea ketso ea ketso e lebisang ho phumants'o. Lintlha tsena li fana ka maikutlo a hore ts'ebetso ea moputso ho OFC e ka ba mofuta o sebetsang oa phapang e amanang le lilemo, leha ts'ebetso ea neuronal ea boitšoaro le boits'oaro li lekana lipakeng tsa lihlopha.
Le ha sekhahla sa ho thunya hoa motheo se ne se tšoana pakeng tsa lihlopha tsa lilemo, tlhahlobo e 'ngoe ea mekhoa ea ho thunya e senotse liphapang tse ling. Bacha ba OFC ba bonts'itse phapang e kholo ha e bapisoa le batho ba baholo ba ho thunya ho feta liteko tse ngata, joalo ka ha ho hlahlojoa ke ntho ea bohlokoa, e fanang ka phapang e fapaneng le maemo mme e ka baloa ka phapang ea teko ea ross ka moelelo oa teko ea sefapano (Churchland et al., 2010). Phapang ena e kanna ea bonts'a ho se hlophisehe hantle ha neural coding ea liketsahalo tse amanang le moputso, hobane ho fapana ka sekhahla ho nyenyefatsa puisano e sebetsang ea libaka tsa potoloho ka tšebelisano ea tšimo ea libaka (Meketso, 2005 'me Churchland et al., 2010). Habohlokoa, sepheo sena se fana ka maikutlo a hore mehato e fetang e bobebe ea ho thunya e ka hlokahala bakeng sa ho bona phapang e sebetsanang le tšebetso ea methapo pakeng tsa lihlopha tsa lilemo, mohlomong le lipakeng tsa taolo e phetseng hantle le bakuli ba kulang kapa ba kotsing.
OFC e bapala karolo ea bohlophisi boikhethong bo sa susumetsoang, bo hlalosoang e le khetho bakeng sa meputso / khotsofalo e potlakileng (Winstanley, 2007). Bacha ba lilemong tsa bocha le likhoto li ekelitse khetho ea bona ea ho ikhotsofatsa hanghang ha e bapisoa le batho ba baholo le likhoto, mme sena se bile le tšusumetso ho ts'ebeliso e mpe ea lithethefatsi ea bongoana le boits'ila bo fosahetseng (Adriani le Laviola, 2003, Doremus-Fitzwater et al., 2012, Mitchell le Potenza, 2014 'me Stanis le Andersen, 2014). Ho etsa liqeto ka potlako ho amana le mathata a mangata a kelello (Bechara et al., 2001, Ahn et al., 2011 'me Nolan le al., 2011), hape ke selelekela sa tšebeliso e mpe ea lithethefatsi le phello ea ho pepesetsoa lithethefatsi tsa tlhekefetso ka nako e telele (Simone et al., 2007, Perry et al., 2008, Anker et al., 2009, de Wit, 2009 'me Mendez et al., 2010). Kahoo, ho ka ba le boemo ba ho iphepa ba ho fepa ba bang moo batho ba nang le ts'oaetso ea kelello e kenyeletsang taolo e sa potlakileng ea ts'ebeliso e mpe ba ka sebelisang lithethefatsi hampe, e leng se eketsang tšusumetso ea bohlasoa (Garavan le Stout, 2005 'me Setlow et al., 2009). Lintlha tsa rona li fana ka maikutlo a hore phapang ea lilemo ho impulifity e ka ba, ka karolo e 'ngoe, ka lebaka la liphapang tsa ts'ebetso ea methapo ho OFC, ha OFC e kenyelletsa leseli mabapi le ho lieha ho amanang le moputso (Roesch le Olson, 2005 'me Roesch et al., 2006). Ts'ebetso e fapaneng haholo ea neural ts'ebetsong eohle ea ts'ebetso (joalo ka ha e lekantsoe ke setšoantšo sa mabaka) le karabelo e matlafalitsoeng ea moputso e fumanoang bongoaneng ba BOTSO, ka hona, e ka amana le lipontšo tse sa tsitsang tsa liketsahalo tse amanang le moputso. Seo re se bonang se kanna sa amana le bokhoni bo sa bonahaleng ba ho koala ho lieha ha nako e telele lipakeng tsa liketso le liphetho, mosebetsi o amanang le li-neuron tsa OFC (Roesch et al., 2006). Sena se ne se tla thusa ho khetha boikemisetso bo sa feleng ba ho ithabisa kapele.
