Dopamine ka taolo ea ho susumetsa: ho putsa, ho pheta-pheta le ho lemosa (2010)

Sheba lihlooho tse ling ho PMC hore Qotsa sehlooho se hatisitsoeng.

Li-neurop tsa dbamine tsa Midbrain li tsebahala ka likarabo tsa tsona tse matla ho meputso le karolo ea tsona ea bohlokoa ho khothaletso e ntle. Ho ntse ho hlaka le ho feta, leha ho le joalo, hore li-dopamine neurons le tsona li fetisetsa lipontšo tse amanang le boiphihlelo bo sa reroang empa bo sa ruiseng letho bo kang liketsahalo tse khopisang le tse lemosang. Mona re lekola tsoelo-pele ea morao-rao ho utloisisa moputso le mesebetsi e seng moputso ea dopamine. Motheong oa data ena, re etsa tlhahiso ea hore li-dopamine neurons li tle ka mefuta e mengata e hokahaneng le marang-rang a fapaneng a boko mme li na le likarolo tse fapaneng taolong e susumetsang. Tse ling tsa dopamine neurons encode boleng bo susumetsang, tse tšehetsang marang-rang a boko bakeng sa ho batla, ho lekola, le ho ithuta boleng. Ba bang ba koala sekhahla se tšehetsang kelello, se ts'ehetsang marang-rang a ts'ebetso ea kelello, tlhalohanyo le tšusumetso e akaretsang. Mefuta ka bobeli ea li-neurop tsa dopamine e matlafatsoa ke lets'oao le lemosang la ho lemoha ka potlako mekhoa e mebe ea bohlokoa ea maikutlo. Re nahana ka botebo hore litsela tsena tsa dopaminergic bakeng sa boleng, sesapo, le tlhokomeliso li sebelisana ho ts'ehetsa boitšoaro bo feto-fetohang.

Dopamine ka ts'usumetso ea liketso tse batlang meputso

Dopamine haesale e tsejoa e le ea bohlokoa bakeng sa ho matlafatsa le ho susumetsa liketso. Lithethefatsi tse sitisang phetisetso ea DA li kena-kenana le ho ithuta ho matlafatsa, athe litsoantso tse ntlafatsang phetisetso ea DA, joalo ka ts'usumetso ea boko le lithethefatsi tse tlatsetsang, hangata li sebetsa e le litlatsetso (Bohlale, 2004). Ho fetisoa ha DA ho bohlokoa bakeng sa ho theha boemo ba tšusumetso ea ho batla meputso (Berridge le Robinson, 1998; Salamone et al., 2007) le ho theha mehopolo ea mekhatlo ea moputso oa cue (Dalley et al., 2005). Ho lokolloa ha DA ha ho hlokahale bakeng sa mefuta eohle ea ho ithuta ka moputso mme e kanna ea se ke ea 'ratoa' ka nako ea maikutlo a ho thabisa, empa ho bohlokoa bakeng sa ho etsa hore lipheo li 'hloloe' ka mohopolo oa liketso tse susumetsang ho li fihlela (Berridge le Robinson, 1998; Palmiter, 2008).

Khopolo-taba e 'ngoe mabapi le kamoo dopamine e tšehetsang thuto ea matlafatso ke hore e ntlafatsa matla a khokahano ea synaptic pakeng tsa methapo ea kutlo. Mofuta o hlakileng ka ho fetesisa oa hypothesis ena ke hore dopamine e laola polasetiki ea synaptic ho latela molao o fetotsoeng oa Sehebberia o ka reng ke "li-neurons tse tukang terata hammoho, ha feela li fumana ho phatloha ha dopamine". Ka mantsoe a mang, haeba sele A e kenya ts'ebetsong sele ea B, mme sele ea B e baka ts'ebetso ea boits'oaro e lebisang moputsong, dopamine e tla lokolloa mme khokahano ea A → B e matlafatsoe (Montague et al., 1996; Schultz, 1998). Mochine ona o ka lumella setho sa 'mele hore se ithute khetho e nepahetseng ea liketso hore se fumane meputso haeba se na le boiphihlelo bo lekaneng ba phoso le phoso. E lumellana le khopolo ena, dopamine e na le tšusumetso e matla ea polasetiki ea synaptic libakeng tse ngata tsa boko (Surmeier et al., 2010; Goto et al., 2010; Molina-Luna et al., 2009; Marowsky et al., 2005; Lisman le Mohau, 2005). Maemong a mang dopamine e nolofalletsa polasetiki ea synaptic hammoho le mela ea molao oa Mahebrania o hlalositsoeng kaholimo, ka tsela e lumellanang le boits'oaro ba ho batla moputso (Reynolds et al., 2001). Ntle le litlamorao tsa eona tsa polasetiki ea synaptic ea nako e telele, dopamine e ka boela ea fana ka taolo ea hanghang holim'a li-circuits tsa neural ka ho fetolela ts'ebetso ea spike ea neural le likhokahano tsa synaptic pakeng tsa li-neurons (Surmeier et al., 2007; Robbins le Arnsten, 2009), maemong a mang ho etsa joalo ka mokhoa o tla khothaletsa liketso tsa ho batla moputso hanghang (Frank, 2005).

Dopamine neuron lipontšo tsa moputso

Bakeng sa ho susumetsa liketso tse lebisang meputsong, dopamine e lokela ho lokolloa nakong ea liphihlelo tse khotsofatsang. Ho joalo, boholo ba li-neuron tsa DA li matlafalitsoe ka matla ke meputso ea mantlha e sa lebelloang joalo ka lijo le metsi, hangata e hlahisa 'menyetla "ea phasic ea ts'ebetso (Schultz, 1998) (thabo)Mohau le Bunney, 1983)). Leha ho le joalo, lithuto tsa bo-pula-maliboho tsa Wolfram Schultz li bonts'itse hore likarabo tsena tsa DA tsa neuron ha li bakoe ke tšebeliso ea meputso ka bobeli. Sebakeng seo ba tšoana le 'phoso ea polelo ea moputso', ba tlaleha phapang lipakeng tsa moputso o amoheloang le moputso o neng o boletsoe esale pele hore o tla etsahala (Schultz et al., 1997) (Setšoantšo sa 1A). Kahoo, haeba moputso o le moholo ho feta o boletsoeng esale pele, li-neuron tsa DA li thabile haholo (phoso e boletsoeng esale pele ea ho bolela, Setšoantšo sa 1E, khubelu); haeba moputso o le monyane ho feta o boletsoeng esale pele kapa o hloleha ho hlaha ka nako e behiloeng, li-neuron tsa DA li thibeloa ka mokhoa o pharaletseng (phoso e mpe ea ho bolela esale pele, Setšoantšo sa 1E, putsoa); mme haeba moputso o buloa esale pele e le hore boholo ba eona bo ka boleloa ka botlalo, li-neurons tsa DA ha li na karabelo e fokolang kapa ha li na karabo (phoso e boletsoeng esale pele ea zero, Setšoantšo sa 1C, e sootho). Molao-motheo o ts'oanang o ts'oara likarabo tsa DA lithutong tsa maikutlo tse fanang ka tlhaiso-leseling e ncha ka meputso ea nako e tlang. Li-neuron tsa DA li nyakalla ha setšoantšo se bontša ho nyoloha ha moputso oa nako e tlang (Setšoantšo sa 1C, bofubelu), ho thibetsoe ha cue e bontša ho fokotseha ha boleng ba moputso bo tlang (Setšoantšo sa 1C, boputsoa), mme ka kakaretso ha ba na karabelo e nyane ho litloaelo tse sa hlahiseng tlhahisoleseling e ncha ea moputso (Setšoantšo sa 1E, e sootho). Likarabo tsena tsa DA li tšoana le mofuta o itseng oa phoso ea ho bolela moputso kapa phoso ea "TD phoso", e reriloeng ho sebetsa e le lets'oao la ho matlafatsa boleng ba ho ithuta boleng ba liketso le linaha tsa tikoloho (Houk et al., 1995; Montague et al., 1996; Schultz et al., 1997). Mefuta ea likhomphutha e sebelisang lets'oao le matlafatsang la TD e ka hlalosa likarolo tse ngata tsa ho matlafatsa thuto ho batho, liphoofolo le li-neuron tsa DA ka bo tsona (Sutton le Barto, 1981; Waelti et al., 2001; Montague le Berns, 2002; Dayan le Niv, 2008).

 

Setšoantšo sa 1

Lekopolla likh'outu tsa liphoso tsa ho bolela moputso le khetho ea tlhaiso-leseling

Tlhahlobo e makatsang ea liteko li bonts'itse hore matšoao a DA a emetse likhakanyo tsa moputso ka tsela e lumellanang haufi le litabatabelo tsa boitšoaro, ho kenyelletsa le khetho ea moputso o moholo ho feta tse nyane (Tobler et al., 2005) meputso e ka bang teng bakeng sa e ke keng ea etsahala (Fiorillo et al., 2003; Satoh et al., 2003; Morris et al., 2004) le moputso o potlakileng bakeng sa tse liehang (Roesch et al., 2007; Fiorillo et al., 2008; Kobayashi le Schultz, 2008). Ho na le bopaki hape ba hore li-neurons tsa DA ho batho li kenyelletsa boleng ba moputso (Zaghloul et al., 2009). Ntle le moo, matšoao a DA a hlaha nakong ea ho ithuta ka nako e ts'oanang ea linako tsa boits'oaro ba ho bolela meputso (Hollerman le Schultz, 1998; Satoh et al., 2003; Takikawa et al., 2004; Letsatsi le al., 2007) mme e amana le methati e ikhethileng ea khetho ea moputso (Morris et al., 2006). Liphumano tsena li thehile li-neuron tsa DA e le e 'ngoe ea mehlala e utloisisitsoeng hantle le e phetoang ka ho fetisisa ea moputso bokong. Ka lebaka leo, lipatlisiso tsa morao-rao li kentse li-neurons tsa DA ho hlahlobisisa haholo ho fumana hore na ba hlahisa liphatlalatso tsa moputso joang le hore na matšoao a bona a sebetsa joang mehahong e tlase ho laola boitšoaro.

Dopamine neuron lipontšo tsa moputso

Khatelo-pele ea morao-rao ho utloisiseng matšoao a moputso oa DA e tsoa ho ho latela lipotso tse tharo tse pharaletseng: Boko ba DA bo ithuta joang likhakanyo tsa moputso? Seo ba se boletseng esale pele se nepahetse hakae? Mme ke eng se ba se tsayang e le se se duelang?

Li-neurons tsa DA li ithuta joang likhakanyo tsa moputso? Likhopolo tsa khale li fana ka maikutlo a hore likhakanyo tsa moputso li ithutoa ka ts'ebetso ea ho matlafatsa butle butle e hlokang li-pair tsa moputso tsa khothaletso (Rescorla le Wagner, 1972; Montague et al., 1996). Nako le nako ha khothalletso ea A e lateloa ke moputso o sa lebelloang, boleng bo hakantsoeng ba A bo eketsehile. Lintlha tsa morao-rao li bontša hore li-neurons tsa DA ha li felle feela ka ho ithuta ka mokhoa o bonolo oa ho khothatsa le ho bolela esale pele tumelo e thehiloeng holima tumelo ea sebopeho sa lefatše. Li-neuron tsa DA li ka bolela moputso hantle esita le maemong a sa tloaelehang moo meputso e entsoeng ka khothalletso e etsang hore a fokotseha boleng ba khothalletso (Satoh et al., 2003; Nakahara et al., 2004; Bromberg-Martin et al., 2010ckapa etsa hore ho be le phetoho boleng ba khothalletso e fapaneng ka ho fetesisa (Bromberg-Martin et al., 2010b). Li-neuron tsa DA li ka fetolela lipontšo tsa moputso ho latela lipalo-taelo tse phahameng tsa kabo ea meputso, joalo ka ho phahamisa matšoao a boletsoeng esale pele a nepileng ho latela phapang ea bona e lebelletsoeng (Tobler et al., 2005) le 'ho hlaphoheloa' likarabo tsa bona ka boithatelo ho likarabo tsa meputso e timelitsoeng (Pan et al., 2008). Tsena tsohle li etsahala ka mokhoa o ts'oanang o tšoanang le litlamorao tse ts'oanang tse bonoang mohopolong oa maikutlo le oa koloi (Braun et al., 2010; Fairhall et al., 2001; Shadmehr et al., 2010), ho fana ka maikutlo a hore li ka bonts'a mokhoa o tloaelehileng oa neural oa ho ithuta esale pele.

Bolepi ba moputso oa DA bo nepahetse hakae? Phuputso ea morao tjena e bontšitse hore li-neurons tsa DA li fetola ka nepo lipontšo tsa tsona tsa moputso hore li ikarabelle bakeng sa mehloli e meraro ea ho se kholoe esale pele. Taba ea mantlha, batho le liphoofolo ba utloisoa bohloko ke lerata la nako e ba thibelang ho bolela esale pele ka linako tse telele tsa meputso (Gallistel le Gibbon, 2000). Kahoo, haeba tebello ea moputso oa cue-khutšoane e le khutšoanyane (metsotsoana ea 1-2) nako e boletsoeng esale pele e nepahetse mme phumants'o ea meputso e baka karabelo e nyane ea DA, empa bakeng sa tebello e teletsana ea tefello ea nako ea tefo e tla fetoha e sa ts'epahale mme meputso e etsa hore ho be le menyetla e hlakileng ea DA (Kobayashi le Schultz, 2008; Fiorillo et al., 2008). Taba ea bobeli, litloaelo tse ngata bophelong ba letsatsi le leng le le leng ha li makatse, ho hlakisa ho ajoa ha linako tse ngata tsa kabo ea meputso. Li-neurons tsa DA li boetse li bonts'a mokhoa ona oa ho hloka bonnete ba nako: li thibelloa butle-butle nakong ea ho lieha ha meputso, joalokaha eka ho tšoaea liphoso tse boletsoeng esale pele tsa moputso ka nako eohle moputso o hloleha ho hlaha (Fiorillo et al., 2008; Bromberg-Martin et al., 2010a; Nomoto et al., 2010). Kamora nako, litloaelo tse ngata li rarahane ka tsela e hlakileng, li hloka tlhahlobo e qaqileng ho fihlela qeto e tiileng mabapi le boleng ba moputso. Maemong a joalo lipontšo tsa moputso oa DA li ba teng ka bolelele bo bolelele le ka mokhoa o tsoelang pele, ho bonahala o bonts'a phallo ea butle-butle ea tlhaiso-leseling ha boleng ba ts'usumetso bo phatlalalitsoe (Nomoto et al., 2010).

Ke liketsahalo life feela tse etsang hore methapo ea DA e utloe e le moputso? Likhopolo tse tloaelehileng tsa thuto ea moputso li khothaletsa hore li-neurons tsa DA li abele boleng bo thehiloeng ho palo e lebelletsoeng ea moputso oa mantlha oa ka moso (Montague et al., 1996). Leha ho le joalo leha tekanyo ea moputso oa mantlha e lula e tšoaretsoe khafetsa, batho le liphoofolo hangata ba bontša khetho e eketsehileng bakeng sa boikemisetso - ba batla tikoloho eo boholo ba moputso, monyetla le nako li ka tsejoang pele ho nako (Daly, 1992; Chew le Ho, 1994; Ahlbrecht le Weber, 1996). Phuputso ea morao-rao ea litšoene e fumane hore li-neurons tsa DA li supa khetho ena (Bromberg-Martin le Hikosaka, 2009). Litšoene li bontšitse boikhethelo bo matla ba ho shebella lintlha tse ka ba thusang tse tla ba lumella ho bolela esale pele boholo ba moputso oa nako e tlang, ho fapana le lintlha tse sa fetoheng tse sa faneng ka tlhaiso-leseling e ncha. Ka mokhoa o tšoanang, li-neuron tsa DA li ne li natefeloa ke monyetla oa ho bona lintlha tse fanang ka tsebo ka tsela e lumellanang le litabatabelo tsa phoofolo.Setšoantšo sa 1B, D). Sena se fana ka maikutlo a hore li-neurons tsa DA ha li khothaletse liketso feela ho fumana meputso empa li boetse li susumetsa liketso ho etsa liqeto tse nepahetseng mabapi le meputso eo, e le ho netefatsa hore meputso e ka lebelloa ka nepo le ho itokisetsa esale pele.

Ha re kopantsoe hammoho, liphumano tsena li bonts'a hore lipontšo tsa phoso ea moputso oa DA li nahanela lintlha tsa morao-rao tse tsebisang batho meputso ea batho le liphoofolo, ho kenyelletsa le lipalo tsa lipalo tsa meputso e holimo, moputso o sa tsitsang, le khetho ea tlhaiso-leseling.

Liphello tsa lipontšo tsa moputso oa phasic dopamine liporong tse tlase

Likarabo tsa moputso oa DA li hlaha ho phatloha ho hoholo ha phasic (Joshua et al., 2009b), mokhoa oa karabo o bopang DA e lokollang mehahong ea sepheo (Gonon, 1988; Zhang et al., 2009; Tsai et al., 2009). Ke khale ho nahanoa hore litaba tsena tsa phasic li susumetsa ho ithuta le ho susumetsa ka tsela e fapaneng le ea tšebetso ea DA ea tonic.Mosa, 1991; Mohau et al., 2007; Schultz, 2007; Lapish et al., 2007). Mahlale a morao-rao a ntlafalitsoeng a entse hore ho khonehe ho netefatsa khopolo ena ka ho laola ts'ebetso ea neuron ea DA ka tatellano e ntle ea sebaka le sa nakoana. Ts'usumetso ea Optogenetic ea li-neurons tsa VTA DA e etsa hore motho a rate sebaka se matla se etsahalang ha feela ts'usumetso e sebelisoa ka mokhoa oa ho phatloha (Tsai et al., 2009). Ka lehlakoreng le leng, ho hlaseloa ka liphatsa tsa lefutso tsa li-receptor tsa NMDA ho li-neuron tsa DA, tse sitisang ho phatloha ha o tlohela ts'ebetso ea tonic haholo, ho baka ho senyeha hoa mefuta e ikhethileng ea ho ithuta ka moputso (Zweifel et al., 2009; Parker et al., 2010) (Le ha ho le joalo hlokomela hore ho kokota hona ho senya leeme la DA la neuron synaptic (Zweifel et al., 2008)). Ho phatloha ha DA ho ka ntlafatsa thuto ea moputso ka ho ntlafatsa li-circuits tsa lehae tsa lehae. Haholo, ho phatloha ha mofufutso oa DA esale pele ho romeloa libakeng tse ikhethang tsa li-bokellase, 'me libaka tsena li na le liketsahalo tse phahameng haholo tsa ho rera moputso esale pele (Cheer et al., 2007; Owesson-White et al., 2009).

Ha ho bapisoa le ho phatloha ho phasic, hanyane ho tsejoa ka bohlokoa ba ho khefutsa ha boqhetseke mosebetsing oa spiking bakeng sa liphoso tse boletsoeng esale pele tsa moputso. Khefutso ena e baka liphetoho tse nyane ka sekhahla sa spike, ha e fetoloe ka tebello ea moputso (Bayer le Glimcher, 2005; Joshua et al., 2009a; Nomoto et al., 2010), hape e ka ba le litlamorao tse nyane ho ithuteng (Rutledge et al., 2009). Le ha ho le joalo, mefuta e meng ea thuto e fosahetseng ea ho bolela esale pele e hloka VTA (Takahashi et al., 2009), ho fana ka maikutlo a hore likhechana tsa phasic li kanna tsa hlalosoa ka meaho e tlase.

Kaha ho phatloha le ho phomola ho baka mekhoa e fapaneng haholo ea ho lokolloa ha DA, ba kanna ba susumetsa meaho e thellang ka methati e ikhethileng. Ho na le bopaki ba morao-rao ba khopolo ena ho sepheo se le seng se seholo sa li-neurons tsa DA, e leng dorsal striatum. Dorsal striatum proion neuron e tla ka mefuta e 'meli e bontšang li-receptor tse fapaneng tsa DA. Mofuta o mong o hlahisa li-receptor tsa D1 le merero ho "tsela e tobileng" ea basal ganglia ho tsamaisa motsamao oa 'mele; mofuta oa bobeli o hlahisa li-receptor tsa D2 le merero ho "tsela e sa tsejoeng" ho hatella ho sisinyeha ha mmele (Setšoantšo sa 2) (Albin le al., 1989; Gerfen et al., 1990; Kravitz et al., 2010; Hikida et al., 2010). Ho ipapisitsoe le likarolo tsa litsela tsena le li-receptor, ho tlalehiloe hore li-DA li phatloha li hlahisa maemo a DA a phahameng, li kenya li-receptor tsa D1, 'me li etsa hore tsela e otlolohileng e khethe mekhatlo ea boleng bo holimo (Setšoantšo sa 2A), athe DA e khefutsa e hlahisa maemo a DA a tlase, inhibit D2 receptors, 'me e etsa hore tsela e sa tobang e hatelle metsamao e theko e tlase (Setšoantšo sa 2B) (Frank, 2005; Hikosaka, 2007). E lumellana le khopolo ena, ts'ebetso e phahameng ea receptor ea DA e phahamisa bokhoni ba li-synapses tsa cortico-striatal tseleng e tobileng (Shen et al., 2008) le ho ithuta ho tsoa liphellong tse ntle (Frank le al., 2004; Voon et al., 2010), ha D1 li-receptor blockade ka mokhoa o khethang li senya meketjana ho lipehelo tse fuoeng (Nakamura le Hikosaka, 2006). Ka mokhoa o makatsang, ts'ebetso e tlase ea li-receptor tsa DA e phahamisa bokhoni ba li-synapses tsa cortico-striatal tseleng e sa tobang (Shen et al., 2008) le ho ithuta ho tsoa liphellong tse mpe (Frank le al., 2004; Voon et al., 2010), ha D2 ea li-receptor blockade e khetha ka mokhoa o khethollang ho sisinyeha ho lipheo tse sa lefuoeng (Nakamura le Hikosaka, 2006). Karohano ena ea D1 le D2 li-receptor li sebetsa ka taolo ea susumetso e hlalosa litlamorao tse ngata tsa liphatsa tsa lefutso tse amanang le DA ka boitšoaro ba motho (Ullsperger, 2010; Frank le Fossella, 2010) hape e ka fetela ka nq'ane ho dorsal striatum, kaha ho na le bopaki ba karohano e ts'oanang ea basebetsi ho "ventral striatum"Mohau et al., 2007; Lobo et al., 2010).

 

Setšoantšo sa 2

Dopamine taolo ea tšusumetso e ntle le e mpe ho dorsal striatum

Le ha morero ona o kaholimo o fana ka setšoantšo se bonolo sa taolo ea boits'oaro ea DA ka litlamorao tsa eona litšoantšong, setšoantšo se felletseng se rarahane haholo. DA e susumetsa boitšoaro bo amanang le moputso ka ho sebetsa likarolong tse ngata tsa boko ho kenyeletsa le cortex ea pele (Hitchcott et al., 2007), rhinal cortex (Liu et al., 2004), hippocampus (Packard le White, 1991; Grecksch le Matties, 1981) le Amygdala (Phillips le al., 2010). Litlamorao tsa DA li kanna tsa fapana haholo lipakeng tsa libaka tsena ka lebaka la phapang ea botsitso ba DA bolulo, batsamaisi ba DA, li-enzyme tsa metabolic, li-autoreceptors, li-receptor, le li-receptor coupling litseleng tse sa tsebeng ho tsebahala (Neve et al., 2004; Bentivoglio le Morelli, 2005; Frank le Fossella, 2010). Ntle le moo, bonyane VV, li-neuron tsa DA li ka ba le thepa e fapaneng ea maqhubu ho latela lipheo tsa bona tsa projeke (Lammel et al., 2008; Margolis et al., 2008), mme ba bang ba na le bokhoni bo tsotehang ba ho fetisa glutamate hammoho le dopamine (Litlatsetso et al., 2008; Chuhma et al., 2009; Hnasko et al., 2010; Tecuapetla et al., 2010; Stuber et al., 2010; Birgner et al., 2010). Ka hona, tekanyo e felletseng ea taolo ea neuron ea DA holim'a ts'ebetso ea neural e se e qala ho senoloa.

Likarabo tse fapaneng tsa dopamine ketsahalong e khopisang

Karabelo ea neuron liketsahalong tse khutsitseng e fana ka teko ea bohlokoa ea mesebetsi ea eona ho taolo e susumetsang (Schultz, 1998; Berridge le Robinson, 1998; Redgrave et al., 1999; Horvitz, 2000; Joseph et al., 2003). Ka litsela tse ngata, re etsa lintho tse thabisang le tse tlohelang batho ba bang ka mekhoa e fapaneng, re bontša hore ba fapaneng le bona boleng bo susumetsang. Re batla meputso mme re ba fa boleng bo nepahetseng, ha re ntse re qoba liketsahalo tse tlosang tse ling ebe re li beha ka boleng bo seng botle. Ka tsela tse ling, re etsa lintho tse monate le tse khathollang ka mekhoa e tšoanang boikaketsi bo susumetsang [FOOTNOTE1]. Liketsahalo tse khotsofatsang le tse ferekanyang li etsa hore batho ba tsepamise maikutlo, ba sebetse hantle, 'me se eketsehe ka matla.

Ke efe ea mesebetsi ee eo li-neurons tsa DA li e tšehetsang? Ke khale ho tsejoa hore liphihlelo tse sithabetsang le tse nyenyefatsang li baka liphetoho tse kholo menahanong ea DA maemong a bongo a theohileng, le hore boitšoaro bo amang liphihlelo tsena bo fetotsoe haholo ke bo-rahistori ba DA, bahanyetsi le liso (Salamone, 1994; Di Chiara, 2002; Pezze le Feldon, 2004; Young le al., 2005). Lithuto tsena li hlahisitse mefuta e mengata e makatsang ea liphetho, leha ho le joalo (Levita et al., 2002; Di Chiara, 2002; Young le al., 2005). Lithuto tse ngata li tsamaisana le li-neurons tsa DA tsa maiketsetso tse susumetsang. Ba tlaleha hore liketsahalo tse nyenyefatsang li eketsa maemo a DA le hore boits'oaro bo bobe bo ts'ehetsoa ke maemo a phahameng a phetiso ea DA (Salamone, 1994; Joseph et al., 2003; Ventura et al., 2007; Barr et al., 2009; Fadok et al., 2009) ho kenyelletsa le ho phatloha hoa ha DAZweifel et al., 2009). Empa lithuto tse ling li tsamaisana hantle le boleng ba tšusumetso ea li-neurons tsa DA. Ba tlaleha hore liketsahalo tse khopisang li fokotsa maemo a DA le hore boits'oaro bo bobe bo ts'ehetsoa ke maemo a tlase a phetiso ea DA (Mark et al., 1991; Shippenberg et al., 1991; Liu et al., 2008; Roitman et al., 2008). Maemong a mangata liphetho tsena tse tsoakaneng li fumanoe lithutong tse le 'ngoe, ho bonts'a hore liphihlelo tse fapaneng le tse ling li baka mekhoa e fapaneng ea ho lokolloa ha DA ka likarolo tse fapaneng tsa boko (Thierry et al., 1976; Besson le Louilot, 1995; Ventura et al., 2001; Jeanblanc et al., 2002; Bassareo et al., 2002; Pascucci et al., 2007), le hore lithethefatsi tse amanang le DA li ka hlahisa motsoako oa litla-morao tsa boits'oaro le boits'oaro bo ts'oanang le bo bakoang ke liphihlelo tse monate le tse nyarosang (Ettenberg, 2004; Wheeler et al., 2008).

Phapang ena ea mekhoa ea tšebetso ea DA e lokollotsoeng le mesebetsi e thata ho lumellana le mohopolo oa hore li-neurons tsa DA li fetisetsa letšoao le tšoanang la tšusumetso ho likarolo tsohle tsa boko. Likarabo tsena tse fapaneng li ka hlalosoa, leha ho le joalo, haeba li-neuron tsa DA ka botsona li entsoe - li entsoe ka lipalo tse ngata tsa neural tse tšehetsang likarolo tse fapaneng tsa ts'ebetso e khelohang. Pono ena e tšehetsoa ke lithuto tsa ho rekota tsa neural ho liphoofolo tse sa tsitsang. Lithuto tsena li bonts'itse hore sepheo se seng le se seng sa ts'usumetso se tsosa likhakanyo tse ling tsa methapo ea DA empa e le thibelo ho li-neuron tse ling tsa DA (Chiodo et al., 1980; Maeda le Mogenson, 1982; Schultz le Romo, 1987; Mantz et al., 1989; Gao et al., 1990; Coizet et al., 2006). Habohlokoa le ho feta, likarabo tsa thabo le tse thibeloang li etsahala ho li-neurons tse netefalitsoeng e le dopaminergic e sebelisang label ea juxtacellular (Brischoux et al., 2009) (Setšoantšo sa 3). Phapang e tšoanang ea likarabo tse khopameng e etsahala nakong ea boitšoaro bo sebetsang. Lihlopheng tse fapaneng tsa li-neuron tsa DA li thabile haholo kapa li thibetsoe ke liketsahalo tse ferekanyang ho kenyelletsa letlalo le sa tsotelleng (Kiyatkin, 1988a; Kiyatkin, 1988b), likhakanyo tsa maikutlo tse bolelang esale pele litšisinyeho tse matla (Guarraci le Kapp, 1999), linonyana tse futhumatsang moea (Matsumoto le Hikosaka, 2009b), le likhakanyo tsa maikutlo tse boletsoeng esale pele ka lifofane tse ferekanyang (Matsumoto le Hikosaka, 2009b; Joshua et al., 2009a). Ntle le moo, ha li-neuron tse peli tsa DA li tlalehiloe ka nako e le ngoe, likarabo tsa tsona tse fapaneng hangata ka kakaretso li na le kamano e fokolang ea liteko.Joshua et al., 2009b), ho fana ka maikutlo a hore likarabo tse akaretsang ha li hokahane ho baahi ba DA ka kakaretso.

 

Setšoantšo sa 3

Likarabo tse fapaneng tsa dopamine neuron li arabela liketsahalong tse fapaneng

Ho utloisisa mesebetsi ea likarabo tsena tse fapaneng tse fapaneng, re hloka ho tseba hore li kopantsoe joang le likarabo tsa moputso ho hlahisa lets'oao le nang le sepheo. Phuputso ea morao-rao e fuputse sehlooho sena mme ea senola hore li-neuron tsa DA li arotsoe ka lihlopha tse ngata tse nang le matšoao a ikhethang a susumetsang (Matsumoto le Hikosaka, 2009b). Baahi ba bang ba thabisoa ke liketsahalo tse putsa le tse thibeloang ke liketsahalo tse nyarosang, joalo ka ha eka li na le likhakanyo boleng bo susumetsang (Setšoantšo sa 4A). Baahi ba bobeli ba thabisoa ke liketsahalo tse monate le tse nyarosang ka mekhoa e tšoanang, joalokaha eka ke ho encodisa boikaketsi bo susumetsang (Setšoantšo sa 4B). Ka bobeli ba batho bana li-neuron tse ngata li ikemiselitse ho putsa le ho hakanya lintho tse boletsoeng esale pele: li arabela ha liketsahalo tse thabisang li le molemo ho feta tse boletsoeng esale pele le ha liketsahalo tse nyarosang li hlasimolla ho feta tse boletsoeng esale pele (Matsumoto le Hikosaka, 2009b). Sena se bonts'a hore likarabo tsa bona tse khopameng li hlile li bakoa ke boiketsetso mabapi le liketsahalo tse nyarosang, ho hlahisa monyetla oa hore li ka bakoa ke lintho tse sa ikhethileng tse kang tlatsetso ea maikutlo a mantle kapa mekhatlo e ikopantseng le moputso (Schultz, 2010). Baahi bana ba babeli ha ba tšoane, ka mokhoa o hlakileng oa khoutu ea bona ea ho nahana esale pele. Bohlokoa ba ho beha kh'outu ea li-neurons tsa DA li kenyelletsa letšoao le nepahetseng la phoso ea ho bolela esale pele, ho kenyelletsa le thibelo e matla ka ho siea ha meputso le boikhathollo bo bonolo ka ho siea liketsahalo tse khelohang (Setšoantšo sa 4A, hantle). Ka lehlakoreng le leng, lisishene tse susumetsang tsa ho ngola li-neurons tsa DA li arabela ha liketsahalo tse ikhethang li le teng empa eseng ha li le sieo (Setšoantšo sa 4B, ka ho le letona), e lumellanang le maikutlo a khopolo ea ho tsosa (Lang le Davis, 2006) [FOOTNOTE2]. Bopaki ba bongata bona ba li-neuron tsa DA bo bonoe le ha ts'ebetso ea neural e hlahlojoa ka mokhoa o ikhethileng. Kahoo, lithuto tse shebileng likarolo tse fapaneng tsa sistimi ea DA li fumaneng matšoao a DA a phasic a kenyelletsa liketsahalo tse nyarosang ka thibelo (Roitman et al., 2008), e ts'oanang le ho kh'outa ka boleng ba susumetsang, kapa ka thomello (Joshua et al., 2008; Anstrom et al., 2009), e ts'oanang le ho kh'outa tsa sesapo se susumetsang.

 

Setšoantšo sa 4

Disopamine dopamine neuron e bopa ea encoding boleng bo susumetsang le masapo

Liphumano tsena tsa morao-rao li ka utloahala li hanana le tlaleho ea pejana ea hore li-neurons tsa DA li khetha ka mokhoa o ikhethileng ho lefa moputso ho e-na le ho khalemela ba bang (Mirenowicz le Schultz, 1996). Ha e hlahlojoa ka hloko, leha ho le joalo, le thuto eo e tsamaellana ka botlalo le boleng ba DA le ho kh'outa ha salience. Thutong eo ea meputso ho ile ha lebisa liphellong tsa moputso ka monyetla o moholo (> 90%) ha mekhoa e fapakaneng e lebisitse liphethong tse khelosang tse nang le menyetla e tlase (<10%). Kahoo boleng le ho bokella salions tsa DA li ne li ke ke tsa arabela hanyane ka hofetisisa, ka nepo li kentse boemo ba bona bo tlase ba ho se tsotelle.

Karolo e sebetsang ea boleng bo susumetsang le matšoao a boits'oaro

Ha ho kopanngoa hammoho, liphumano tse kaholimo li bonts'a hore li-neuron tsa DA li arotsoe ka likarolo tse ngata tse loketseng likarolo tse ikhethileng taolong ea susumetsang. Bohlokoa ba "coding" ba li-neurons tsa DA bo lumellana hantle le likhopolo tsa hona joale tsa dopamine neurons le ts'ebetso ea moputso (Schultz et al., 1997; Berridge le Robinson, 1998; Bohlale, 2004). Li-neurons tsena li fupere pontšo e felletseng ea phoso ea ho bolela esale pele, 'me li behe liketsahalo tse monate le tse pharalletseng. Ka hona, li-neurons tsena li fana ka pontšo e loketseng e thusang bakeng sa ho batla, ho lekola, le ho ithuta boleng (Setšoantšo sa 5). Haeba ts'usumetso e etsa hore boleng ba li-neurons tsa DA li nyakalle, re lokela ho e atamela, re e abele boleng bo phahameng, 'me re ithute liketso ho e batla hape nakong e tlang. Haeba ts'usumetso e etsa hore boleng ba li-neurons tsa DA bo thibeloe re lokela ho bo qoba, re bo behe boleng bo tlase, 'me re ithute liketso ho bo qoba hape nakong e tlang.

 

Setšoantšo sa 5

Mesebetsi ea Hypothesized ea bohlokoa bo susumetsang, sesose, le matšoao a lemosang

Ka lehlakoreng le leng, li-neurons tsa DA tse sebetsang li lumellana hantle le likhopolo tsa dopamine neurons le ho sebetsa ha liketsahalo tse ikhethang (Redgrave et al., 1999; Horvitz, 2000; Joseph et al., 2003; Kapur, 2003). Li-neuron tsena li khahlisoa ke liketsahalo tse monate le tse tlatsetsang ho tsona ebile li na le likarabo tse fokolang liketsahalong tse sa jeleng paate, li fana ka leseli le loketseng bakeng sa ho tsamaisoa ha neural ho ithuta ho tseba, ho bolela esale pele le ho arabela maemong a bohlokoa haholo. Mona re tla hlahloba lits'ebetso tse tharo joalo tsa boko (Setšoantšo sa 5). Taba ea pele, li-circuits tsa neural bakeng sa ho shebella le ho lebisa tlhokomelo li khahloa ho fumana tlhaiso-leseling ka mefuta eohle ea liketsahalo, e monate le e khahlisang. Mohlala, litheko tse khahlisang le tse khahlisang li hohela litšusumetso tse sebetsang hamolemo ho feta mekhoa e sa nke lehlakore (Lang le Davis, 2006; Matsumoto le Hikosaka, 2009b; Austin le Duka, 2010). Taba ea bobeli, maemo a putsoang le a khefutsang a kenyelletsa mekhoa ea methapo ea kutlo bakeng sa taolo ea kutloisiso le khetho ea liketso - re hloka ho kenya letsoho mohopolong oa ho boloka tlhaiso-leseling ka kelellong, tharollo ea likhohlano ho nka qeto mabapi le ketso, le mohopolo oa nako e telele ho hopola sephetho se hlahang (Bradley et al., 1992; Botvinick et al., 2001; Savine et al., 2010). Taba ea boraro, maemo a thabisang le a khopisang a hloka keketseho ea kakaretso ea ho matlafatsa liketso le ho etsa bonnete ba hore li etsoa hantle. Ka sebele, li-neurons tsa DA li bohlokoa haholo ho susumetseng boiteko ba ho fihlela lipheo tsa boleng bo holimo le ho fetolela tsebo ea litlhoko tsa mosebetsi hore o sebetse ka mokhoa o tšepahalang oa makoloi (Berridge le Robinson, 1998; Mazzoni et al., 2007; Niv et al., 2007; Salamone et al., 2007).