Phapang e amanang le lilemo le eona e bonoa libakeng tse sa tsotelleng le tsa pele tsa PFC, tse amehang ho rala le ho fana ka maikutlo, tlhokomelo, le thibelo ea karabo (Goldman-Rakic, 1995, Fuster, 2001, Killcross le Coutureau, 2003, Magno et al., 2006, Peters et al., 2008, Burgos-Robles et al., 2013 'me Pezze et al., 2014). Ha ts'ebetso ea methapo ea kutlo e e-so rekotoe libakeng tsena mabapi le ho etsa liphoofolo tse lilemong tsa bocha, litlatsetso tsa nts'etsopele tsa ntlafatso ea meputso li se li senotsoe ka ho hlalosa liphatsa tsa lefutso tsa pele. Kamora ho itaola hoa heroin, bacha ba bontšitse keketseho ea li-neurons tse ntle tsa Fos ho li-cortices tsa prelimbic le infralimbic ha li bapisoa le batho ba baholo, e leng sesupo sa ts'ebetso e fokolitsoeng ea ts'ebetso ea bocha ea PFC ka ho batla moputso oa lithethefatsi (Doherty et al., 2013). Litlaleho tsa ts'ebetso e ts'oeroeng ke nicotine lia hanyetsana, li bonts'a keketseho e ntlafalitsoeng ea Arc kapa liphetoho tse tšoanang ho li-cfos lilemong tsa bocha ha li bapisoa le li-PFC tsa batho ba baholo (Leslie et al., 2004 'me Schochet et al., 2005). Kamora nako, ho pepesetsoa ha koae ho bakile polelo ea c-fos ho PFC ea lilemong tsa bocha (Cao et al., 2007). Le ha lithuto tsena li fana ka datha e sebetsang, litekanyo tse tobileng tsa ts'ebetso ea neural ea meputso ea lithethefatsi le ea tlhaho ho PFC ea bongaka ea bocha e tla fana ka tlhahisoleseling ea nakoana mabapi le ts'ebetso ea PFC ea medial ea bongoana.
Dopamine receptor expression in prelimbic cortex litlhōrō tsa bohlankana (Andersen et al., 2000). Li-receptor tsa D1 dopamine, ka ho khetheha, li hokahane le boitšoaro bo susumetsoang ba bongoaneng. Likhoto tsa lilemong tsa bocha li bontša ho ba kotsing e kholo ea likotsi tse amanang le lithethefatsi ha li bapisoa le likhoto tsa batho ba baholo (Leslie et al., 2004, Brenhouse le Andersen, 2008, Brenhouse et al., 2008 'me Kota et al., 2011); ho thibela li-receptor tsa D1 ho cortex ea bongoaneng e fokotsa kutloelo-bohloko ho lintlha tsena (Brenhouse et al., 2008). Ntle le moo, li-receptor tse matla tsa D1 tse tebileng ho batho ba baholo ba prelimbic cortex li fetotse litšekamelo tsa boitšoaro tsa bongoana, ho kenyelletsa ho susumetsa le ho eketseha ha maikutlo ho lithahasello tse amanang le lithethefatsi (Sonntag et al., 2014). Manolo a li-receptor a D1 a boetse a ntlafatsa maikutlo a boitšoaro ho amphetamine ka tekanyo e kholo ho bacha ho feta batho ba baholo.Mathews le McCormick, 2012).
2.2. Striatum
Kholiso ea methapo nakong ea bocha e ntse e tsoela pele ho striatum (Sowell et al., 1999, Ernst et al., 2006, Casey et al., 2008, Geier et al., 2010 'me Somerville et al., 2011). Striatum e kenella ho ho ithuta, ho lokisa moputso le ho sisinyeha, mme e bile le tšusumetso e matla mathateng a mafu a kelello ho kenyeletsa schizophrenia le ho lemalla (Kalivas le Volkow, 2005, Everitt et al., 2008 'me Horga le Abi-Dargham, 2014). Bobeli ba "ventral" le dorsal striatum ba fumana menahano ea dopaminergic e tsoang ho letsoele, mme phetiso ea dopamine e bontšitsoe khafetsa hore e fapana lipakeng tsa ho ba motho e moholo le lilemong tsa bocha.Adriani le Laviola, 2004, Volz et al., 2009 'me McCutcheon et al., 2012). Ha ho ntse ho e-na le leruo la lintlha tse tsoang mehlaleng ea liphoofolo tse hlalosang phapang ea neuroanatomical le pharmacological ho striatum lipakeng tsa litoeba tsa bocha le ba baholo (Andersen et al., 1997, Bolanos et al., 1998 'me Tarazi et al., 1998), ho na le lintlha tse fokolang tse hlalosang liphapang tse amanang le lilemo tsa ts'ebetso ea neural. Boholo ba lithuto tsa neano tsa ho nahana ka bongoana tse entsoeng lithutong tsa bongoana tsa batho li tsepamisitse maikutlo ho intral striatum (VS) haholo-holo li-nucleus accumbens (NAc), tse kenyelletsoeng ts'usumetso, ho ithuta le ts'ebetsong ea cue (Robbins le Everitt, 1996, Kelley, 2004, Ernst et al., 2006, Galvan et al., 2006, Geier et al., 2010 'me Hart et al., 2014). Leha ho le joalo, dorsal striatum (DS), e amehang ho ithuteng, khetho ea likhato le ho theha tloaelo (Packard le White, 1990, Balleine et al., 2007 'me Kimchi et al., 2009), e hlokomolohuoe haholo joaloka sebaka sa phapang ea ntlafatso. Ho lekola le ho bapisa li-logative tsa neural tsa ts'ebetso ea moputso libakeng ka bobeli tse tiileng, litebele tsa rona li tlalehe tšebetso ea kantle-kantle ho DS le NAc ea litheko tsa batho ba baholo le ba lilemong tsa bocha nakong ea boitšoaro bo tataisitsoeng.