Hlaola lintho tse u khothatsang ka ho lemosa likeleli tsa maikutlo

Ntle le litlatsetso tsa bona tsa boleng le tšusumetso ea maqhubu, bongata ba li-neuron tsa DA bo boetse bo na le likarabo tse fapaneng mefuta e fapaneng ea liketsahalo tsa maikutlo tse sa amaneng ka kotloloho le liphihlelo tse monate kapa tse nyarosang. Likarabo tsena li filoe maikutlo ho latela lintlha tse 'maloa tsa neural le kelello, ho kenyelletsa ho kenya maikutlo ka kotlolloho, ho makala, ho hlaka, ho tsosa maikutlo, tlhokomelo, ho se sebe, ho akaretsa le maemo a pseudo (Schultz, 1998; Redgrave et al., 1999; Horvitz, 2000; Lisman le Mohau, 2005; Redgrave le Gurney, 2006; Joshua et al., 2009a; Schultz, 2010).

Mona re tla leka ho kopanya mehopolo ena le ho ikarabella bakeng sa likarabo tsena tsa DA ho latela lets'oao le le leng le ka tlasa, a tlhokomeliso (Setšoantšo sa 5). Lentsoe 'tlhokomeliso' le sebelisitsoe ke Schultz (Schultz, 1998) joalo ka polelo e akaretsang bakeng sa liketsahalo tse hohelang tlhokomelo. Mona re tla e sebelisa ka mokhoa o ikhethang. Ka ketsahalo e hlokomelisang, re bolela senepe se sa lebelloang sa maikutlo se lebisang tlhokomelo ho latela tlhahlobo e potlakileng ea bohlokoa ba eona, re sebelisa likarolo tse bonolo tse kang sebaka sa eona, boholo le mokhoa oa maikutlo. Liketsahalo tse joalo tse fanang ka temoso hangata li etsa hore batho ba iphetetse hang-hang ho bona le ho fumana moelelo o nepahetseng. Ka tsela ena mats'oao a ho hlokomelisa DA hangata a hlaha libakeng tse khutšoane, a ipapisitse le likarolo tse sa utloahaleng tsa khothatso, 'me li tsamaisana hantle le karabelo tse potlakileng joalo ka karabelo e potlakileng (Schultz le Romo, 1990; Joshua et al., 2009a; Schultz, 2010). Sena se fapane le matšoao a mang a susumetsang a li-neuron tsa DA tse atisang ho hlaha ka nako e telele, li nahanne ka tšusumetso e nepahetseng, 'me li amana hantle le liketso tse nkoang e le liqeto tse lokelang ho atameloa kapa ho qojoa (Schultz le Romo, 1990; Joshua et al., 2009a; Schultz, 2010).

Likarabo tse fanang ka temoso ea DA li ka bakoa ke liketsahalo tse makatsang tse kang khanya e sa lebelloang le likhakanyo tsa lenaneong, tse hlahisang thabo e kholo ho 60-90% ea li-neurons tsa DA ho pholletsa le SNc le VTA (Strecker le Jacobs, 1985; Horvitz et al., 1997; Horvitz, 2000) (Setšoantšo sa 6A). Likarabo tsena tse lemosang li bonahala li bonts'a tekanyo eo mohopolo o khothalletsang ho makatsa hape o hohelang tlhokomelo; lia fokotseha haeba ts'usumetso e etsahala ka nako e boletsoeng esale pele, haeba tlhokomelo e kenella kae-kae, kapa nakong ea boroko (Schultz, 1998; Takikawa et al., 2004; Strecker le Jacobs, 1985; Steinfels et al., 1983). Mohlala, molumo oa ho tobetsa o sa lebelloang o tsosa lerata le tummeng la DA ha katse e le boemong ba boithati bo khutsitseng, empa ha e na thuso ha katse e etsa lintho tse batlang tlhokomelo joalo ka ho tsoma rat, ho fepa, ho itlhophisa le ho ts'oaroa. ka sebali, joalo-joalo (Strecker le Jacobs, 1985) (Setšoantšo sa 6A). Ka mokhoa o ts'oanang, likarabo tsa DA tse phatlohileng li ts'oaroa ke liketsahalo tse fokolisang 'mele empa li lemosa ka lebaka la bohlanya ba bona (Ljungberg et al., 1992; Schultz, 1998). Likarabo tsena li lula li le teng ha thuto ea nalane e qala ho tloaela, ka ho tsamaisana le mokhoa oa ho feto-fetoha oa maikutlo (Setšoantšo sa 6B). Tumellanong le liphumano tsena, liketsahalo tse makatsang le tse ncha li etsa hore DA e lokolloe meahong e theohelang.Lisman le Mohau, 2005) le ho kenya tšebetsong li-circuits tsa boko tse amanang le DA ka tsela e bonoang ho lokisa moputso (Zink le al., 2003; Davidson et al., 2004; Duzel et al., 2010).

 

Setšoantšo sa 6

Dopamine likarabo tse thabisang tsa neuron mabapi le liketsahalo tse lemosang

Likarabo tse lemosang tsa DA li boetse li ts'oaroa ke mekhoa e sa lebelloang ea maikutlo e nang le monyetla oa ho fana ka tlhaiso-leseling e ncha ka liketsahalo tse itlhommeng pele. Joalokaha ho lebelletsoe bakeng sa lets'oao le lemosang la nako e khuts'oanyane, likarabo tsena ha li khethoe: li susumetsoa ke ts'usumetso efe kapa efe e ikemiselitseng feela e tšoana khoele e tšehetsang ka mokhoa o susumetsang, le haeba ho tšoana ho nyane haholo (ketsahalo e bitsoang kakaretso) (Schultz, 1998). Ka lebaka leo, li-neurons tsa DA hangata li arabela ho ts'usumetso e nang le motsoako oa matšoao a mabeli: lets'oao le lemosang le potlakisang ho tiisa taba ea hore khothalletso ke mohlomong ea bohlokoa, 'me lets'oao la bobeli le a kenyelelitse sebele moelelo o khotsofatsang kapa o khopisang (Schultz le Romo, 1990; Waelti et al., 2001; Tobler et al., 2003; Letsatsi le al., 2007; Kobayashi le Schultz, 2008; Fiorillo et al., 2008; Nomoto et al., 2010) (Bona (Kale le Dayan, 2002; Joshua et al., 2009a; Schultz, 2010) bakeng sa tlhahlobo). Mohlala o ka bonoa sethaleng sa li-neurons tsa ts'usumetso ea ts'usumetso ea DA tse bontšitsoeng ho Setšoantšo sa 6C (Bromberg-Martin et al., 2010a). Li-neuron tsena li ne li khahliloe ke moputso le menyenyetsi e khahlisang, empa hape li ne li thabisitsoe ke polelo e sa jeleng paate. Cue ea ho se nke lehlakore e ne e so ka e raloa ka sephetho se susumetsang, empa e ne e ts'oana le moputso le likotsi tse nyarosang.

Likarabo tsena tse fanang ka temoso li bonahala li matahane haholo le bokhoni ba leseli ba ho ferekanya maikutlo a ho e hlahloba le ho e utloisisa. Sena se ka bonoa maemong a mararo a hlokomelehang. Taba ea pele, likarabo tse lemosang li ba teng feela bakeng sa litakatso tsa maikutlo tse tlamehang ho hlahlojoa ho bona moelelo oa tsona, eseng bakeng sa liketsahalo tse thabisang kapa tse nyarosang tse kang ho tsamaisoa ha lero kapa linonyana (airpuffs).Schultz, 2010). Taba ea bobeli, likarabo tse lemosang li etsahala feela ha cue e le bohlokoa ka mokhoa o fapaneng mme e khona ho tsosa maikutlo, eseng ha cue e sa amane le mosebetsi o teng mme e sa sitise ho sisinya maikutlo (Schultz le Romo, 1990). Taba ea boraro, likarabo tse fanang ka temoso li ntlafatsoa maemong ao mekhoa e ka etsang hore batho ba lebise tlhokomelo e sa lebelloang - ha e hlaha ka nako e sa lebelloang kapa e le bohareng ba mahlo (Bromberg-Martin et al., 2010a). Kahoo ha mekhoa e susumetsang e hlahisoa ka nako e sa lebelloang e tsosa karabelo e potlakileng le karabelo e akaretsang ea temoso ea DA - ho khahlisoa ke mekhoa eohle ho kenyelletsa lintlha tse sa nke lehlakore (Setšoantšo sa 6C, e sootho). Empa haeba nako ea bona e etsoa esale pele - mohlala, ka ho bonts'a lihlooho tse nang le "teko ea ho qala nyeoe" e hlahisitsoeng motsotsoana pele likhokahano li hlaha - likhetsi ha li sa hlahisa karabelo e lemosang (Setšoantšo sa 6D, bohlooho). Sebakeng seo, karabelo e lemosang e fetohela temaneng ea ho qala nyeoe - ketsahalo ea pele ea nyeoe e nang le nako e sa lebelloang le e tsosang karabelo e lebisang (3)Setšoantšo sa 6D, e sootho).

Mochini o hlahisang li-neuron tsa tlhokomeliso ea neuron ke ofe? Khopolo-taba e 'ngoe ke hore likarabo tse lemosang ke feela lipontšo tse tloaelehileng tsa moputso tse boletsoeng libakeng tse khutšoane, tse kenyelletsang boleng ba moputso bo lebelletsoeng pele ho khothatsoa pele ho khethollo e felletseng (Kale le Dayan, 2002). Bopaki ba haufinyane tjena, leha ho le joalo, bo fana ka maikutlo a hore matšoao a lemosang a ka hlahisoa ke mochine o ikhethileng ho tsoa lipontšong tse tloaelehileng tsa moputso oa DA (Satoh et al., 2003; Bayer le Glimcher, 2005; Bromberg-Martin et al., 2010a; Bromberg-Martin et al., 2010c; Nomoto et al., 2010). Ho makatsang ka ho fetisisa, karabelo e lemosang ho qalo ea nyeoe ha e felle feela mesebetsing e khotsofatsang; e ka ba le matla a lekanang nakong ea mosebetsi o khelohang empa ho sa tlisoe meputso (Setšoantšo sa 6C, D, tlase, "mosebetsi oa ho phethola"). Sena se etsahala leha e le lipontšo tse tloaelehileng tsa moputso oa DA ka li-neuron tse tšoanang li supa hantle hore mosebetsi o moputso o na le boleng bo holimo bo lebelletsoeng ho feta mosebetsi o phephetsang (Bromberg-Martin et al., 2010a). Matšoao ana a lemosang ha se mofuta oa boleng ba ho ngola likhakanyo kapa mofuta oa mofuta oa ho etsa litšitiso, hobane li hlahella boholo ba bohlokoa bo susumetsang le ho seta li-neurons tsa DA (Bromberg-Martin et al., 2010a). Ho ikarola ka bobeli ho ka bonoa ka tsela eo li-neurons tsa DA li bolelang esale pele meputso ea nako e tlang ho latela mohopolo oa liphetho tsa nakong e fetileng (Satoh et al., 2003; Bayer le Glimcher, 2005). Le ha matšoao a moputso a tloaelehileng a DA a laoloa ke mohopolo oa nako e telele o hopoloang moputso o nepahetseng, likarabo tse mabapi le teko ea ho qala nyeoe li laoloa ke setšoantšo se ka thoko sa memori se tšoanang le se bonoang mokhoeng o haufi oa ho etsa joalo (Bromberg-Martin et al., 2010c). Ho arohana ha boraro ho ka bonoa ka tsela eo lipontšo tsena li fetisoang ka bongata ho li-neuron tsa DA. Le ha matšoao a tloaelehileng a moputso a DA a le matla ho SNc e kenelletseng, tsebiso e arabisang karabo ea nyeoe e qalang (le ho mekhoa e meng e sa lebelloang e sa lebelloang) e phatlalatsoa ho SNc (Nomoto et al., 2010).

Ho fapana le karohano ena ho tsoa lipontšong tse tloaelehileng tsa moputso, lipontšo tse lemosang tsa DA li hokahanngoa le lebelo la ho shebella le ho arabela likarabo ketsahalong e lemosang (Satoh et al., 2003; Bromberg-Martin et al., 2010a; Bromberg-Martin et al., 2010c). Sena se fana ka maikutlo a hore matšoao a lemosang a hlahisoa ke ts'ebetso ea neural e susumetsang karabelo e potlakileng ea ho etsa lipatlisiso tsa liketsahalo tse ka bang bohlokoa. Nakong ea hona joale, ka bomalimabe, ha ho tsejoe letho ka hore na ketsahalo ena e nka e le ea bohlokoa hakae. Mohlala, na ho lemosa likarabo ka mokhoa o ts'oanang ho liketsahalo tse thabisang le tse fetohelang? Likarabo tse fanang ka temoso li tsejoa li hlaha bakeng sa sepheo se ts'oanang le meputso e tšoanang kapa moputso o tšoanang (mohlala ka ho arolelana maikutlo a tšoanang). Empa ha ho tsejoe hore na likarabelo tse lemosang li ba teng ka lebaka la ts'usumetso e batlang e tšoana le lintlha tse khelohang.

Mosebetsi o sebetsang oa ho bonts'a matšoao a dopamine

Joalokaha re bone, mats'oao a lemosang a kanna a hlahisoa ke mochine o ikhethileng ho tloha boleng bo susumetsang le matšoao a sephara. Leha ho le joalo, mats'oao a tlhokomeliso a romelloa ho boleng ba bohlokoa bo susumetsang le li-salience tsa ho kenya li-neurons tsa DA, ka hona li ka khona ho laola ts'ebetso ea ts'ebetso ea boko le boitšoaro ka mokhoa o ts'oanang le matšoao a boleng le ho se sebe (Setšoantšo sa 5).

Matšoao a hlokomelisoang a rometsoeng ho li-activist tsa ho taka tsa boikemisetso ba li-neurons tsa DA li tla tšehetsa ho lebisa tlhokomelo ho ts'usumetso e lemosang, ho kenella ha lisebelisoa tsa tlhokomeliso ho fumana moelelo le ho etsa qeto ka moralo oa ts'ebetso, le ho eketsa maemo a ts'usumetso ho kenya ts'ebetsong moralo ona ka nepo (Setšoantšo sa 5). Litlamorao tsena li ka hlaha ka litlamorao tsa ts'ebetsong ea neural kapa ka ho tiisa liketso tse lebisitseng ho boneng ketsahalo e lemosang. Karolo ena e sebetsang e tsamaisana hantle le kamano lipakeng tsa likarabo tse lemosang tsa DA le karabelo e potlakileng ea boits'oaro ho khothaletso e lemosang, hape ka maikutlo a hore likarabo tsa nakoana tsa tatellano ea DA ea neuron li kentse letsoho ho lebiseng tlhokomelo, ho tsosa mofuthu, ntlafatso ea ts'ebetso ea kelello, le karabelo ea hang-hang ea boitšoaro (Redgrave et al., 1999; Horvitz, 2000; Joseph et al., 2003; Lisman le Mohau, 2005; Redgrave le Gurney, 2006; Joshua et al., 2009a).

Ho ba teng ha mats'oao a ho lemosa ka bohlokoa ba li-neurons tsa DA ho thata ho ho hlalosa. Li-neurons tsena li fetisa matšoao a boleng bo susumetsang a loketseng ho batla, ho lekola sephetho le ho ithuta boleng; leha ho le joalo ba ka boela ba hlasimoloha ka ho lemosa liketsahalo tse kang melumo ea ho tobetsa e sa lebelloang le ho qala ha liteko tse matla. Ho latela tsela ea rona e hloahloa (Setšoantšo sa 5), sena se ka etsa hore liketsahalo tse hlokomelisang li fuoe boleng bo nepahetseng mme li batloe ka mokhoa o ts'oanang le moputso! Le ha o makatsa ha o habanya feela, ho na le lebaka la ho belaela hore liketsahalo tse lemosang li ka nkuoa e le lipheo tse ntle. Matšoao a lemosang a fana ka temoso ea pele ea hore ketsahalo e bohlokoa e haufi e tla etsahala, ka hona e fana ka monyetla oa pele oa ho nka khato ho laola ketsahalo eo. Haeba mekhoa ea tlhokomeliso e teng, liketsahalo tse ikhethileng li ka bonoa, tsa boleloa esale pele tsa ba tsa lokisetsoa esale pele; haeba tlhokomeliso e le sieo, liketsahalo tse ikakasang ka linako tsohle li etsahala ka mokhoa o sa lebelloang. Ho joalo, hangata batho le liphoofolo ba bontša ho rata libaka tse monate, tse khutsitseng esita le tse senang maikutlo tse senang maikutlo tse sa amaneng le maikutlo li ka boleloa esale pele (Badia et al., 1979; Herry et al., 2007; Daly, 1992; Chew le Ho, 1994) le li-neurons tse ngata tsa DA li bontša mokhoa oa ho khetha ho bona tlhaiso-leseling e boletsoeng esale pele (Bromberg-Martin le Hikosaka, 2009). Maemo a lemosang a DA a ka ts'ehetsa likhetho tsena ka ho beha boleng bo nepahetseng maemong ao ho ka lebelloang lintlha tsa bohlokoa tsa maikutlo pele ho nako.

Libaka tsa anatomical tsa boleng le li-neurons tsa coding neurons

Phuputso ea morao-rao e bontsitse libaka tsa moputso oa DA le matšoao a pharalatsang bohareng ba 'mele bo kenyelletsang karolo ea SNc le karolo ea morao-rao ea VTA (Matsumoto le Hikosaka, 2009b). Bohlokoa ba ho susumetsa le matšoao a boikaketsi ba ts'usumetso li fetisitsoe sebakeng sena ka gradient ea anatomical. Matšoao a boleng bo susumetsang a fumanoe a fumaneha hangata ho li-neuron ho SNc ea ventromedial le VTA ea morao-rao, ha matšoao a susumetsang a ts'usumetso a fumaneha hangata ho li-neurons ho dorsolateral SNc (Setšoantšo sa 7B). Sena se lumellana le litlaleho tsa hore moputso oa boleng ba moputso oa DA o matla ho SNc ea ventromed (Nomoto et al., 2010) ha maikutlo a ferekanyang hangata a ba matla ho feta hamorao (Mirenowicz le Schultz, 1996). Boithuto bo bong bo ithutile mofuta oa mofuta oa bo-maleshoane bo atileng haholoanyane. Lithuto tsena li fumane motsoako oa likarabo tse nyakallisang le tse thibelang maikutlo ho se na phapang e kholo libakeng tsa bona, leha ho le joalo ka mokhoa oa thabo e nyarosang ho ba teng ka mokhoa o bulehileng haholoanyane (Guarraci le Kapp, 1999; Brischoux et al., 2009) (Setšoantšo sa 7C).