Ka mokhoa o makatsang, ts'ebetso e hlahisang mosebetsi ho NAc e ne e sa fapana haholo pakeng tsa likhoto tsa batho ba baholo le ba lilemong tsa bocha (Sturman le Moghaddam, 2012). Phapang e amanang le lilemo tse matla, leha ho le joalo, e ile ea bonoa ho DS. Li-neuron tsa bongoana li ile tsa qobelloa pele ho ts'ebetso ea ho batla moputso, athe li-neurons tsa batho ba baholo ha lia ka tsa araba ho fihlela kamora ho qeta ketso (Feie. 1B). Li-neurons tsa bongoana ho DS le tsona li ile tsa qala ho etsoa pele ho phumano ea moputso, ha li-neuron tsa batho ba baholo li ne li thibetsoe ke moputso (Feie. 1B). Sena se bonts'itse hore boko ba bongoana bo hira circry ea DS pejana le ka tekanyo e kholo ho feta ea batho ba baholo nakong ea ho khutlisa moputso.
Le ha methapo ea kutlo ea DS ea lilemong tsa bocha e amohela meputso, ho lokolloa ha dopamine ea amphetamine ho bapisoa le batho ba baholo ba sebakeng sena. Maemo a tlase a amphetamine-evoke dopamine efflux ho DS, empa, hape, eseng NAc ea likhoto tsa bocha ha e bapisoa le batho ba baholo (Matthews le al., 2013). Ho khahlisang, phello e fapaneng e bonehetse le lithethefatsi tsa dopaminergic tse sebetsang e le tse thibelang likokoana-hloko tse kang cocaine le methylphenidate, tse bakang ntlafatso ea dopamine e le lilemong tsa bocha ha li bapisoa le DS e moholo.Walker le Kuhn, 2008 'me Walker et al., 2010). Joalo ka amphetamine, phello ena ea koae e amanang le lilemo e ile ea boleloa haholo ho DS ho feta NAc (Frantz et al., 2007 'me Walker le Kuhn, 2008). Phapang ena pakeng tsa ho lokolloa ha dopamine ea DS e ka ba mosebetsi oa ho fumaneha ha dopamine ea mantlha, kaha ho fokotseha ha dopamine ho fumanehang ha li-dopamine neurons ho ka ama lithethefatsi tse nolofalletsang ho lokolloa ha dopamine (joalo ka amphetamine) ka tekanyo e kholo ho feta lithethefatsi tse bolokang dopamine ho synapse (joalo ka joalo ka koae) Ka hona, tyrosine hydroxylase, enzyme e amanang le ho hlophisoa ha dopamine, e fokotsehile ho DS ea lilemong tsa bocha empa eseng NAc (Matthews le al., 2013). Phokotso ena ea evopamine-dopamine neurotransication e fana ka maikutlo a hore monahano oa dopamine ho DS, o hlahang ka lebaka la compacta eaantiantig nigra (Ungerstedt, 1971 'me Lynd-Balta le Haber, 1994), e kanna ea ba le kelello nakong ea bocha. Dopamine e na le tšusumetso ho thibela li-spiny neurons tse mahareng ka striatum (Kreitzer le Malenka, 2008). Hypoactive dopamine neurotransication nakong ea bocha DS e ka ba le tlatsetso mosebetsing oa rona o ntlafalitsoeng oa meputso ho li-neuron tsa DS. Boithuto ba nako e tlang ho rekota ho tloha ho dopamine ho ea ho mocha oa DS bo tla sebetsana ka kotloloho le mochine ona.
Sebaka sa striatum ka setso se amanang le ho hlahisa boleng le tšusumetso ho li-cu le meputso ke VS (Robbins le Everitt, 1996, Kelley, 2004, Cooper le Knutson, 2008 'me Flagel et al., 2011). Ka hona, likhopolo tse ngata tsa bokuli ba bongoana le ho ba kotsing ea boitšoaro li itšetleha ka boitšoaro bo susumetsoang bo amanang le moputso le karabelo ea ho potoloha ha boko bo amanang le moputso (Bjork et al., 2004, Galvan et al., 2006, Geier et al., 2010 'me Van Leijenhorst et al., 2010). Lintlha tse fetileng, ka lehlakoreng le leng, li fana ka maikutlo a hore liphapang tse amanang le lilemo le meputso li ka ba kholo le ho feta ho DS (Sturman le Moghaddam, 2012 'me Matthews le al., 2013). Le ha tsena li sa thibele karolo ea VS e ntseng e hola ea ho ba le boitšoaro ba bacha le ho ba kotsing ea mafu, ba fana ka maikutlo a hore DS le eona e ka bapala karolo e kholo litakatsong tsa boits'oaro ba bocha.
DS e amana haholo le ho ithuta le ponahatso ea 'mele ea boitšoaro bo sa rateheng (Robbins le Everitt, 1992, Packard le Knowlton, 2002 'me Gittis le Kreitzer, 2012). Haholo-holo sebaka sa dorsomedial striatum (DMS), kapa sebaka se kopaneng sa DS se bohlokoa ho hokahanya liketso le liphetho tse atlehileng, ha maqeba a DMS a felisa ho ithuta le ho hlahisa boitšoaro bo tataisoang ke sepheo (Yin le Knowlton, 2004 'me Ragozzino, 2007), mme ts'ebetso ea DMS le eona e kentsoe tšebetsong ea ho kenyelletsa mekhoa e bonolo ea karabelo (Kimchi le Laubach, 2009). Ka lehlakoreng le leng, dorsolateral striatum (DLS) e ameha ka kopanyo le tlhahiso ea boits'oaro, nakong eo liketso li sa hlolehang ho emela boemeli ba sephetho (Yin et al., 2004 'me Yin et al., 2009). Lithuto tsa ts'ebetso ea methapo ea bongoana le ho lokolloa ha dopamine li hlalositsoe ka botlalo ponong ena (Sturman le Moghaddam, 2012 'me Matthews le al., 2013) li ne li fumaneha ka kotlolloho ho DMS, li hatisa bohlokoa ba sebaka sena ntshetsopeleng malebana le boitšoaro ba bongoana le ho ba kotsing ea bokuli. Tumellanong le mohopolo ona, ho hlokometsoe liphapang tse 'maloa tšebetsong e sebetsang lipakeng tsa likhoto tsa batho ba baholo le ba lilemong tsa bocha,' me bacha ba bonts'a phapang ea boits'oaro, ho kenyelletsa le phapang ea takatso ea takatso ea lijo, ho fokotseha ha nako, ho sitisoa ha karabelo le bokhoni ba ho feto-fetoha ha liphetoho liketsong. sephetho sa maemo (Friemel et al., 2010, Sturman et al., 2010, Andrzejewski et al., 2011, Spear, 2011, Burton le Fletcher, 2012 'me Naneix et al., 2012). Ntle le moo, bacha ba bonts'a bokhoni ba ho fokotsa karabelo kapele ka mor'a ho emisa (Simone et al., 2013), e ts'oanang le phello e bonoang kamora ho ruruha ha DMS (Ntsu le Robbins, 2003).