Mafelo a matšoao a boleng bo susumetsang

Ho ea ka tšibollo ea rona, boleng ba "coding" ba li-neurons tsa DA bo lokela ho kenella libakeng tsa boko tse kenyellelitsoeng ho liketso tsa ho qoba, ho hlahloba sephetho le ho ithuta boleng (Setšoantšo sa 5). Ka sebele, projeke ea "NN" ea VNc le VTA e kenella ho "cortex" e tlang pele ho nako.Williams le Goldman-Rakic, 1998) ho kenyelletsa orbitofrontal cortex (OFC) (Porrino le Goldman-Rakic, 1982) (Setšoantšo sa 7A). OFC e 'nile ea kenyelletsoa khafetsa ho boleng ba ho kenya likhouto lithutong tse sebetsang tsa ho nahana (Anderson et al., 2003; Tse nyenyane le al., 2003; Jensen et al., 2007; Litt et al., 2010le lirekoto tsa neuron e le 'ngoe (Morrison le Salzman, 2009; Roesch le Olson, 2004). OFC e nahanoa ho lekola likhetho tsa khetho (Padoa-Schioppa, 2007; Kable le Glimcher, 2009), litebello tsa sephetho sa encode (Schoenbaum et al., 2009), le ho ntlafatsa litebello tsena nakong ea ho ithuta (Walton et al., 2010). Ntle le moo, OFC e nka karolo ho ithuteng liphosong tse mpe tse boletsoeng esale pele tsa moputso (Takahashi et al., 2009) tse matla ka ho coding DA li-neurons (Setšoantšo sa 4).

Ntle le moo, likarolo tsa medial tsa projeke ea dopaminergic midbrain ho "ventral striatum" ho kenyelletsa le nucleus accumbens shell (NAc shell) (Haber et al., 2000) (Setšoantšo sa 7A). Phuputso ea morao-rao e bonts'itse hore khetla ea NAc e fumana lipontšo tsa "phasic" tsa DA tse akaretsang boleng bo susumetsang ba liphetho tsa tatso (Roitman et al., 2008). Matšoao ana a kanna a baka ho ithuta ha boleng hobane ho kenngoa ka kotlolloha ha lithethefatsi tsa DA ka khetla ea NAc ho matlafatsa haholo (Kemoto, 2010) ha kalafo tse fokotsang tlatsetso ea DA khetla li ka tsosa (Liu et al., 2008). Lebaka le leng ke hore lithuto tsa NAc tsa DA li lokollotsoe nako ea nako e telele (metsotso) li hlahisitse litholoana tse fapaneng, tse ling li tsamaisana le khoutu ea boleng le tse ling tse nang le mokhoa oa ho seta (mohlala (Bassareo et al., 2002; Ventura et al., 2007)). Sena se fana ka maikutlo a hore matšoao a boleng a ka fokotsoa feela libakeng tse ikhethang ka har'a khetla ea NAc. Haholo, libaka tse fapaneng tsa khetla ea NAc li khetheloa ho laola takatso ea lijo le boiphetetso (Reynolds le Berridge, 2002), tseo ka bobeli li hlokang ho kenya maikutlo ho tsoa ho li-neuron tsa DA (Faure et al., 2008).

Kamora nako, li-neurons tsa DA ho theosa le boholo ba SNc li romella lipalo tse boima ho dorsal striatum (Haber et al., 2000), ho fana ka maikutlo a hore dorsal striatum e ka fumana boleng ba susumetso le maseli a li-signature tsa DA ((Setšoantšo sa 7A). Bohlokoa ba ho kenya kh'outu ea li-neurons tsa DA bo ka fana ka leseli le nepahetseng bakeng sa potoloho e amanang le ho ithuta ka boleng, joalo ka ho ithuta ka mekhoa ea ho arabela ka khothatso (Faure et al., 2005; Yin le Knowlton, 2006; Balleine le O'Doherty, 2010). Ha li-neuron tsena tsa DA li phatloha, ba ne ba kenella tseleng e tobileng ea ho ithuta ho fumana moputso; ha ba khefutsa, ba ne ba nka tsela e sa tobang ho ithuta ho qoba sephetho se setle (Setšoantšo sa 2). Ho joalo, ho na le bopaki ba morao-rao ba hore litsela tse tiileng li latela karohano ena ea mosebetsi bakeng sa moputso le ts'ebetso e khelohileng (Hikida et al., 2010). Leha ho le joalo, ha e ntse e sa tsejoe hore na li-neurons tse tseleng tsena li arabela joang liketsahalong tse monate le tse fapaneng nakong ea boitšoaro. Bonyane ho dorsal striatum ka kakaretso, seteishene sa methapo se arabela liketsahalong tse itseng tse thabisang le tse nyarosang ka mekhoa e ikhethileng (Ravel et al., 2003; Yamada et al., 2004, 2007; Joshua et al., 2008).

Lits'oaetso tsa matšoao a ts'usumetso ea matla a ho susumetsa

Ho ea ka tšibollo ea rona, li-neurons tsa DA tse sebetsang ka sepheo sa ts'ebetso li lokela ho kenella libakeng tsa boko tse kenyelletsang ho tsepamisang kelello, tšebetsong kelello le ho susumetsa ka kakaretso (Setšoantšo sa 5). Ka sebele, li-neurons tsa DA tse tšebetsong methapo ea methapo e tsamaisang methapo ea maoto li romella likhakanyo ho lisele tsa bongoaneng le tsa morao-rao (Williams le Goldman-Rakic, 1998) (Setšoantšo sa 7A), sebaka se kentsoeng ts'ebetsong ea ho lemoha joalo ka ho batla ka hloko, mohopolo oa ho sebetsa, taolo ea kelello, le ho etsa liqeto lipakeng tsa liphetho tse susumetsang (Williams le Castner, 2006; Lee le Seo, 2007; Bohlale, 2008; Kable le Glimcher, 2009; Wallis le Kennerley, 2010). Mesebetsi ea kelello ea mantlha ea bokhachane e laoloang ka thata ke maemo a DA (Robbins le Arnsten, 2009) mme ba reretsoe ho latela ts'ebetso ea phasic DA neuron (Cohen et al., 2002; Lapish et al., 2007). Haholo-holo, seteishene sa methapo ea kutlo se tlang pele ho marang-rang se arabela linthong tse bonoang tse atlehang le tse thibelang maikutlo, 'me boholo ba bona bo arabela ka lehlakoreng le ts'oanang le ho fana ka maikutlo a ho se sebetse hantle (Kobayashi et al., 2006). Ntle le moo, tšebetso ea methapo ena ea kutlo e tsamaisana le katleho ea boits'oaro mosebetsing oa ho etsa mesebetsi ea memori e sebetsang (Kobayashi et al., 2006). Leha mokhoa ona oa dorsolateral DA → dorsolateral frontal cortex way e bonahala e khethehile ho li-primates (Williams le Goldman-Rakic, 1998), tsela e ts'oanang e sebetsang e kanna ea ba teng ho mefuta e meng. Haholo-holo, mesebetsi e mengata ea boiqapelo ea prortal dorsolateral prefrontal cortex e etsoa ke rodent medial prefrontal cortex (Uylings et al., 2003), mme ho na le bopaki ba hore sebaka sena se fumana matšoao a susumetsang a DA le ho laola boits'oaro bo amanang le salience (Mantz et al., 1989; Di Chiara, 2002; Joseph et al., 2003; Ventura et al., 2007; Ventura et al., 2008).

Ha ho fanoa ka bopaki ba hore VTA e na le mahlaseli a mabeli a boleng le li-coding neurons le hore boleng ba matšoao a ho beha likh'outu li romelloa mokokotlong oa NAc, matšoao a salience a ka romelloa mokokotlong oa NAc (Setšoantšo sa 7A). Ehlile, motheo oa NAc (empa eseng khetla) e bohlokoa bakeng sa ho susumetsa tšusumetso ea ho hlola litšenyehelo tsa karabelo joalo ka boiteko ba 'mele; ts'ebetso ea mesebetsi e feto-fetohang e hlokang ho feto-fetoha ha kelello; le bakeng sa ho thusa meputso ea meputso ho khothaletsa khothatso e akaretsang (Ghods-Sharifi le Floresco, 2010; Floresco et al., 2006; Hall et al., 2001; Mok'hadinale, 2006). E lumellana le ho hlophisoa ha bohlanya ba ho susumetsoa ke tšusumetso, mantlha ea NAc e amohela ho phatloha ho hoholo ha DA nakong ea liphihlelo tse peli tse monate (Letsatsi le al., 2007) le liphihlelo tse fetolang (Anstrom et al., 2009).

Qetellong, joalo ka ha ho hlalositsoe kaholimo, mekhoa e meng ea "li-neurons" ea DA e ka ipabola ho dorsal striatum (Setšoantšo sa 7A). Leha libaka tse ling tsa dorsal striatum li kentse letsoho mesebetsing e amanang le boleng ba ts'ebetso ea thuto, dorsal striatum e boetse e kenella mesebetsing e lokelang ho etsoa bakeng sa liketsahalo tsohle tse ikhethang, tse kang ho tsepamisa kelello, ho tsepamisa mohopolo, ho etsa memori e sebetsang, le ts'usumetso e akaretsang (Hikosaka et al., 2000; Klingberg, 2010; Palmiter, 2008). Ka 'nete, sephutheloana sa methapo ea kutlo e arabelang ka ho fetelletseng liketsahalong tse monate le tse fetohang ho feta liketsahalong tse sa nke lehlakore.Ravel et al., 1999; Blazquez et al., 2002; Yamada et al., 2004, 2007), leha karolo ea bona ea boqhekanyetsi boitsholong bo susumetsang e e-so tsejoe.

Mehloli ea matšoao a boleng bo susumetsang

Tlhahlobo ea morao-rao ea lipatlisiso e bontša hore li-neurons tsa DA li fumana matšoao a bohlokoa a susumetsang ho tloha mookotong o monyane ho epithalamus, habenula (LHb) ea morao (Hikosaka, 2010) (Setšoantšo sa 8). LHb e fana ka taolo e matla ea taolo ho li-neuron tsa DA: Ts'usumetso ea LHb e thibela li-neurons tsa DA maemong a makgutshwane (Christoph et al., 1986) hape e ka laola ho ithuta ka tsela e fapaneng le ea ho khothaletsa VTA (Shumake et al., 2010). E lumellana le letšoao la taolo e mpe, li-neurons tse ngata tsa LHb li na le likarabo tsa phasic tse kentsoeng ka seiponeng tsa li-neuron tsa DA: LIH neurons ke e thibetsoe ka liphoso tse boletsoeng esale pele tsa moputso mme thabile ka liphoso tse boletsoeng esale pele tsa moputso (Matsumoto le Hikosaka, 2007, 2009a; Bromberg-Martin et al., 2010a; Bromberg-Martin et al., 2010c). Maemong a 'maloa matšoao ana a hlaha maemong a makhuts'oane ho LHb, a lumellana le phetisetso ea LHb → DA (Matsumoto le Hikosaka, 2007; Bromberg-Martin et al., 2010a).

 

Setšoantšo sa 8

Mehloli ea Hypothesised ea bohlokoa bo susumetsang, sesose, le matšoao a lemosang

LHb e khona ho laola li-neuron tsa DA ho pholletsa le bohareng ba mpa, empa lintlha tse 'maloa tsa bopaki li fana ka maikutlo a hore e fana ka taolo e khethollang matla a khothalletsang boleng ba li-neurons tsa DA. Taba ea mantlha, li-neurons tsa LHb li kenyelletsa boleng bo susumetsang ka mokhoa o lekanyang boleng ba li-neurons tsa DA - li behile liphoso tse mpe le tse mpe tsa ho bolela moputso ebe li arabela ka lehlakoreng le fapaneng liketsahalong tse lehlohonolo le tse phetetsang (Matsumoto le Hikosaka, 2009a; Bromberg-Martin et al., 2010a). Ntlha ea bobeli, ts'usumetso ea LHb e na le litlamorao tse matla ho li-neuron tsa DA tseo thepa ea eona e lumellanang le boleng ba ho kenyelletsa lintlha, ho kenyelletsa ho thibeloa ke likhakanyo tsa moputso le sebaka sa anatomical ho SNc ea ventromedial (Matsumoto le Hikosaka, 2007, 2009b). Taba ea boraro, ho ruruha ha LHb ho sitisa likarabo tsa thibelo ea methapo ea DA ho liketsahalo tse nyarosang, ho fana ka maikutlo a karolo ea ho etsa LHb ho hlahiseng matšoao a boleng ba DA (Gao et al., 1990).

LHb ke karolo ea tsela e kholo ea neural eo ka eona li-neurons tsa DA li ka laoloang ke basal ganglia (Setšoantšo sa 8). LHb e amohela lipontšo tse ts'oanang le liphoso tsa ho bolela moputso ka ho hakanngoa ho tsoa ho palo ea li-neuron tse haufi le moeli oa globus pallidus (GPb) (Hong le Hikosaka, 2008). Hang ha lipontšo tsena li fihla ho LHb, li tla romelloa ho li-neuron tsa DA ka tsela e sa bonahaleng eo LHb e natefisang methapo ea kutlo ea methapo ea methapo ea methapo ea methapo (Gb neurons) eo ka tsela e thibelang li-neuron tsa DA (Ji le Shepard, 2007; Omelchenko et al., 2009; Brinschwitz et al., 2010). Sena se ka hlaha ka ho reroa ha LHb lipakeng tsa VTA le khubung e haufi ea Gaba-ergic e bitsoang rostromedial tegmental nucleus (RMTg) (Jhou et al., 2009b) (e boetse e bitsoa mohatla oa "caudal oa VTA" (Kaufling et al., 2009)). Haholo-holo, li-neuron tsa RMTg li na le likarolo tsa karabelo tse tšoanang le LHb neurons, boleng ba encode, hape li na le projeke e boima ea thibelo ea dopaminergic midbrain (Jhou et al., 2009a). Kahoo, tsela e felletseng ea basal ganglia ea ho romella lipontšo tsa boleng bo botle ho li-neuron tsa DA e kanna ea ba GPb → LHb → RMTg → DA (Hikosaka, 2010).

Potso ea bohlokoa bakeng sa lipatlisiso tsa nako e tlang ke hore na matšoao a boleng bo susumetsang a channoa feela ka LHb kapa hore na li tsamaisoa ke litsela tse ngata tse kenyellelitsoeng. Haholo-holo, lithibelo tsa DA tse thibeloang ke makhopho a laolehang li laoloa ke ts'ebetso ho mesopontine parabrachial nucleus (PBN) (Coizet et al., 2010) (Setšoantšo sa 8). Lenonyeletso lena le na le li-neurons tse fumanang ho kenella ka kotloloho ho tsoa lesapo la mokokotlo le koahelang menahano e seng kotsi mme e ka thibela li-neuron tsa DA ka lipalo tse nyarosang ho RMTg (Coizet et al., 2010; Gauriau le Bernard, 2002). Sena se fana ka maikutlo a hore LHb e romella lipontšo tsa boleng ba "li-neurons" tsa DA bakeng sa melemo le litlamorao tse khahlisang ha PBN e fana ka karolo ea lets'oao la boleng le amanang ka kotloloho le sephetho sa boiketo.

Mehloli ea matšoao a ts'usumetso ea ts'usumetso

Ho nyane ho tsebahala ka mohloli oa matšoao a ts'usumetso ea ts'usumetso ho li-neuron tsa DA. Moemeli e mong ea khahlang ke karolo ea bohlokoa ea amygdala (CeA) e 'nileng ea ts'oaroa khahlano le maikutlo, maikutlo le likarabelo tse akaretsang nakong ea liketsahalo tse monate le tse nyarosang (Holland le Gallagher, 1999; Baxter le Murray, 2002; Merali et al., 2003; Balleine le Killcross, 2006) (Setšoantšo sa 8). CeA le li-nery tse ling tsa amygdala li na le li-neuron tse ngata tse bonts'ang melaetsa e lumellanang le ts'usumetso ea ts'usumetso: li supa liketsahalo tse thabisang le tse nyarosang ka lehlakoreng le le leng, lia matlafatsoa ha liketsahalo li etsahala ka tshohanyetso, mme li amana le mekhoa ea boits'oaro ea mofets'e (Nishijo et al., 1988; Belova et al., 2007; Shabel le Janak, 2009). matšoao ana e ka romeloa ho Da methapo ea kutlo hobane CeA e theoha dipeakanyo ho brainstem eo tsoele lesedi putsang le aversive (Lee le al., 2005; Pascoe le Kapp, 1985) le CeA ea hlokahala bakeng sa tokollo ea DA nakong ea liketsahalo tse amanang le moputso (Phillips et al., 2003a). Ntle le moo, CeA e nka karolo le li-neuron tsa DA litseleng tse tsamaellanang le marang-rang a rona a ts'ebetso ea ts'ebetso le ts'ebetso bakeng sa boiketlo bo susumetsang. Mmila o kenyelletsang CeA, SNc, le dorsal striatum oa hlokahala bakeng sa ho ithuta tloaelo ho litekanyetso tsa lijo (Han et al., 1997; Lee le al., 2005; El-Amamy le Holland, 2007). Tumellanong le karolo ea rona ea masapo le boleng ba bohlokoa ba tsela, tsela ena e ea hlokahala bakeng sa ho ithuta ho latela mekhoa ea lijo empa eseng bakeng sa ho ithuta ho fihlela sephetho sa lijo (Han et al., 1997). Tsela ea bobeli, ho kenyelletsa mokokotlo oa CeA, SNc, VTA, le NAc, e hlokahala bakeng sa mekhoa ea meputso ho etsa hore keketseho e kholo ea khothatso ea ho etsa liketso tse batlang meputso (Hall et al., 2001; Corbit le Balleine, 2005; El-Amamy le Holland, 2007).

Ntle le CeA, li-neurons tsa DA li ka fumana matšoao a matla a ho hlohlona ho tsoa mehloling e meng joalo ka methapo ea methapo ea methapo e teng mothong (basal)Lin le Nicolelis, 2008; Richardson le DeLong, 1991) le li-neuron ho PBN (Coizet et al., 2010), leha litsela tsena li ntse li lokela ho hlahlojoa.