Ho fapana le DMS ea lilemong tsa bocha, boteng ba liphapang tsa nts'etsopele ho DLS ha bo hlake hantle. Nakong ea polelo ea boitšoaro bo tataisoang ke sepheo, liketso li qala ho hokahanngoa ka thata le boemeli ba sephetho. Kamora ho bala ka ho fetelletseng, liketso li fetoha tse fokolang ho susumetsoa ke tlhahiso ea sephetho, 'me li iketsetse ("tloaelo") (Dickinson, 1985). Plasticity e amanang le mokhoa ona oa ho ithuta e hlaha ho DLS (Yin et al., 2009, Balleine le O'Doherty, 2010 'me Thorn et al., 2010), le phetoho ho tloha ho sepheo se lebisitsoeng ho ea ho boits'oaro bo tloaelo e arolelanoa ka karolo phetiso ea dopamine ho DS (Packard le White, 1991 'me Belin le Everitt, 2008). Ho na le lintlha tse hanyetsanang mabapi le nts'etsopele ea sebopeho sa tloaelo ho likotsi tsa batho ba lilemong tsa bocha le ho likhoto tse kholo. Likhoto tsa lilemong tsa bocha li bonts'a ho se khone ho fetola ho araba liphetoho maemong a tšohanyetso, hammoho le boits'oaro bo eketsehang mosebetsing oa ho matlafatsa boleng (Naneix et al., 2012 'me Hammerslag le Gulley, 2014). Ho na le bopaki ba ho tiea ha boits'oaro kapa ho fetoha ha maemo lithutong tsa bocha mabapi le mosebetsi o fetotsoeng ho bapisoa le batho ba baholo, ho ipapisitsoe le moralo oa mesebetsi le liparamente (Leslie et al., 2004, Newman le McGaughy, 2011 'me Snyder et al., 2014). Mesebetsi e meng e rarahaneng e bonahala e fana ka maemo a mangata ho feta a bocha. Mosebetsi oa ho fetola likhetho tse 'ne, o hlokang hore motho a lemohe haholo ho feta sebopeho se feto-fetohang sa likhetho tse peli, o senotse ho feto-fetoha ha maemo ho feta tekaneng ha ho bapisoa le litoeba tsa batho ba baholo (Johnson le Wilbrecht, 2011). Ntle le moo, tlhaiso-leseling ea morao-rao e bonts'a hore, kamora ho ithuta ho hana ho nka khato moo ho nang le boiphihlelo, likhoto tsa bocha li fumana mohopolo oo kapele joalo ka sekhahla se khothatsang sa Pavlovian sa moputso, joalo ka ha se lekantsoe ke keketseho ea boits'oaro ba mokhoa oa moputso. Sena se khothalelitse hore bacha ba khona ho fetola boleng ba mohopolo o neng o le bohlokoa pele (o fapaneng le mesebetsi e khutlisetsang morao morao, eo hangata e kenyelletsang boleng ba mohopolo o neng o sa lefuoe pele). Teko ea haufinyane laboratoring ea rona e lekile bokhoni bona ba ho ikamahanya le liphetoho tse mabapi le boitsebahatso bo eketsehileng ka ho koetlisa likhoto ka paradigm ea seletsa, nakong eo ho ileng ha hlahisoa leseli la leseli (leseli kapa molumo), 'me nko e kenang koung e bonesitsoeng phepelo ea pellet ea lijo. Ha ho phapang ea likarabo tse nepahetseng lipakeng tsa batho ba baholo le bacha e bonoeng mosebetsing ona (F(1,12) = .23, p = .64; n = 7 / sehlopha sa lilemo; Feie. 2). Karolong ea bobeli ea teko ena, mohopolo oa liletsa o ile oa fetoloa ka mokhoa o ikhethileng oa 10 s Pavlovian cue. Kamora phetoho ea kamano ea sephetho sa maikutlo, bacha ba bontšitse liperesente tse phahameng tsa mokhoa oa Pavlovia nakong ena ea maikutlo ho feta batho ba baholo, joalo ka ha ho lekantsoe ka nako e sebelisitsoeng ka sejelong sa lijo nakong ea khetsi (F(1,12) = 6.96, p = .023; Feie. 2). Thutong ea taolo, likhoto tsa bocheng le tsa batho ba baholo li ile tsa fumana mokhoa oa Pavlovian ho cate ea rialo ka tekanyo e lekanang, li bonts'a hore phello ena e ne e sa amane le phapang e amanang le lilemo ho bokhoni bo akaretsang ba ho ithuta kapa ho etsa boemo ba Pavlovian (F(1,12) = .26, p = .62). Lintlha tsena, ka hona, li bonts'a hore, ha mohopolo o sebetsa joalo ka lets'oao la ho emisa kapa la ho tsamaea ka har'a sesebelisoa, liphetoho likamanong tsa sephetho sa maikutlo li ka fumanoa habonolo ke likhoto tsa bocha kapele ho feta batho ba baholo. Tšobotsi ena ea boko ba bocha e ne e tla e lumella ho ikamahanya le liphetoho tsa boleng ba lintlha kapa maemo a neng a le bohlokoa pele ho feta boko ba motho e moholo. Ena ke phumano e khahlisang hobane lipatlisiso tse ngata ho bacha li shebane le boits'oaro bo sa lokang, athe ho feto-fetoha ha boits'oaro hangata ho khothalletsoa hore e be tšobotsi e ntle.
- Setšoantšo sa 2.
(A) Likhoto tsa batho ba baholo le ba lilemong tsa bocha ba ithutile ho etsa ketso ea bohlokoa bakeng sa moputso kamora tlhahiso ea sebui. (B) Cue eona eo e ile ea fetisetsoa ntlong ea Pavlovian, eo moputso o neng o se o sa tlisoe karabo, empa o ne o lula o fanoa joalo ka ha cue e felisoa. Likhoto tsa lilemong tsa bocha li fumane karabelo ea Pavlovian ho cue (e hlalositsoeng e le nako e sebelisitsoeng leqebelong la lijo tse lebelletseng nakong ea cue) ka potlako ho feta batho ba baholo.
Lintlha tse akaretsang li bontša hore likhoto tsa bocha li ka kenyelletsa likamano lipakeng tsa litakatso le liphetho moo litumelo li neng li utloahala ho feta bonolo ho feta batho ba baholo (Simone et al., 2013; Feie. 2), kapa maemong a nang le mojaro o mongata oa ho lemoha (Johnson le Wilbrecht, 2011). Boitšoaro bo matlafatsang ba kutlo bo bonoang ho DMS ea lilemong tsa bocha nakong ea liketsahalo tse amanang le moputso (Sturman le Moghaddam, 2012) e ka khothaletsa bokhoni bo eketsehileng ba ho fetola maano a boits'oaro (Kimchi le Laubach, 2009). E tla ba ntho e khahlisang ho rekota ho tloha DLS ea lilemong tsa bocha, e amanang le ho ithuta le mokhoa oa tloaelo, ho bona hore sebaka sena se na le tšusumetso e ntle ha se bapisoa le batho ba baholo. Ho thehoa tloaelo e potlakileng ho khothaletsoa ho khothaletsa bokhoba, hobane tloaelo ea lithethefatsi tse tloaelo ha e amehe ka litlamorao tse mpe tsa tlhekefetso ea lithethefatsi le ho lemalla.Everitt et al., 2008 'me Hogarth et al., 2013). Kahoo, thuto e tsoelang pele ea karolo ea DS e ntseng e hola ho theha tloaelo e bohlokoa haholo pakong ea bokhoba ba lithethefatsi tsa bongoana.
Ka bobeli DS le VS li kentse letsoho ho etsa liqeto tse kotsi (Mok'hadinale, 2006, Simone et al., 2011, Kohno et al., 2013 'me Mitchell et al., 2014), e hlalosoang e le khetho bakeng sa meputso e bolokehileng ho feta meputso e bolokehileng. Boitšoaro bo kotsi ke lets'oao la bohlankana, 'me bo hokahane le ts'ebeliso e mpe ea lithethefatsi (Bornovalova et al., 2005 'me Balogh et al., 2013). Ho feta moo, bopaki ba morao-rao bo tsoang mohlaleng oa likhakanyo tsa ho etsa liqeto tse kotsi bo bontša hore boits'oaro bo kotsi ho bacha bo bolela esale pele hore ho tla ba le taolo ea koae.Mitchell et al., 2014), e ka nolofalletsang ts'ebeliso e mpe ea lithethefatsi le ho lemalla tlhekefetso lilemong tsa bocha (Adriani le Laviola, 2004, Merline et al., 2004 'me Doremus-Fitzwater et al., 2010). Phokotso ea dopamine receptor e fumanehang libakeng ka bobeli tse nang le bokhoni ba ho nka liqeto tse kotsi likerekeng, le ts'oaetso ea lehae ea litheko tsa "dopamine agonist" tse etsahalang ka mokhoa o hlophisehileng kapa tse kenang lilemong tsa bocha li fokotsa boitšoaro bo kotsi (Simone et al., 2011 'me Mitchell et al., 2014). Ka ho tšoanang, likhoto tsa bocha li bonts'a karabelo ea dopamine e fokolang le polelo ea TH ho DS (Matthews le al., 2013), e ka fanang ka mokhoa o batlang o lekana bakeng sa boits'oaro bo bobe ba lilemong tsa bocha. Ho etsa liqeto ka kotsi le hona ho amana le ts'ebetso ea neuronal le polelo ea dopamine receptor ho OFC (Eshel et al., 2007, Van Leijenhorst et al., 2010, Simone et al., 2011 'me O'Neill le Schultz, 2013). Ho ka etsahala hore likarabo tsa moputso oa hyperactive ho bobeli ba OFC le DS (Sturman le Moghaddam, 2011b 'me Sturman le Moghaddam, 2012) li amana le ho etsa liqeto tse kotsi le tse ngata ka nako e itseng lilemong tsa bocha. Boithuto bo tsoelang pele ba potoloho ena bo ka hlahisa datha tse khahlisang le likhetho tsa kalafo bakeng sa methati ea pele ea mafu a khetholloang ka boits'oaro bo kotsi bo hlahang nakong ea bocha, ho kenyelletsa le bokhoba, lefu la kelello le khatello ea maikutlo (Ludewig et al., 2003, Bornovalova et al., 2005 'me Taylor Tavares et al., 2007).