Mehloli ea matšoao a lemosang

Ho na le likhetho tse 'maloa tse ntle bakeng sa ho fana ka li-neurons tsa DA ka matšoao a lemosang. Mohlomong mokhethoa o motle ka ho fetisisa ke colliculus (SC) e phahameng haholo, e amohelang maikutlo a mokhuts'oanyane a morao-rao a tsoang mehopolong e mengata ea kutlo le ho laola maikutlo le maikutlo (Redgrave le Gurney, 2006) (Setšoantšo sa 8). SC e na le tatelano e tobileng ho SNc le VTA (May et al., 2009; Comoli et al., 2003). Liphoofolong tse sa koaheloang ke letho ke mokhoa oa bohlokoa bakeng sa matšoao a bokhutšoanyane a morao ho fihlella li-neuron tsa DA le ho tsosa ho lokolloa ha DA ka meaho e thellang (Comoli et al., 2003; Dommett et al., 2005). Tsela ea SC-DA e loketse hantle haholo ho fetisa matšoao a lemosang ho fapana le moputso le lipontšo tsa ho kheloha, hobane li-neurons tsa SC ha li na karabelo e fokolang ho iseng phano ea moputso mme li na le tšusumetso e nyane feela ho likarabelo tsa DA tse arohaneng (Coizet et al., 2006). Sena se fana ka maikutlo a tatellano ea liketsahalo tseo ho tsona li-neur neurons (1) li e khethang e le bohlokoa, (2) e e khethang e le bohlokoa ka ho fetesisa, (3) e tsosa karabelo ea ho shebisisa ts'usumetso, le (4) ka nako e ts'oanang e tsosang karabelo ea tlhokomeliso ea DA e bakang ho phatloha ha DA ho meaho e tlase (Redgrave le Gurney, 2006).

Lekhetho la bobeli bakeng sa ho romella matšoao a lemosang ho li-neuron tsa DA ke LHb (Setšoantšo sa 8). Haholo, ho qala ho sa lebelloeng ha teko ea ho qala ha teko ho thibela li-neuron tse ngata tsa LHb ka tsela e sa lumellaneng le temoso ea tlhokomeliso ea neuron ea DA, mme karabelo ena e etsahala motsotsong o mokhuts'oane ho LHb e lumellanang le tataiso ea LHb → DA ea phetiso (Bromberg-Martin et al., 2010a; Bromberg-Martin et al., 2010c). Re boetse re hlokometse ka mokhoa o hlakileng hore li-neuron tsa LHb hangata li thibelloa ke litšoantšo tse sa lebelloang tsa pono le molumo ka tsela e hanyetsanang ho thabo ea DA (MM, ESB-M., Le OH, lipatlisiso tse sa phatlalatsoang) leha sena se emetse tlhahlobo e hlophisehileng haholoanyane.

Kamora nako, moemeli oa boraro bakeng sa ho romella matšoao a lemosang ho li-neuron tsa DA ke pedunculopontine tegmental nucleus (PPTg), e sebetsang ho SNc le VTA mme e kenya letsoho ts'ebetsong ea khothalletso (Ntlo, 2006) (Setšoantšo sa 8). PPTg e bohlokoa bakeng sa ho nolofalletsa ho phatloha ha li-neuron tsa VTA DA (Mohau et al., 2007) ho kenyelletsa likarabo tsa ho phatloha hoa meputsoPan le Hyland, 2005). E lumellana le letšoao le lemosang, li-neuron tsa PPTg li na le likarabo tse khutšoane ho latsoa mefuta e mengata ea maikutlo ebile li mafolofolo nakong ea karabelo e lebisang (Ntlo, 2006). Ho na le bopaki ba hore likarabo tsa maikutlo a PPTg li susumetsoa ke boleng ba moputso le litlhoko tsa ketso e potlakileng (Dormont et al., 1998; Okada et al., 2009) (empa bona (Pan le Hyland, 2005)). Li-neurons tse ling tsa PPTg le tsona li arabela liphellong tse khotsofatsang kapa tse khelohang ka botsona (Dormont et al., 1998; Kobayashi et al., 2002; Ivlieva le Timofeeva, 2003b, a). Ho tla ba bohlokoa ho hlahloba hore na lipontšo tseo PPTg e li romellang ho li-neuron tsa DA li amana ka ho khetheha le ho lemosa kapa hore na li na le matšoao a mang a susumetsang a kang boleng le ho hloka kelello.

Mosebetsi ona o ile oa tšehetsoa ke lenaneo la lipatlisiso la intramera Setsing sa Sechaba sa Leihlo. Re boetse re leboha Amy Arnsten ka lipuisano tsa bohlokoa.

Boitlhotlhollo ba Mohoeletsi: Ena ke faele ea PDF ea mongolo o ngotsoeng ka letsoho o sa amoheloang o amoheletsoeng bakeng sa ho hatisoa. Joaloka tšebeletso ho bareki ba rona re fana ka phetolelo ena ea pele ea mongolo o ngotsoeng ka letsoho. Mongolo o ngotsoeng ka letsoho o tla ba le ho kopitsa, ho kenya lihlopha le ho hlahloba bopaki bo hlahang pele o hatisoa ka mokhoa oa oona oa ho qetela. Ka kopo hlokomela hore nakong ea mekhoa ea tlhahiso ea lihlahisoa e ka fumanoa e ka amang litaba, le litsebiso tsohle tsa molao tse sebetsang koranteng e amanang le eona.

^FOOTNOTE1By boikaketsi bo susumetsang re bolela palo e phahameng bakeng sa liketsahalo tse monate le tse tlatsetsang ho tsona ebile e le tlase bakeng sa liketsahalo tse sa nke lehlakore (tse sa ruiseng letho le tse sa amaneng). Sena se tšoana le tlhaloso e fanoeng ke (Berridge le Robinson, 1998). Hlokomela hore ho hlohlona ho susumetsang ho fapane le maikutlo a mang a ho se sebetse ka ts'ebetso ea methapo ea kutlo, joalo ka ts'usumetso ea ts'usumetso (e sebetsang feela liketsahalong tse lakatsehang; (Berridge le Robinson, 1998)) le sesapo sa pono (se sebetsang liketsahalong tse sa nke lehlakore tse kang lintho tse tsamaeang le mabone a mebala;Bisley le Goldberg, 2010)).]

^FOOTNOTE2Hlokomela hore matšoao a khothaletso a ho ngola li-neuron tsa DA a fapane le maikutlo a "ho ts'oaroa" le "phetoho ho ts'oanelehang" e reriloeng ho laola sekhahla sa thuto ea matlafatso (mohlala (Pearce le Hall, 1980)). Likhopolo tse joalo li re liphoofolo li ithuta (le ho fetola litekanyetso tsa ho ithuta) ho tsoa liphosong tse mpe tse mpe tse boletsoeng esale pele. Leha li-neuron tsena tsa DA li ka kenya letsoho ho ithuteng ho tsoa liphosong tse boletsoeng esale pele, ka nako eo li ka ba le karabelo e matla (mohlala ho tlisa phano e sa lebelloang), li kanna tsa se ke tsa kenya letsoho ho ithuteng liphosong tse mpe tse boletsoeng esale pele, tseo ka tsona ba ka bang le karabelo e nyane kapa ea ho se be le tsona ( mohlala, ho lahla moputso o sa lebelloang) (Setšoantšo sa 4B).