2.3. Sebaka se ikhethileng sa boteng
Dopamine neurons, haholo-holo tse fumanehang sebakeng sa ventral tegmental area (VTA), li kenelletse ts'ebetsong ea meputso, ho ithuta ka likamano, le pathophysiology ea lithethefatsi, mathata a maikutlo le lefu la kelello (schizophrenia)Bohlale le Bozarth, 1985, Schultz, 1998, Bohlale, 2004, Sesack le Grace, 2010 'me Howes et al., 2012). Sisteme ea dopamine e bile le tšusumetso e matla litekanyetsong tsa bocha tsa bocha le bokuli (Luciana et al., 2012, Matthews le al., 2013 'me Niwa et al., 2013), le likarolo tsa phetisetso ea dopamine le ts'ebetso ea VTA li fapane ho batho ba baholo le ba lilemong tsa bocha (Robinson et al., 2011, McCutcheon et al., 2012 'me Matthews le al., 2013). Ntle le moo, dopamine neurons morerong oa VTA ho preortal cortex le ventral striatum, libaka tse ntseng li tsoela pele nakong ea bocha. Ha ho tsejoehale leha ho le joalo, mabapi le hore na li-neurons tsa VTA tsa lilemong tsa bocha li etsa moputso joang liketsahalong tse amanang le batho ba baholo. K'hamphani ea morao-rao ea ho rekota ea liketsahalo tsa kantle ho lisele tsa VTA ho likhoto tsa batho ba baholo le ba lilemong tsa bocha e bontša hore li-neurons tsena li na le sekhahla se tšoanang sa ho thunya le ho arabela likotlo tse amanang le moputso (Kim le Moghaddam, 2012), mme mosebetsi o ntse o tsoela pele o tla lekola ts'ebetso ea moputso oa bongoana ho sena, le libakeng tse ling tsa dopaminergic.
2.4. Kakaretso ea moputso oa potoloho
Bacha ba bonts'a boitshwaro bo matlafatsang, ho ipeha kotsing, ho se tsitse, ho ts'oara lithethefatsi le ho fumana moputso, le ho fetoha ha maemo ha ho bapisoa le batho ba baholo. Joalokaha ho hlalositsoe ka holimo, yuniti e le 'ngoe ea electrophysiology e senotse liphapang tse amanang le lilemo ho ts'ebetsong ea moputso tse ka amehang litakatsong tsena tsa boits'oaro. Bacha ba bonts'a ts'ebetso ea hyper-activation ho putsa motho ea amanang le batho ba baholo ka bobeli ba OFC le DS (Feie. 3). OFC e kenella ka kotloloho ho DS, bonyane ka litoeba tsa batho ba baholo, e fana ka maikutlo a hore ho hokahana ha OFC-DS le hona ho ka kenya letsoho liphellong tsena tse bonoang (Berendse et al., 1992 'me Reep et al., 2003). Dopaminergic neurons tse qalang ho tloha ho projekeanti nigra le tsona li tla ho DS (Voorn et al., 2004), le ts'ebetso e khahlisang ea meputso methapong ena ea methapo e ka kenya letsoho ts'ebetsong ea meputso ea DS nakong ea bocha. Motsoako o theotsoeng oa dopamine o bonoang ho DS o latelang tlhahiso ea amphetamine o fana ka maikutlo a hore li-neuron tsena ka sebele li ka ba le khatello ea maikutlo ha li bapisoa le batho ba baholo, leha liteko tse ling li hlokahala ho netefatsa phapang ena e sebetsang. Ts'ebetso e hlahisitsoeng ke moputso ho DLS, e fumanang phepelo e matla ka ho fetisisa ea dopaminergic e tsoang hoantiantia nigra (Groenewegen, 2003 'me Voorn et al., 2004), hape e kanna ea fapana lipakeng tsa batho ba baholo le ba lilemong tsa bocha, ha kholo ea mekhoa ea boitšoaro e fapana ho pholletsa le nako ea bophelo (Johnson le Wilbrecht, 2011, Newman le McGaughy, 2011, Simone et al., 2013 'me Snyder et al., 2014).
- Setšoantšo sa 3.
Potoloho ea moputso e fetotsoeng bakeng sa boko ba bocha. Khokahano ea "mekhahlelo e tloaelehileng ea moputso" e bontšoa ka bosoeu le ho kenyelletsa li-nucleus accumbens (NAc), sebaka sa ventral tegmental (VTA), le medial prefrontal cortex (mPFC). Seo re se fumaneng ho bacha se supa tsela e tlatselletsang ea moputso e bonahatsoang ka bofubelu. Re fumana hore litebello tsa dopamine ho dorsal striatum (DS), tse hlahang ka lebaka laantiantia nigra (SNc) li ka ba le tšibollo ea maikutlo ho bacha (Matthews le al., 2013) ha orbitofrontal cortex (OFC) le li-neurons tsa DS tsa bongoana li arabela hantle ho feta moputso ha li bapisoa le batho ba baholo (Sturman le Moghaddam, 2011a, Sturman le Moghaddam, 2011b 'me Sturman le Moghaddam, 2012). Ka lehlakoreng le leng, NAc dopamine e lokolloang le ts'ebetso e fuoang moputso, le ho chechisa methapo ea methapo ea dopamine sebakeng se bitsoang (ventral tegmental sebakeng seo (VTA)) li ka bapisoa pakeng tsa batho ba baholo le bacha (Kim le Moghaddam, 2012 'me Matthews le al., 2013).
Ho khahlisang, ha ho phapang e kholo e amanang le lilemo e ileng ea bonoa ts'ebetsong ea moputso oa NAc, leha VS e le ntho e hlahelletseng mehlaleng ea bosholu ba boits'oaro ba bacha (Ernst et al., 2009 'me Geier et al., 2010). Ts'ebetso ena e tšoanang ea neural lipakeng tsa lihlopha tsa lilemo e lumellana le litlaleho tsa ho se tšoane ho amanang le lilemo mokhoeng oa dopamine o hlahisitsoeng ke lithethefatsi ho NAc (Frantz et al., 2007 'me Matthews le al., 2013), leha lithuto tse mabapi le polelo ea dopamine receptor ho NAc li hanyetsana (Teicher et al., 1995 'me Tarazi le Bal)arini, 2000). Ho haella ha liphapang ho ts'ebetso ea moputso oa NAc ha ho qobelle tšusumetso ea NAc ea bongoaneng ea ho ba le boits'oaro bo bobe ba boitšoaro le psychopathology; leha ho le joalo, phapang e lemohileng lits'ebetsong tsa ts'usumetso nakong ea bohlankana (Spear, 2011) e ka hlaha ho tsoa tšebetsong tšebetso ea neural libakeng tsa DS le PFC ho ea holimo ho feta NAc. Ka kopanelo, liphumano tsena li fana ka maikutlo a hore lipallo tsa bongaka ba moetlo li lokela ho fetoloa bakeng sa bacha (Feie. 3).
3. Qetello
Liphumano tse hlahlojoang mona li tsebisa lipatlisiso tsa nakong ea bongoana ka mekhoa e 'meli: (1) ts'ebetso ea mantlha kapa karabelo molemong oa maikutlo joalo ka ha meputso e boletsoeng esale pele e sa angoe, kapa e amehile hofeta, ho feta ts'ebetso ea neuronal ho potoloha nako ea moputso. Kahoo, ho tsepamisa maikutlo karabong ea moputso ho ka fana ka biomarker e ntle bakeng sa ho ba kotsing ea ho hlaseloa habonolo ke mafu a susumetsang le a amang. (2) Likarabo tsa neuronal tse matla li ile tsa bonoa libakeng tse sa amaneng le ts'ebetso ea moputso ho batho ba baholo. Kahoo, potoloho ea matla ea boits'oaro bo susumetsang e kanna ea fapana le mehlala ea rona ea batho ba baholo mme e kenyelletsa libaka tsa cortical le basal ganglia tse sa amaneng le ts'ebetso ea moputso. Ho hatella nako e tlang libakeng tse kang DS ho ka ntlafatsa tsebo ea rona ea potoloho ena e matla le tlatsetso ea eona bothateng ba mafu a batho ba kotsing.
Likhohlano tsa thahasello
Bangoli ba phatlalatsa hore ha ho na likhohlano tsa phaello.
Liteboho
Mosebetsi ona o ne o tšehelitsoe ke DA EA 035050 (NWS) le MH048404-23 (BM).
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