1. Ahlbrecht M, Weber M. Tharollo ea ho hloka botsitso: thuto ea liteko. Tlaleho ea litsebi tsa moruo oa setheo le thuto ea bolumeli. 1996; 152: 593-607.
2. Albin RL, Young AB, Penney JB. Anatomy e sebetsang ea mathata a basal ganglia. Mekhoa ea li-neuroscience. 1989; 12: 366-375. [E fetotsoe]
3. Anderson AK, Christoff K, Stappen I, Panitz D, Ghahremani DG, Glover G, Gabrieli JD, Sobel N. Lipontšo tsa neural tse ikhethileng tsa matla le bokhopo mosebetsing oa batho. Nat Neurosci. 2003; 6: 196-202. [E fetotsoe]
4. Anstrom KK, Miczek KA, Budygin EA. Keketseho ea phasic dopamine e tšoaeang tseleng ea mesolimbic nakong ea tlhaselo ea sechaba ka likhoto. Neuroscience. 2009; 161: 3-12. [E fetotsoe]
5. Aragona BJ, Day JJ, Roitman MF, Cleaveland NA, Wightman RM, Carelli RM. Ho khetheha ha tikoloho ho nts'etsopele ea nako ea 'nete ea phasic dopamine phetisetso ea nako nakong ea ho fumana mokhatlo oa cue-cocaine ho likhoto. Koranta ea Europe ea neuroscience. 2009; 30: 1889-1899. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
6. Austin AJ, Duka T. Mekhoa ea ho tsepamisa maikutlo bakeng sa litabatabelo le sepheo se setle sa maemo a Pavlovian. Patlisiso e mabapi le boitsoaro. 2010; 213: 19-26. [E fetotsoe]
7. Badia P, Harsh J, Abbott B. Ho khetha lipakeng tsa maemo a tšohanyetso le a sa lebelloang: Tšoaetso le Thopolo. Bulletin ea kelello. 1979; 86: 1107-1131.
8. Balleine BW, Killcross S. Parallel ts'usumetso ea ts'usumetso: pono e kopaneng ea ts'ebetso ea amygdala. Mekhoa ea li-neuroscience. 2006; 29: 272-279. [E fetotsoe]
9. Balleine BW, O'Doherty JP. Li-homology tsa batho le tsa litoeba tse laoloang ke liketso: li-corticostriatal tse khethollang sepheo se tataisoang ke sepheo le tloaelo. Neuropsychopharmacology. 2010; 35: 48-69. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
10. Barr GA, Moriceau S, Shionoya K, Muzny K, Gao P, Wang S, Sullivan RM. Liphetoho lithutong tsa masea li fetoloa ke dopamine ho amygdala. Nat Neurosci. 2009; 12: 1364-1366. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
11. Bassareo V, De Luca MA, Di Chiara G. Tlhaloso e Ikhethileng ea Lits'usumetso tsa Matla a Ho Tsosa Motheo ka Dopamine ho Nucleus Accumbens Shell e bapisoang le Coreex le Cortex ea Pele. J Neurosci. 2002; 22: 4709-4719. [E fetotsoe]
12. Bassareo V, Di Chiara G. Karabelo e fapaneng ea phetiso ea dopamine ho lijo-tse tsosang ho li-compartments tsa li-nucleus accumbens. Neuroscience. 1999; 89: 637-641. [E fetotsoe]
13. Baxter MG, Murray EA. Monate le moputso. Nat Rev Neurosci. 2002; 3: 563-573. [E fetotsoe]
14. Bayer HM, Glimcher PW. Midbrain dopamine neurons e kenyelletsa pontšo ea phoso ea moputso e bongata. Neuron. 2005; 47: 129-141. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
15. Belova MA, Paton JJ, Morrison SE, Salzman CD. Tebello e modulates likarabo tsa neural ho tse monate le tse khutsitseng ho li-amygdala tsa mehleng. Neuron. 2007; 55: 970-984. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
16. Bentivoglio M, Morelli M. Mokhatlo le li-circuits tsa mesencephalic dopaminergic neurons le kabo ea li-receptor tsa dopamine bokong. Handbook of Chemical Neuroanatomy. 2005: 1-107.
17. Berridge KC, Robinson TE. Karolo ea dopamine moputsong ke efe: tšusumetso ea hedonic, ho ithuta ka moputso, kapa ho hlohlona? Patlisiso ea Brain. 1998; 28: 309-369. [E fetotsoe]
18. Besson C, Louilot A. Ho kenya letsoho ha mesmimbic dopaminergic neurons ka pono e amehang. Neuroscience. 1995; 68: 963-968. [E fetotsoe]
19. Birgner C, Nordenankar K, Lundblad M, Mendez JA, Smith C, le Greves M, Galter D, Olson L, Fredriksson A, Trudeau LE, et al. VGLUT2 ho dopamine neurons e hlokahala bakeng sa ts'ebetso ea ts'ebetso ea ts'ebetso ea psychostimulant. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2010; 107: 389-394. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
20. Bisley JW, Goldberg ME. Tlhokomeliso, sepheo, le pele ho parietal lobe. Tlhahlobo ea selemo le selemo ea neuroscience. 2010; 33: 1-21. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
21. Bjorklund A, Dunnett SB. Dopamine methapo ea kutlo bokong: ntjhafatso. Mekhoa ea li-neuroscience. 2007; 30: 194-202. [E fetotsoe]
22. Blazquez PM, Fujii N, Kojima J, Greybiel AM. Tlhahiso ea marang-rang ea monyetla oa ho arabela ho striatum. Neuron. 2002; 33: 973-982. [E fetotsoe]
23. Botvinick MM, Braver TS, Barch DM, Carter CS, Cohen JD. Tlhahlobo ea likhohlano le taolo ea kelello. Psychol Rev. 2001; 108: 624-652. [E fetotsoe]
24. Bradley MM, Greenwald MK, Petry MC, Lang PJ. Ho hopola litšoantšo: monyaka le tse monate kelellong. J Exp Psychol Ithute Mem Cogn. 1992; 18: 379-390. [E fetotsoe]
25. Braun DA, Mehring C, Wolpert DM. Moralo oa ho ithuta ka ketso. Patlisiso e mabapi le boitsoaro. 2010; 206: 157-165. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
26. Brinschwitz K, Dittgen A, Madai VI, Lommel R, Geisler S, Veh RW. Li-axon tsagic tsa glutamatergic tse tsoang habenula ea morao-rao li khaola haholo ho li-neurons tsa GABAergic tsa "ventral midbrain". Neuroscience. 2010; 168: 463-476. [E fetotsoe]
27. Brischoux F, Chakraborty S, Brierley DI, Ungless MA. Phasic excavation ea dopamine neurons ho ventral VTA ka tšusumetso e se nang boikemisetso. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2009; 106: 4894-4899. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
28. Bromberg-Martin ES, Hikosaka O. Midbrain dopamine neurons sesupo sa khetho bakeng sa tlhaiso-leseling e mabapi le meputso e tlang. Neuron. 2009; 63: 119-126. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
29. Bromberg-Martin ES, Matsumoto M, Hikosaka O. distinct activic tonic le phasic antential activate in lateral habenula le dopamine neurons. Neuron. 2010a; 67: 144-155. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
30. Bromberg-Martin ES, Matsumoto M, Hong S, Hikosaka O. Tsela ea pallidus-habenula-dopamine e tsamaisang litekanyetso tsa tšusumetso. J Neurophysiol. 2010b; 104: 1068-1076. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
31. Bromberg-Martin ES, Matsumoto M, Nakahara H, Hikosaka O. Li-timescales tse ngata tsa mohopolo ho habenula ea habenula le dopamine neurons. Neuron. 2010c; 67: 499-510. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
32. Brown MTC, Henny P, Bolam JP, Magill PJ. Ketsahalo ea li-neurochemically heterogenible dopaminergic neurons ho "antiantianti "ka bongata nakong ea ho fetoha ha boithatelo le ho tsamaisoang hoa boko. J Neurosci. 2009; 29: 2915-2925. [E fetotsoe]
33. Mok'hadinale RN. Litsamaiso tsa Neural li kenyelletsa ts'ebelisong e liehang le e matlafatsang. Neural Netw. 2006; 19: 1277-1301. [E fetotsoe]
34. Cheer JF, Aragona BJ, Heien ML, Seipel AT, Carelli RM, Wightman RM. Khokahano e kopantsoeng ea "dopamine" e kopanetsoeng le ts'ebetso e tsamaisitsoeng ea neural. Neuron. 2007; 54: 237-244. [E fetotsoe]
35. Cheramy A, Kemel ML, Gauchy C, Desce JM, Galli T, Barbeito L, Glowinski J. Role oa li-amino acid tse thabisang ka ho otloloha le ka kotloloho melaoana ea dopamine e lokollotsoeng litsong tsa methapo ea methapo ea methapo ea methapo ea methapo. Amino Acids. 1991; 1: 351-363. [E fetotsoe]
36. Chew SH, Ho JL. Tšepo: thuto e susumetsang ea maikutlo mabapi le nako ea tharollo e sa tsitsang. Tlaleho ea Kotsi le Ho se Tsitse. 1994; 8: 267-288.
37. Chiodo LA, Antelman SM, Caggiula AR, Lineberry CG. Tsela e matla ea ho fetola likhahla tsa li-dopamine (DA) li-neurons: bopaki ba mefuta e 'meli e sebetsang ea lisele tsa DA tse khōlō tsa nigra. Resin ea Boko. 1980; 189: 544-549. [E fetotsoe]
38. Christoph GR, Leonzio RJ, Wilcox KS. Ho hlohlona ha habenula ea morao-rao ho thibela li-dopamine tse nang le li-neuron sebakeng sa substantia nigra le ventral tegmental ea rat. J Neurosci. 1986; 6: 613-619. [E fetotsoe]
39. Chuhma N, Choi WY, Mingote S, Rayport S. Dopamine neuron glutamate cotransication: modulation e itšetlehileng ka maqhubu ho projeke ea mesoventromedial. Neuroscience. 2009; 164: 1068-1083. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
40. Cohen JD, Braver TS, Brown JW. Mehopolo e mabapi le dopamine e sebetsa ka pele ho cortex. Maikutlo a hona joale ho neurobiology. 2002; 12: 223-229. [E fetotsoe]
41. Coizet V, Dommett EJ, Klop EM, Redgrave P, Overton PG. Leqhubu la parabrachial ke sehokelo sa bohlokoa phetisetsong ea leseli le khuts'oanyane la latency nocicrit ho li-neuron tsa midbrain dopaminergic neurons. Neuroscience. 2010; 168: 263-272. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
42. Coizet V, Dommett EJ, Redgrave P, Overton PG. Likarabo tse se nang kelello tsa li-neurones tsa "midbrain dopaminergic neurones" li fetoloa ke "colliculus" e phahameng ka ho fetisisa ho rat. Neuroscience. 2006; 139: 1479-1493. [E fetotsoe]
43. Comoli E, Coizet V, Boyes J, Bolam JP, Cantera NS, Quirk RH, Overton PG, Redgrave P. Takanyo e tobileng ho tsoa ho colliculus e kholo ho fihlela haholoanti nigra bakeng sa ho fumana liketsahalo tse ikhethang. Nat Neurosci. 2003; 6: 974-980. [E fetotsoe]
44. Corbit LH, Balleine BW. Ho ikarola habeli ka marang-rang a lesapo la basolateral le bohareng ba amygdala ka mefuta e akaretsang le e tobileng ea phetisetso ea pavlovian-instrumental. J Neurosci. 2005; 25: 962-970. [E fetotsoe]
45. Diking JW, Laane K, Theobald DE, Armstrong HC, Corlett PR, Chudasama Y, Robbins TW. Nako e lekanyelitsoeng ea nako ea memori ea ho rata mahlo ea Patlovian ka D1 le li-receptor tsa NMDA ho li-nucleus accumbens. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2005; 102: 6189-6194. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
46. Daly HB. Khetho ea ho se ikemisetse e khutlisetsoa morao ha lintho tse sa lebelloang tse se nang sepheo li le matla: lits'ebetso, datha, le likhopolo tsa ho rata ho bona karabelo. Ka: Gormezano I, Wasserman EA, bahlophisi. Ho ithuta le ho hopola: Likarolo tsa Behavial and Biological. Litho tsa; 1992. maq. 81-104.
47. Davidson MC, Horvitz JC, Tottenham N, Fossella JA, Watts R, Ulug AM, Casey BJ. Phapang e phapang le ts'ebetso ea ts'ebetso e latela ts'usumetso e sa lebelloang. NeuroImage. 2004; 23: 1039-1045. [E fetotsoe]
48. Letsatsi JJ, Roitman MF, Wightman RM, Carelli RM. Ho ithuta ho kopanetsanang ho fetoha le liphetoho tse matla ho dopamine ho saena ho li-nucleus tsa li-nucleus. Nat Neurosci. 2007; 10: 1020-1028. [E fetotsoe]
49. Dayan P, Niv Y. Ho ithuta ho matlafatsa: botle, bobe le bobe. Maikutlo a hona joale ho neurobiology. 2008; 18: 185-196. [E fetotsoe]
50. Descarries L, Berube-Carriere N, Riad M, Bo GD, Mendez JA, Trudeau LE. Glutamate ho dopamine neurons: transaptic khahlanong le phetiso. Litlhahlobo tsa lipatlisiso tsa Brain. 2008; 58: 290-302. [E fetotsoe]
51. Di Chiara G. Nucleus e bokella likhetla le dopamine ea mantlha: Karolo e khethollang boits'oarong le bokhobeng. Patlisiso e mabapi le boitsoaro. 2002; 137: 75-114. [E fetotsoe]
52. Dommett E, Coizet V, Blaha CD, Martindale J, Lefebvre V, Walton N, Mayhew JE, Overton PG, Redgrave P. How visual activate dopaminergic neurons at latency e khuts'oane. Mahlale. 2005; 307: 1476-1479. [E fetotsoe]
53. Dormont JF, Conde H, Farin D. Karolo ea sekhahla sa "patunculopontine tegmental nucleus" mabapi le ts'ebetso ea koloi e maemong a katse. I. Ketsahalong e le 'ngoe e amanang le ts'ebeliso ea ts'ebeliso ea ts'ebetso Patlisiso ea kelello ea kelello. Liteko Hirnforschung. 1998; 121: 401-410. [E fetotsoe]
54. Duzel E, Bunzeck N, Guitart-Masip M, Duzel S. NOvelty-tlhotlheletso e amanang le tebello le ho fuputsoa ke dopamine (NOMAD): litlamorao bakeng sa botsofali bo phetseng hantle. Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2010; 34: 660-669. [E fetotsoe]
55. El-Amamy H, Holland PC. Litlamorao tse khahlisang tsa ho khaola li-amygdala khubu ea mantlha ho tsoa sebakeng sa tefo ea karohano kapa substantia nigra ho thuto e tataisang e susumetsang le tšusumetso. Koranta ea Europe ea neuroscience. 2007; 25: 1557-1567. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
56. Ettenberg A. Mohanyetsi o sebetsa thepa ea k'hok'heine e itlhommeng pele. Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2004; 27: 721-728. [E fetotsoe]
57. Fadok JP, Dickerson TM, Palmiter RD. Dopamine e ea hlokahala bakeng sa maemo a tšabo a itšetlehileng ka cue. J Neurosci. 2009; 29: 11089-11097. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
58. Fairhall AL, Lewen GD, Bialek W, de Ruyter Van Steveninck RR. Ho sebetsa hantle le ho nepahala ha khoutu e lumellanang ea neural. Tlhaho. 2001; 412: 787-792. [E fetotsoe]
59. Faure A, Haberland U, Conde F, El Massioui N. Lesion ho sisteme ea nigrostriatal dopamine e sitisa sebopeho sa tloaelo ea karabelo. J Neurosci. 2005; 25: 2771-2780. [E fetotsoe]
60. Faure A, Reynolds SM, Richard JM, Berridge KC. Mesolimbic dopamine ka takatso le tšabo: ho susumetsa ho susumetsoa ke litšitiso tsa lehae tsa glutamate tse fumanehang ho li-nucleus accumbens. J Neurosci. 2008; 28: 7184-7192. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
61. CDF Filloillo, Newsome WT, Schultz W. Ho nepahala ha nakoana ha ponelopele ea moputso ho dopamine neurons. Nat Neurosci. 2008; 11: 966-973. [E fetotsoe]
62. Fiorillo CD, Tobler PN, Schultz W. Discodcoding ea menyetla ea meputso le ho hloka botsitso ke li-dopamine neurons. Mahlale. 2003; 299: 1898-1902. [E fetotsoe]
63. Floresco SB, Ghods-Sharifi S, Vexelman C, Magyar O. Likarolo tse sa lumellaneng tsa li-nucleus li ka bokella motheo le khetla ho laola ho fetoha ha nako. J Neurosci. 2006; 26: 2449-2457. [E fetotsoe]
64. Ford CP, Gantz SC, Phillips PE, Williams JT. Taolo ea extracellular dopamine libakeng tsa dendrite le axon. J Neurosci. 2010; 30: 6975-6983. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
65. Frank MJ. Phetoho ea matla a dopamine ka har'a basal ganglia: ak'haonte ea methapo ea methapo ea kutlo e amanang le taolo ea methapo le methapo ea Parkinsonism e sa tsebeng letho. Tlaleho ea "neuroscience" e etsang hore motho a utloisise. 2005; 17: 51-72. [E fetotsoe]
66. Frank MJ, Fossella JA. Neurogenetics le pharmacology ea ho ithuta, ho susumetsa, le kelello. Neuropsychopharmacology. 2010 [Tlhahiso ea mahala ea PMC] [E fetotsoe]
67. Frank MJ, Seeberger LC, O'Reilly RC. Ka rantipole kapa ka lere: ho ithuta ho matlafatsoa hoa temohisiso ho parkinsonism. Mahlale. 2004; 306: 1940-1943. [E fetotsoe]
68. Gallistel CR, Gibbon J. Nako, sekhahla le boemo. Psychol Rev. 2000; 107: 289-344. [E fetotsoe]
69. Gao DM, Jeaugey L, Pollak P, Benabid AL. Likarabo tse sa tsotelleng tse tsoang ho tsebiso e matla e tsoang ho li-neurons tsa "dopaminergic neurons" tsa "substantia nigra", "compacta" ka "rat" le litokiso tsa tsona ka liphumano tsa habenula tsa morao. Brain Res. 1990; 529: 315-319. [E fetotsoe]
70. Gauriau C, Bernard JF. Litsela tse bohloko le lipotoloho tse parabrachial ka hara rat. Boikoetliso ba 'mele. 2002; 87: 251-258. [E fetotsoe]
71. Geisler S, Zahm DS. Liphetoho tsa tikoloho e kengoang kahare ea teropo ho sub -atum ea ratomical bakeng sa mesebetsi e kopanetsoeng. Lingoloa tsa neurology e bapisang. 2005; 490: 270-294. [E fetotsoe]
72. Gerfen CR, Engber TM, Mahan LC, Susel Z, Chase TN, Monsma FJ, Jr, Sibley DR. D1 le D2 dopamine receptor-regulated gene expression of striatonigral and striatopallidal neurons. Mahlale. 1990; 250: 1429-1432. [E fetotsoe]
73. Ghods-Sharifi S, Floresco SB. Litlamorao tse fapaneng le phokotso ea boiteko bo hlahisoang ke inactivations ea nucleus e bokella mantlha kapa khetla. Behahlangual neuroscience. 2010; 124: 179-191. [E fetotsoe]
74. Gonon FG. Kamano e sa amaneng le pakeng tsa phallo ea tšusumetso le dopamine e ntšitsoeng ke "rat" midbrain dopaminergic neurons joalo ka ha e ithutile ho vivo electrochemistry. Neuroscience. 1988; 24: 19-28. [E fetotsoe]
75. Goto Y, Yang CR, Otani S. Ts'ebetso ea polasetiki e sebetsang le e sa sebetseng ho polasetiki ea pele: likarolo maikutlong a mafu a kelello. Tlholeho ea kelello. 2010; 67: 199-207. [E fetotsoe]
76. Mohau AA. Phasic bapisoa le tokiso ea dopamine ea phasic le phallo ea ts'ebetso ea phetoho ea dopamine: phetisetso bakeng sa etiology ea schizophrenia. Neuroscience. 1991; 41: 1-24. [E fetotsoe]
77. Mohau AA, Bunney BS. Intracellular le extracellular electrophysiology ea nigral dopaminergic neurons-1. Boitsebahatso le semelo. Boithuto ba kelello. 1983; 10: 301-315. [E fetotsoe]
78. Mohau AA, Floresco SB, Goto Y, Lodge DJ. Taolo ea ho thunya ha li-neurop tsa dopaminergic le taolo ea boits'oaro bo lebisitsoeng ho sepheo. Mekhoa ea li-neuroscience. 2007; 30: 220-227. [E fetotsoe]
79. Grecksch G, Matties H. Karolo ea methapo ea dopaminergic ho hippocampus ea rat ho kopanya ka khethollo e khanyang. Psychopharmacology (Berl) 1981; 75: 165-168. [E fetotsoe]
80. Guarraci FA, Kapp BS. Tšobotsi e khethiloeng ea elektrophysiological ea sebaka se ka hare sa mokokotlo oa dopaminergic neurons nakong ea phapang e fapaneng ea tšabo ea pavlovia ho 'mutla o tsohileng. Patlisiso e mabapi le boitsoaro. 1999; 99: 169-179. [E fetotsoe]
81. Moqapi SN, Fudge JL, McFarland NR. Litsela tsa Striatonigrostriatal li-primates li theha holimo ho tloha ho khetla ho ea ho li-dorolateral striatum. J Neurosci. 2000; 20: 2369-2382. [E fetotsoe]
82. Hall J, Parkinson JA, Connor TM, Dickinson A, Everitt BJ. Ho kenyelletsa karolo e bohareng ea "amygdala" le "nucleus" e bokellana ho matlafatsong ea ts'usumetso ea ts'usumetso ea Pavlovian ka boits'oaro ba sebopeho. Koranta ea Europe ea neuroscience. 2001; 13: 1984-1992. [E fetotsoe]
83. Han JS, McMahan RW, Holland P, Gallagher M. Karolo ea tsela ea bo-amygdalo-nigrostriatal thutong e amanang. J Neurosci. 1997; 17: 3913-3919. [E fetotsoe]
84. Harris GC, Aston-Jones G. Arousal le moputso: dichotomy ts'ebetsong ea orexin. Mekhoa ea li-neuroscience. 2006; 29: 571-577. [E fetotsoe]
85. Herry C, Bach DR, Esposito F, Di Salle F, Perrig WJ, Scheffler K, Luthi A, Seifritz E. Ho sebetsanoa le boiphetetso ba nakoana ho amygdala ea batho le liphoofolo. J Neurosci. 2007; 27: 5958-5966. [E fetotsoe]
86. Hikida T, Kimura K, Wada N, Fun fan K, Nakanishi S. distinct likarolo tse phetisetso ea phetiso ea testaptic ka kotloloho le ka kotloloho mekhoa e metle ea ho putsa le boits'oaro bo khathollang. Neuron. 2010; 66: 896-907. [E fetotsoe]
87. Hikosaka O. Basal ganglia mekhahlelo ea ho tsamaisa mahlo e lebisang ho moputso. Liphatlalatso tsa New York Academy of Science. 2007; 1104: 229-249. [E fetotsoe]
88. Hikosaka O. habenula: ho tloha ho khatello ea maikutlo ho ea ho liqeto tse thehiloeng holim'a boleng. Nat Rev Neurosci. 2010; 11: 503-513. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
89. Hikosaka O, Takikawa Y, Kawagoe R. Karolo ea sehlopha sa basal ganglia taolong ea methapo ea mahlo e reriloeng e nang le morero. Tlhahlobo ea 'mele. 2000; 80: 953-978. [E fetotsoe]
90. Hitchcott PK, Quinn JJ, Taylor JR. Ho feto-fetoha ha maikutlo ho Bidirectional ea liketso tse lebisitsoeng ho sepheo ke dopamine ea pele. Cereb Cortex. 2007; 17: 2820-2827. [E fetotsoe]
91. Hnasko TS, Chuhma N, Zhang H, Goh GY, Sulzer D, Palmiter RD, Rayport S, Edward RH. Vesicular glutamate ea lipapatso e phahamisa polokelo ea dopamine le glutamate elementlease ho vivo. Neuron. 2010; 65: 643-656. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
92. Holland PC, Gallagher M. Amygdala potolohong ea tlhokomeliso le boemeli. Mekhoa ea mahlale a kelello. 1999; 3: 65-73. [E fetotsoe]
93. Hollerman JR, Schultz W. Dopamine neurons e tlaleha phoso e boletsoeng esale pele ea moputso nakong ea ho ithuta. Nat Neurosci. 1998; 1: 304-309. [E fetotsoe]
94. Holroyd CB, Coles MG. Motheo oa neural oa ts'ebetso ea liphoso tsa motho: ho ithuta ho matlafatsa, dopamine, le negativity e amanang le phoso. Psychol Rev. 2002; 109: 679-709. [E fetotsoe]
95. Hong S, Hikosaka O. Global globus pallidus e romela lipontšo tse amanang le moputso ho habenula ea morao. Neuron. 2008; 60: 720-729. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
96. Horvitz JC. Mesolimbocortical le nigrostriatal dopamine li arabela liketsahalong tse se nang moputso. Khopolo-taba. 2000; 96: 651-656. [E fetotsoe]
97. Horvitz JC, Stewart T, Jacobs BL. Ketsahalo e senyang ea ventral tegmental dopamine neurons e hlahisoa ke tšusumetso e tsosang takatso ea ts'oaetso cat. Resin ea Boko. 1997; 759: 251-258. [E fetotsoe]
98. Houk JC, Adams JL, Barto AG. Mohlala oa kamoo sehlopha sa basal ganglia se hlahisang le ho sebelisa lipontšo tsa neural tse bolelang ho matlafatsa. Ka: Houk JC, Davis JL, Beiser DG, bahlophisi. Mefuta ea Ts'ebetso ea Tlhahisoleseling ho Basal Ganglia. Cambridge, MA: Press ea MIT; 1995. maq. 249-274.
99. Ikemoto S. Brain ajoa ka kotloloho ho feta sisteme ea mesolimbic dopamine: Khopolo ea neurobiological. Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2010 [Tlhahiso ea mahala ea PMC] [E fetotsoe]
100. Ito R, Diking JW, Howes SR, Robbins TW, Everitt BJ. Dissociation in Conditioned Dopamine Release in Nucleus Accumbens Core le Shell Karabelo ho Cues Cocaine le nakong ea Cocaine-Go Batla Ho Behaeve ho Rats. J Neurosci. 2000; 20: 7489-7495. [E fetotsoe]
101. Ivlieva NY, Timofeeva NO. Ts'ebetso ea Neuron mokokotlong oa pedunculopontine nakong ea ts'ebetso e amanang le lijo e nang le boemo ba Reflex. Neuroscience le Boitšoaro ba Bohlale. 2003a; 33: 919-928. [E fetotsoe]
102. Ivlieva NY, Timofeeva NO. Ts'ebetso ea Neuron mokokotlong oa pedunculopontine nakong ea "reflex" e sebetsang. Neuroscience le Boitšoaro ba Bohlale. 2003b; 33: 499-506. [E fetotsoe]
103. Jalabert M, Aston-Jones G, Herzog E, Manzoni O, Georges F. Karolo ea mokokotlo oa bethe ea stria terminalis taolong ea ventral tegmental area dopamine neurons. Tsoelo-pele ho neuro-psychopharmacology le psychiatry ea likokoana-hloko. 2009; 33: 1336-1346. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
104. Jeanblanc J, Hoeltzel A, Louilot A. Ho arohana ha ho kenella tšebetsong ea methapo ea methapo ea methapo ea methapo e kenang methapong ea mantlha le likhetla ha li-nucleus li bokellana ka thibelo ea morao-rao le pono e amehang. Neuroscience. 2002; 111: 315-323. [E fetotsoe]
105. Jensen J, Smith AJ, Willeit M, Crawley AP, Mikulis DJ, Vitcu I, Kapur S. Karohano ea likarolo tsa kh'outu bakeng sa bosence vs. valence nakong ea polelo ea moputso ho batho. Boko ba motho ba kelello. 2007; 28: 294-302. [E fetotsoe]
106. Jhou TC, Masimo HL, Baxter MG, Saper CB, PC ea Holland. Rostromedial tegmental nucleus (RMTg), e amanang le GABAergic e amanang le methapo ea methapo ea methapo ea methapo, e tsosa mohopolo o fokolisang le likarabo tsa makoloi. Neuron. 2009a; 61: 786-800. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
107. Jhou TC, Geisler S, Marinelli M, Degarmo BA, Zahm DS. Mesopontine rostromedial tegmental nucleus: Sebopeho se lebisitsoeng ke habenula ea morao-rao e sebetsang sebakeng sa likarolo tse ka hare tsa Tsai le substantia nigra compacta. Lingoloa tsa neurology e bapisang. 2009b; 513: 566-596. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
108. Ji H, Shepard PD. Ho hlohlometsa habenula ho hlohlelletsa litla-morao tsa "cell" "dopamine neurons ka mochine o amohelang li-receptor-mediated. J Neurosci. 2007; 27: 6923-6930. [E fetotsoe]
109. Jin X, Costa RM. Lipontšo tsa ho qala / ho emisa li hlaha makaleng a nigrostriatal nakong ea ho ithuta ka tatellano. Tlhaho. 2010; 466: 457-462. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
110. Johansen JP, Maseru HL. Glutamatergic activation ea anterior cingrate cortex e hlahisa lets'oao le sa rateheng la ho ruta. Nat Neurosci. 2004; 7: 398-403. [E fetotsoe]
111. Joseph MH, Datla K, Mohlankana AM. Tlhaloso ea tekanyo ea li-nucleus accumbens dopamine ka vivo dialysis: ho raha, ho lakatsa kapa ho tseba? Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2003; 27: 527-541. [E fetotsoe]
112. Joshua M, Adler A, Bergman H. Matla a dopamine taolong ea boitšoaro ba makoloi. Maikutlo a hona joale ho neurobiology. 2009a; 19: 615-620. [E fetotsoe]
113. Joshua M, Adler A, Mitelman R, Vaadia E, Bergman H. Midbrain dopaminergic neurons le li-striatal cholinergic interneurons li kenyelletsa phapang lipakeng tsa moputso le liketsahalo tse sa fetoheng liketsahalong tse fapaneng tsa liteko tsa maemo a maemo a bohlasoa. J Neurosci. 2008; 28: 11673-11684. [E fetotsoe]
114. Joshua M, Adler A, Prut Y, Vaadia E, Wickens JR, Bergman H. Synchronization ea li-neurb tsa dopaminergic neuron e matlafatsoa ke liketsahalo tse thabisang. Neuron. 2009b; 62: 695-704. [E fetotsoe]
115. Kable JW, Glimcher PW. The neurobiology ea qeto: tumellano le ngangisano. Neuron. 2009; 63: 733-745. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
116. Ke S, Dayan P. Dopamine: kakaretso le li-bonase. Li-network tsa Neural. 2002; 15: 549-559. [E fetotsoe]
117. Kapur S. Psychosis joalo ka boemo ba ho se sebetse hantle: moralo o hokahanyang biology, phenomenology, le pharmacology ho schizophrenia. Koranta ea Amerika ea mafu a kelello. 2003; 160: 13-23. [E fetotsoe]
118. Kaufling J, Veinante P, Pawlowski SA, Freund-Mercier MJ, Barrot M. Afferents ho mohatla oa GABAergic oa sebaka sa karolelano ea mocheso karolong. Lingoloa tsa neurology e bapisang. 2009; 513: 597-621. [E fetotsoe]
119. Kennerley SW, Wallis JD. Ho lekola likhetho tsa li-neuron tse kahare ho lobe ea pele: boleng ba sephetho bo kenyelelitsoe liqetong tse ngata tse fapaneng tsa liqeto. Koranta ea Europe ea neuroscience. 2009; 29: 2061-2073. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
120. Kim H, Sul JH, Huh N, Lee D, Jung MW. Karolo ea striatum ho ntlafatsa boleng ba liketso tse khethiloeng. J Neurosci. 2009; 29: 14701-14712. [E fetotsoe]
121. Kiyatkin EA. Mesebetsi ea tšebetsong ea li-dopamine tse nang le li-neurons tsa sebakeng se itseng sa likarolo tsa kelello ho likhoto tse hlokolosi. Int J Neurosci. 1988a; 42: 21-43. [E fetotsoe]
122. Kiyatkin EA. Morphine e susumetsoang ke morphine ea ts'ebetso ea li-neurons tsa sebakeng sa "ventral tegmental sebakeng" tse sebetsang hantle. Hare J Neuroscience. 1988b; 41: 57-70. [E fetotsoe]
123. Klingberg T. Koetliso le polasetiki ea mohopolo o sebetsang. Mekhoa ea mahlale a kelello. 2010; 14: 317-324. [E fetotsoe]
124. Kobayashi S, Nomoto K, Watanabe M, Hikosaka O, Schultz W, Sakagami M. Tšusumetso ea meputso le likotlo tse bohloko mosebetsing mabapi le macaque lateral preortalal cortex. Neuron. 2006; 51: 861-870. [E fetotsoe]
125. Kobayashi S, Schultz W. Tšusumetso ea ho lieha ha moputso ho likarabo tsa li-neurop tsa dopamine. J Neurosci. 2008; 28: 7837-7846. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
126. Kobayashi Y, Inoue Y, Yamamoto M, Isa T, Aizawa H. Tlatsetso ea methapo ea methapo ea methapo ea kutlo e amanang le ts'ebetso ea kutlo. J Neurophysiol. 2002; 88: 715-731. [E fetotsoe]
127. Koyama T, Tanaka YZ, Mikami A. Nocicepts neurons e hohelang ka ntle ho caate ea macaque nakong ea ho lebella ho utloa bohloko. Neuroreport. 1998; 9: 2663-2667. [E fetotsoe]
128. Kravitz AV, Freeze BS, Parker PR, Kay K, Thwin MT, Deisseroth K, Kreitzer AC. Taolo ea boits'oaro ba koloi ea parkinsonian ka taolo ea optogenetic ea basal ganglia circry. Tlhaho. 2010 [Tlhahiso ea mahala ea PMC] [E fetotsoe]
129. Lammel S, Hetzel A, Hackel O, Jones I, Liss B, Roeper J. Mefuta e ikhethileng ea li-neuropathy tsa mesoprefrontal kahare ho sistimi e kopanetsoeng ea mesocorticolimbic dopamine. Neuron. 2008; 57: 760-773. [E fetotsoe]
130. Lang PJ, Davis M. Emotion, tšusumetso le kelello: metheo ea Reflex lipatlisisong tsa liphoofolo le batho. Tsoelo-pele lipatlisisong tsa boko. 2006; 156: 3-29. [E fetotsoe]
131. Lapish CC, Kroener S, Durstewitz D, Lavin A, Basel JK. Bokhoni ba sistamine ea mesoportical dopamine ho sebetsa ka mekhoa e fapaneng ea nakoana ea nakoana. Psychopharmacology (Berl) 2007; 191: 609-625. [E fetotsoe]
132. Lee D, Seo H. Mechanisms ea ho matlafatsa thuto le ho etsa liqeto ka prortal dorsolateral prefrontal cortex. Liphatlalatso tsa New York Academy of Science. 2007; 1104: 108-122. [E fetotsoe]
133. Lee HJ, Groshek F, Petrovich GD, Cantalini JP, Gallagher M, Holland PC. Karolo ea potoloho ea amygdalo-nigral maemong a tšusumetso ea pono e pakiloeng ka lijo. J Neurosci. 2005; 25: 3881-3888. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
134. Levita L, Dalley JW, Robbins TW. Li-nyuklea li bokella dopamine mme li ithutile tšabo e boelisitsoeng: tlhahlobo le liphetho tse ling tse ncha. Patlisiso e mabapi le boitsoaro. 2002; 137: 115-127. [E fetotsoe]
135. Lin SC, Nicolelis MA. Neuronal Ensemble Bursting Sebakeng sa Basal Forebrain Encali Salience Irrespective of Valence. Neuron. 2008; 59: 138-149. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
136. Lisman JE, Mohau AA. Loop ea hippocampal-VTA: ho laola ho kenella ha tlhahisoleseling mohopolong oa nako e telele. Neuron. 2005; 46: 703-713. [E fetotsoe]
137. Litt A, Plassmann H, Shiv B, Rangel A. Ho khetholla lintlha tsa bohlokoa le matšoao a boits'oaro nakong ea ho etsa liqeto. Cereb Cortex. 2010 mochine oa khatiso. [E fetotsoe]
138. Liu Z, Richmond BJ, Murray EA, Saunders RC, Steenrod S, Stubblefield BK, Montague DM, Ginns EI. DNA e shebiloeng ka liprotheine tsa liginal cortex D2 receptor e thibela habonolo ho ithuta ka mekhoa e boletsoeng esale pele ka moputso. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2004; 101: 12336-12341. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
139. Liu ZH, Shin R, Ikemoto S. Karolo e le 'ngoe ea li-dopamine neuron tsa medial A10 dopamine neurons ho encoding e amehang. Neuropsychopharmacology. 2008; 33: 3010-3020. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
140. Ljungberg T, Apicella P, Schultz W. Likarabo tsa monkey dopamine neurons nakong ea ho ithuta ka boitšoaro bo botle. J Neurophysiol. 1992; 67: 145-163. [E fetotsoe]
141. Lobo MK, Covington HE, 3rd, Chaudhury D, Friedman AK, Sun H, Damez-Werno D, Dietz DM, Zaman S, Koo JW, Kennedy PJ, et al. Tahlehelo ea mofuta o khethehileng oa taolo ea BDNF ea ho supa mimics optogenetic ea moputso oa koae. Mahlale. 2010; 330: 385-390. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
142. Maeda H, Mogenson GJ. Litlamorao tsa ho hlohlelletsa ho potoloha hoa tšebetso ea li-neurons sebakeng sa ho kenella ka hare ho methapo, substantia nigra le midbrain reticular form of rats. Bulletin ea lipatlisiso. 1982; 8: 7-14. [E fetotsoe]
143. Mantz J, Thierry AM, Glowinski J. Phello ea tax ea tax e senang tekanyo ea litšila tsa mesocortical le mesolimbic dopamine neurons: ts'ebetso e khethollang ea mesocortical system. Brain Res. 1989; 476: 377-381. [E fetotsoe]
144. Margolis EB, Lock H, Hjelmstad GO, Masimong HL. Sebaka sa "ventral tegmental" se ntlafalitsoeng: na ho na le "letšoao la" electrophysiological "bakeng sa" dopaminergic neurons "? Tlaleho ea physiology. 2006; 577: 907-924. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
145. Margolis EB, Mitchell JM, Ishikawa J, Hjelmstad GO, Fields HL. Midbrain dopamine neurons: sepheo sa projeke se khetholla menyetla e ka bang teng ea nako le dopamine D (2) receptor inhibition. J Neurosci. 2008; 28: 8908-8913. [E fetotsoe]
146. Mark GP, Blander DS, Hoebel BG. Khothatso e nang le boemo e fokotseha dopamine ea extracellular ka har'a nucleus e bokellanang ka mor'a nts'etsopele ea ho ithuta ho latsoa tatso. Brain Res. 1991; 551: 308-310. [E fetotsoe]
147. Marowsky A, Yanagawa Y, Obata K, Vogt KE. Setsi se ikhethileng sa li-interneurons mediates dopaminergic activization ea mosebetsi oa amygdala. Neuron. 2005; 48: 1025-1037. [E fetotsoe]
148. Matsumoto M, Hikosaka O. Habenula ea morao-rao e le mohloli oa lipontšo tse mpe tsa moputso ho li-dopamine neurons. Tlhaho. 2007; 447: 1111-1115. [E fetotsoe]
149. Matsumoto M, Hikosaka O. Tlhahiso ea boleng bo bobe ba susumetso ho prenne hamorao habenula. Nat Neurosci. 2009a; 12: 77-84. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
150. Matsumoto M, Hikosaka O. Mefuta e 'meli ea dopamine neuron ka nepo e fana ka matšoao a khothatsang le a fosahetseng. Tlhaho. 2009b; 459: 837-841. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
151. Matsumoto M, Matsumoto K, Abe H, Tanaka K. Mosebetsi o ka pele oa sele o bonts'ang liphoso tsa boleng ba liketso. Nat Neurosci. 2007; 10: 647-656. [E fetotsoe]
152. May PJ, McHaffie JG, Stanford TR, Jiang H, Costello MG, Coizet V, Hayes LM, Haber SN, Redgrave P. Tectonigral projeke ka primate: tsela ea ho kenya tsebetso pele ho mamelwang methapo ea kutlo ea methapo ea kutlo ea midbrain dopaminergic neurons. Koranta ea Europe ea neuroscience. 2009; 29: 575-587. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
153. Mazzoni P, Hristova A, Krakauer JW. Hobaneng re sa tsamaye ka lebelo? Lefu la Parkinson, matla a ho sisinyeha, le tšusumetso e felletseng. J Neurosci. 2007; 27: 7105-7116. [E fetotsoe]
154. Merali Z, Michaud D, McIntosh J, Kent P, Anisman H. Ho nka karolo ho fapaneng ha li-amygdaloid CRH system (s) ho salience le valence ea tšusumetso. Tsoelo-pele ho neuro-psychopharmacology le psychiatry ea likokoana-hloko. 2003; 27: 1201-1212. [E fetotsoe]
155. Mirenowicz J, Schultz W. Ts'ebetso e khethehileng ea li-neuron tsa dopamine neurons ka takatso ea lijo ho fapana le ho susumetsa maikutlo. Tlhaho. 1996; 379: 449-451. [E fetotsoe]
156. Molina-Luna K, Pekanovic A, Rohrich S, Hertler B, Schubring-Giese M, Rioult-Pedotti MS, Luft AR. Dopamine ka motor cortex ea hlokahala bakeng sa ho ithuta ka bokhoni le polasetiki ea synaptic. MAHLOMOLA MOTHO. 2009; 4: e7082. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
157. Montague PR, Berns GS. Moruo oa Neural le likarolo tsa tlhaho tsa boleng. Neuron. 2002; 36: 265-284. [E fetotsoe]
158. Montague PR, Dayan P, Sejnowski TJ. Moralo oa litsamaiso tsa dopamine tsa mesencephalic tse thehiloeng ho thupelo ea ho ithuta ea Se-Hebere. J Neurosci. 1996; 16: 1936-1947. [E fetotsoe]
159. Morris G, Arkadir D, Nevet A, Vaadia E, Bergman H. Coincident empa e le melaetsa e hlakileng ea li-dbamine tsa midbrain le li-neurons tse sebetsang tsa "striatal tonics" tse sebetsang. Neuron. 2004; 43: 133-143. [E fetotsoe]
160. Morris G, Nevet A, Arkadir D, Vaadia E, Bergman H. Midbrain dopamine neurons encode liqeto bakeng sa ketso ea nako e tlang. Nat Neurosci. 2006; 9: 1057-1063. [E fetotsoe]
161. Morrison SE, Salzman CD. Khokahano ea tlhaiso-leseling e mabapi le tšusumetso e monate le e nyarosang ea li-neuron tse le 'ngoe. J Neurosci. 2009; 29: 11471-11483. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
162. Nakahara H, Itoh H, Kawagoe R, Takikawa Y, Hikosaka O. Dopamine neurons e ka emela phoso e boletsoeng esale pele ea polelo. Neuron. 2004; 41: 269-280. [E fetotsoe]
163. Nakamura K, Hikosaka O. Karolo ea dopamine karolong e ka pele ea tlhahiso ea "caudate" ho moputso oa li-saccade. J Neurosci. 2006; 26: 5360-5369. [E fetotsoe]
164. Neve KA, Seamans JK, Trantham-Davidson H. Dopamine receptor signaling. Tlaleho ea patlisiso le phetisetso ea transduction 2004; 24: 165-205. [E fetotsoe]
165. Nishijo H, Ono T, Nishino H. Single neuron likarabo ka amygdala ea monkey ea tlhokomeliso nakong ea ts'usumetso e rarahaneng ea maikutlo le bohlokoa bo amanang. J Neurosci. 1988; 8: 3570-3583. [E fetotsoe]
166. Niv Y, Daw ND, Joel D, dayan P. Tonic dopamine: litšenyehelo tsa menyetla le taolo ea matla a karabelo. Psychopharmacology. 2007; 191: 507-520. [E fetotsoe]
167. Nomoto K, Schultz W, Watanabe T, Sakagami M. Ka nako e telele likarabo tsa dopamine ka sepheo sa ho batla moputso esale pele. J Neurosci. 2010; 30: 10692-10702. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
168. Okada K, Toyama K, Inoue Y, Isa T, Kobayashi Y. Mefuta e fapaneng ea "patunculopontine tegmental neurons" e boletsoeng esale pele le moputso oa mosebetsi oa nnete. J Neurosci. 2009; 29: 4858-4870. [E fetotsoe]
169. Omelchenko N, Bell R, Sesack SR. Morero oa habenula oa morao-rao ho dopamine le li-neurons tsa GABA sebakeng sa "rat ventral tegmental" sebakeng. Koranta ea Europe ea neuroscience. 2009; 30: 1239-1250. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
170. Owesson-White CA, Ariansen J, Stuber GD, Cleaveland NA, Cheer JF, Wightman RM, Carelli RM. Encode encode ea boits'oaro ba koae e kopantsoeng, e kopane le phasic dopamine e lokolloang mokokotlong oa li-accumbens le khetla. Koranta ea Europe ea neuroscience. 2009; 30: 1117-1127. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
171. Oyama K, Hernadi I, Iijima T, Tsutsui K. Moputso oa ho profeta liphoso tsa dorsal striatal neurons. J Neurosci. 2010; 30: 11447-11457. [E fetotsoe]
172. Packard MG, White NM. Ho ikarola ha li-system tsa memori ea hippocampus le caudate ka ho romella ka morao ka morao ente ea intracerebral ea dopamine agonists. Behahlangual neuroscience. 1991; 105: 295-306. [E fetotsoe]
173. Padoa-Schioppa C. Orbitof Pambal cortex le kopanyo ea boleng ba moruo. Liphatlalatso tsa New York Academy of Science. 2007; 1121: 232-253. [E fetotsoe]
174. Palmiter RD. Dopamine ho supa ho dorsal striatum ho bohlokoa bakeng sa boits'oaro bo khothalletsoang: lithuto tse tsoang ho litoeba tse haelloang ke dopamine. Liphatlalatso tsa New York Academy of Science. 2008; 1129: 35-46. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
175. Pan WX, Hyland BI. Pedunculopontine tegmental nucleus laola maemo a arabeloang a li-neurb tsa dopamine neurons ka ho etsa litoeba. J Neurosci. 2005; 25: 4725-4732. [E fetotsoe]
176. Pan WX, Schmidt R, Wickens JR, Hyland BI. Mochine oa Tripartite oa ho timela o khothaletsoang ke tšebetso ea dopamine neuron le mofuta oa phapang ea nakoana. J Neurosci. 2008; 28: 9619-9631. [E fetotsoe]
177. Parker JG, Zweifel LS, Clark JJ, Evans SB, Phillips PE, Palmiter RD. Ho ba sieo ha li-receptor tsa NMDA ho dopamine neurons ho etsa hore dopamine e lokolloe empa e se boemo ba boemo nakong ea boemo ba Pavlovian. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2010 [Tlhahiso ea mahala ea PMC] [E fetotsoe]
178. Pascoe JP, Kapp BS. Litšoaneleho tsa electrophysiological tsa amygdaloid khubu ea li-nucleus neurons nakong ea maemo a ts'abo a Pavlovian ho 'mutla. Patlisiso e mabapi le boitsoaro. 1985; 16: 117-133. [E fetotsoe]
179. Pascucci T, Ventura R, Latagliata EC, Cabib S, Puglisi-Allegra S. Cortex ea medial prefrontal e khetholla karabelo ea dopamine ea khatello ea maikutlo ka tšusumetso e hanyetsang ea norepinephrine le dopamine. Cereb Cortex. 2007; 17: 2796-2804. [E fetotsoe]
180. Pearce JM, Hall G. Mohlala oa thuto ea Pavlovian: ho fapana ha ts'ebetso ea maemo empa eseng ho susumetsoa ke maemo. Psychol Rev. 1980; 87: 532-552. [E fetotsoe]
181. Pezze MA, Feldon J. Mesolimbic dopaminergic pathways maemong a tšabo. Tsoelo-pele ho neurobiology. 2004; 74: 301-320. [E fetotsoe]
182. Phillips AG, Ahn S, Howland JG. Ts'ebetso ea Amygdalar ea mesocorticolimbic dopamine system: Litsela tse bapileng tsa boitšoaro bo susumetsoang. Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2003a; 27: 543-554. [E fetotsoe]
183. Phillips GD, Salussolia E, Hitchcott PK. Karolo ea lenane la tšebetso ea dopamine ea machoamygdaloid thutong ea maikutlo. Psychopharmacology. 2010 [E fetotsoe]
184. Phillips PE, Stuber GD, Heien ML, Wightman RM, Carelli RM. Ho lokolloa ha dopamine ea subsecond ho khothaletsa ho batla koae. Tlhaho. 2003b; 422: 614-618. [E fetotsoe]
185. Porrino LJ, Goldman-Rakic ​​PS. Brainstem patlisiso ea cortex ea pele le ea anterior cortex ka litšoelesa tsa rhesus e senotsoe ke ho khutla ha HRP. Lingoloa tsa neurology e bapisang. 1982; 205: 63-76. [E fetotsoe]
186. Puryear CB, Kim MJ, Mizumori SJ. Ho kopanya ka mokhoa o kopanyang oa ho sisinyeha le moputso ka li-neuron tsa sebakeng se kenellang hare sebakeng se tsamaeang ka mokhoa oa bolokolohi. Behahlangual neuroscience. 2010; 124: 234-247. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
187. Ravel S, Legallet E, Apicella P. Li-neuron tse sebetsang ka mokhoa o hlophisitsoeng hantle ka har'a monkey striatum ha li arabe ka mokhoa o khahlisang takatso ea maikutlo. Patlisiso ea kelello ea kelello. Liteko Hirnforschung. 1999; 128: 531-534. [E fetotsoe]
188. Ravel S, Legallet E, Apicella P. Karabelo ea li-neuron tse sebetsang ka mokhoa oa toni ho khethollo ea litheko lipakeng tsa ts'usumetso e hanyetsang. J Neurosci. 2003; 23: 8489-8497. [E fetotsoe]
189. Ravel S, Richmond BJ. Ho araba likarabo tsa li-neuronal ka litšoene tse fanang ka moputso oa pono. Koranta ea Europe ea neuroscience. 2006; 24: 277-290. [E fetotsoe]
190. Redgrave P, Gurney K. Letshwao la dopamine le bokhuts'oanyane: karolo ea ho fumana liketso tsa nalane? Nat Rev Neurosci. 2006; 7: 967-975. [E fetotsoe]
191. Redgrave P, Prescott TJ, Gurney K. Na karabelo ea dopamine e khutšoane haholo e khutšoane haholo ho supa phoso ea moputso? Mekhoa ea li-neuroscience. 1999; 12: 146-151. [E fetotsoe]
192. Rescorla RA, Wagner AR. Khopolo ea boemo ba Pavlovian: ho fapana hoa matlafatso le matlafatso. Ka: Black AH, Prokasy WF, bahlophisi. Boemo ba Classical II: Patlisiso le Khopolo ea Hona Joale. New York, New York: Cropts ea Appleton Century; 1972. maq. 64-99.
193. Reynolds JNJ, Hyland BI, Wickens JR. Mokhoa oa mahlale oa ho ithuta o amanang le moputso. Tlhaho. 2001; 413: 67-70. [E fetotsoe]
194. Reynolds SM, Berridge KC. Tšusumetso e ntle le e mpe khubung ea bokello ea li-nucleus: li-gradients tsa rostrocaudal tse tsoanang bakeng sa ho ja tse etsoang ke GABA, tatso ea "ho rata" / "ho se rate", ho khetha / ho qoba le ho tšaba. J Neurosci. 2002; 22: 7308-7320. [E fetotsoe]
195. Richardson RT, DeLong MR. Lithuto tsa electrophysiological tsa mesebetsi ea li-nucleus basalis ho li-primates. Tsoelo-pele lefapheng la liteko le biology. 1991; 295: 233-252. [E fetotsoe]
196. Robbins TW, Arnsten AF. The neuropsychopharmacology ea ts'ebetso ea molaoli oa fronto: modumo ea monoaminergic. Tlhahlobo ea selemo le selemo ea neuroscience. 2009; 32: 267-287. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
197. Robinson DL, Hermans A, Seipel AT, Wightman RM. Ho beha leihlo puisano ea lik'hemik'hale ka potlako bokong. Litlhahlobo tsa lik'hemik'hale. 2008; 108: 2554-2584. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
198. Roesch MR, Calu DJ, Schoenbaum G. Dopamine neurons e kentse khetho e ntle ho likhoto ho etsa qeto lipakeng tsa meputso e sa liehang kapa e kholo. Nat Neurosci. 2007; 10: 1615-1624. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
199. Roesch MR, Olson CR. Ts'ebetso ea Neuronal e amanang le boleng ba moputso le tšusumetso ho cortex ea pele. Mahlale. 2004; 304: 307-310. [E fetotsoe]
200. Roitman MF, Wheeler RA, Wightman RM, Carelli RM. Likarabo tsa lik'hemik'haleng tsa nako e telele nucleus accumbens li khetholla lintho tse thabisang le tse tsosang takatso. Nat Neurosci. 2008; 11: 1376-1377. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
201. Rutledge RB, Lazzaro SC, Lau B, Myers CE, Gluck MA, Glimcher PW. Lithethefatsi tsa dopaminergic li hlophisa sekhahla sa ho ithuta le mamello ho bakuli ba Parkinson mosebetsing o matla oa ho iphelisa. J Neurosci. 2009; 29: 15104-15114. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
202. Salamone JD. Ho kenya letsoho ha li-nucleus accumbens dopamine molemong oa takatso ea takatso le maikutlo. Patlisiso e mabapi le boitsoaro. 1994; 61: 117-133. [E fetotsoe]
203. Salamone JD, Correa M, Farrar A, Mingote SM. Mesebetsi e amanang le matla ea li-nucleus accumbens dopamine le li-circbrain tse amanang le eona. Psychopharmacology (Berl) 2007; 191: 461-482. [E fetotsoe]
204. Satoh T, Nakai S, Sato T, Kimura M. Correlated coding of susumetsang le sephetho sa qeto ke qeto ea dopamine neurons. J Neurosci. 2003; 23: 9913-9923. [E fetotsoe]
205. Savine AC, Beck SM, Edwards BG, Chiew KS, Braver TS. Ntlafatso ea taolo ea kelello ka ho atamela le ho qoba maemo a susumetsang. Temoho le maikutlo. 2010; 24: 338-356. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
206. Schoenbaum G, Roesch MR, Stalnaker TA, Takahashi YK. Pono e ncha mabapi le karolo ea orbitofrontal cortex ho boiketsi bo sebetsang. Nat Rev Neurosci. 2009; 10: 885-892. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
207. Schultz W. Karabelo ea li-neurop tsa dbamine tsa neuron ho ea boits'oaro bo botle bo susumetsang moeteng. J Neurophysiol. 1986; 56: 1439-1461. [E fetotsoe]
208. Schultz W. Pontšo ea moputso oa ponelo-pele ea li-neurone tsa dopamine. J Neurophysiol. 1998; 80: 1-27. [E fetotsoe]
209. Schultz W. Multiple dopamine e sebetsa ka linako tse fapaneng tsa lithuto. Tlhahlobo ea selemo le selemo ea neuroscience. 2007; 30: 259-288. [E fetotsoe]
210. Schultz W. Dopamine lipontšo bakeng sa boleng ba moputso le kotsi: data ea mantlha le ea morao-rao. Behav Brain Funct. 2010; 6: 24. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
211. Schultz W, Dayan P, Montague PR. Karolo e khutšoane ea ho noha le ho fumana moputso. Mahlale. 1997; 275: 1593-1599. [E fetotsoe]
212. Schultz W, Romo R. Repes of nigrostriatal dopamine neurons to high -ensensity somatosensory stimulation in the monkey anesthetised. J Neurophysiol. 1987; 57: 201-217. [E fetotsoe]
213. Schultz W, Romo R. Dopamine neurons ea monkey midbrain: maemo a tšohanyetso a likarabo ho tsosa maikutlo a boitšoaro bo potlakileng. J Neurophysiol. 1990; 63: 607-624. [E fetotsoe]
214. Seo H, Lee D. Ho fifala ha nakoana hoa lipontšo tsa moputso ka mokokotlong oa dorsal anterior cingrate cortex nakong ea papali ea maano a kopaneng. J Neurosci. 2007; 27: 8366-8377. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
215. Shabel SJ, Janak PH. Ho tšoana ho ts'oanang ho ts'ebetso ea methapo ea amygdala nakong ea takatso ea lijo le maikutlo a sitisang maikutlo. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2009; 106: 15031-15036. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
216. Shadmehr R, Smith MA, Krakauer JW. Khalemelo ea Phoso, Polelo ea Kutloahalo, le Phokotso ea taolo ea makoloi. Tlhahlobo ea selemo le selemo ea neuroscience. 2010 [E fetotsoe]
217. Shen W, Flajolet M, Greengard P, Surmeier DJ. Dichotomous dopaminergic taolo ea striatal synaptic plasticity. Mahlale. 2008; 321: 848-851. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
218. Shimo Y, Wichmann T. Neuronal shughuli ho subthalamic nucleus modulates ho lokolloa ha dopamine ho monkey striatum. Koranta ea Europe ea neuroscience. 2009; 29: 104-113. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
219. Shippenberg TS, Bals-Kubik R, Huber A, Herz A. Neuroanatomical substrates interading theversvers of D-1 dopamine receptor antagonists. Psychopharmacology (Berl) 1991; 103: 209-214. [E fetotsoe]
220. Shumake J, Ilango A, Scheich H, Wetzel W, Ohl FW. Phapang e fapaneng ea li-neuromodulation tsa ho hapa le ho khutlisa thuto ea ho qoba ka sebaka se ka morao sa habenula le sebakeng sa likarolo tse kenang. J Neurosci. 2010; 30: 5876-5883. [E fetotsoe]
221. DM e nyane, Gregory MD, Mak YE, Gitelman D, Mesulam MM, Parrish T. Ho arohana ha boemeli ba neural ba matla le boleng bo ananeloang bathong. Neuron. 2003; 39: 701-711. [E fetotsoe]
222. Stefani MR, Moghaddam B. Ho ithuta le maemo a tšohanyetso ho amana le mekhoa e meng e sa amaneng le ts'ebetso ea dopamine ho "ratort preortal cortex", nucleus accumbens, le dorsal striatum. J Neurosci. 2006; 26: 8810-8818. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
223. Steinfels GF, Heym J, Strecker RE, Jacobs BL. Karabelo ea li-neuron tsa dopaminergic ho katse ho ea litekong tse matla tse hlahisoang ho potoloha boroko. Brain Res. 1983; 277: 150-154. [E fetotsoe]
224. Strecker RE, Jacobs BL. Substantia nigra dopaminergic unit shughuli tsa ho ts'oara likatse: phello ea ho tsosa ha motho a itlotsitse ka boits'oaro le tšebetso ea ho ferekanya kelello. Brain Res. 1985; 361: 339-350. [E fetotsoe]
225. Stuber GD, Hnasko TS, Britt JP, Edward RH, Bonci A. Dopaminergic terminals in the nucleus accumbens but not dorsal striatum coreicela glutamate. Tlaleho ea Neuroscience. 2010; 30: 8229-8233. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
226. Stuber GD, Wightman RM, Carelli RM. Ho felisoa ha ts'ebetso ea ts'ebetso ea cocaine ho senola lipontšo tsa dopaminergic tse sebetsang ka nakoana ho li-nucleus accumbens. Neuron. 2005; 46: 661-669. [E fetotsoe]
227. Sul JH, Kim H, Huh N, Lee D, Jung MW. Likarolo tse fapaneng tsa "rodent orbitof Pambal" le "cortex" ea mantlha pele ho nako ea ho etsa liqeto. Neuron. 2010; 66: 449-460. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
228. Surmeier DJ, Ding J, Letsatsi M, Wang Z, Shen W. D1 le D2 dopamine-receptor modulation of striatal glutamatergic signaling in striatal medium spiny neurons. Mekhoa ea li-neuroscience. 2007; 30: 228-235. [E fetotsoe]
229. Surmeier DJ, Shen W, Letsatsi M, Gertler T, Chan S, Tian X, Plotkin JL. Karolo ea dopamine ho fetoleng sebopeho le tšebetso ea lipotoloho tse potileng. Tsoelo-pele lipatlisisong tsa boko. 2010; 183C: 148-167. [E fetotsoe]
230. Sutton RS, Barto AG. Mabapi le khopolo ea sejoale-joale ea marang-rang a feto-fetohang: litebello le boporofeta. Psychol Rev. 1981; 88: 135-170. [E fetotsoe]
231. Takahashi YK, Roesch MR, Stalnaker TA, Haney RZ, Calu DJ, Taylor AR, Burke KA, Schoenbaum G. Sebaka sa orbitofrontal cortex le ventral tegmental li hlokahala bakeng sa ho ithuta ho tsoa liphellong tse sa lebelloang. Neuron. 2009; 62: 269-280. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
232. Takikawa Y, Kawagoe R, Hikosaka O. Karolo e ka khonehang ea li-neurop tsa dopamine neuron ka nako e khuts'oanyane le ea nako e telele ea li-saccads ho beha limmapa tsa maemo-moputso. J Neurophysiol. 2004; 92: 2520-2529. [E fetotsoe]
233. Tecuapetla F, Patel JC, Xenias H, Senyesemane D, Tadros I, Shah F, Berlin J, Deisseroth K, Rice ME, Tepper JM, Koos T. Glutamatergic signaling ka mesolimbic dopamine neurons ka har'a li-nucleus accumbens. J Neurosci. 2010; 30: 7105-7110. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
234. Thierry AM, Tassin JP, Blanc G, Glowinski J. Khetho ea ts'ebetso ea mesocortical DA system ka khatello ea maikutlo. Tlhaho. 1976; 263: 242-244. [E fetotsoe]
235. Tobler PN, Dickinson A, Schultz W. Coding of Predict Reward Omission ka Dopamine Neurons ho Paramente e Inhibition Paradigm. J Neurosci. 2003; 23: 10402-10410. [E fetotsoe]
236. Tobler PN, Fiorillo CD, Schultz W. Adaptive coding ea boleng ba moputso ka dopamine neurons. Mahlale. 2005; 307: 1642-1645. [E fetotsoe]
237. Tsai HC, Zhang F, Adamantidis A, Stuber GD, Bonci A, de Lecea L, Deisseroth K. Phasic Firing in Dopaminergic Neurons E Lekane bakeng sa Boemo ba ho Ts'oara. Mahlale. 2009 [E fetotsoe]
238. Ullsperger M. liphatsa tsa lefutso tsa lipatlisiso tsa tlhahlobo ea ts'ebetso le ho ithuta ho tsoa phihlelong: karolo ea dopamine le serotonin. Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2010; 34: 649-659. [E fetotsoe]
239. Nts'a MA. Dopamine: bothata bo ka sehloohong. Mekhoa ea li-neuroscience. 2004; 27: 702-706. [E fetotsoe]
240. Uylings HB, Groenewegen HJ, Kolb B. Na likhoto li na le cortex ea pele? Patlisiso e mabapi le boitsoaro. 2003; 146: 3-17. [E fetotsoe]
241. Ventura R, Cabib S, Puglisi-Allegra S. Karabo e arohaneng ea genotype e itšetlehileng ka mesocorticolimbic dopamine ka khatello ea maikutlo. Neuroscience. 2001; 104: 627-631. [E fetotsoe]
242. Ventura R, Latagliata EC, Morrone C, La Mela I, Puglisi-Allegra S. Prefrontal norepinephrine e khetholla tšebetso ea "matla" a ho susumetsa a matla. LITEKOANE ONE. 2008; 3: e3044. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
243. Ventura R, Morrone C, Puglisi-Allegra S. Sistimi ea pele ea khekhe ea litheko le litheko li khetholla tšusumetso ea sepheo sa hau sa ho susumetsa ka bobeli meputso- le sepheo se amanang le ho hlohlona. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2007; 104: 5181-5186. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
244. Vogt BA. Tšebelisano e bohloko le ea maikutlo ka likarolo tsa mofuta oa gingase. Nat Rev Neurosci. 2005; 6: 533-544. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
245. Voon V, Pessiglione M, Brezing C, Gallea C, Fernandez HH, Dolan RJ, Hallett M. Mechanisms e thehiloeng tlas'a moputso oa dopamine bohareng ba boitšoaro bo hatelletseng. Neuron. 2010; 65: 135-142. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
246. Waelti P, Dickinson A, likarabo tsa Schultz W. Dopamine li latela lintlha tsa mantlha tsa thuto ea molao ea thuto. Tlhaho. 2001; 412: 43-48. [E fetotsoe]
247. Wallis JD, Kennerley SW. Lipontšo tsa moputso oa Heterogenible ho cortex ea pele. Maikutlo a hona joale ho neurobiology. 2010; 20: 191-198. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
248. Walton ME, Behrens TE, Buckley MJ, Rudebeck PH, Rushworth MF. Lisebelisoa tse arohaneng tsa ho ithuta bokong ba macaque le karolo ea cortex ea orbitofrontal ho ithuteng e le phehisano. Neuron. 2010; 65: 927-939. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
249. Wheeler RA, Twining RC, Jones JL, Slater JM, Grigson PS, Carelli RM. Boitšoaro bo bobe le boitshwaro bo bobe bo amang matla a boitšoaro bo bobe bo ka ama bokamoso ba ho itaola. Neuron. 2008; 57: 774-785. [E fetotsoe]
250. Wightman RM, Heien MLAV, Wassum KM, Sombers LA, Aragona BJ, Khan AS, Ariansen JL, Cheer JF, Phillips PE, Carelli RM. Ho lokolloa ha dopamine ho atile haholo kahare ho lintho tse nyane tse sebetsanang le li-nucleus tsa "rat". Koranta ea Europe ea neuroscience. 2007; 26: 2046-2054. [E fetotsoe]
251. Williams GV, Castner SA. Tlas'a kotopo: mathata a bohlokoa a ho hlakisa tšebetso ea li-receptor tsa D1 memori e sebetsang. Neuroscience. 2006; 139: 263-276. [E fetotsoe]
252. Williams SM, Goldman-Rakic ​​PS. Tšimoloho e pharaletseng ea sistimi ea mesoprontal dopamine e atileng. Cereb Cortex. 1998; 8: 321-345. [E fetotsoe]
253. Winn P. Tsela e molemohali ea ho lekola sebopeho le tšebetso ea sekhahla sa pedunculopontine tegmental nucleus: bopaki bo tsoang lithutong tsa liphoofolo. Tlaleho ea Saense ea Neurological. 2006; 248: 234-250. [E fetotsoe]
254. RA ea bohlale. Dopamine, ho ithuta le tšusumetso. Nat Rev Neurosci. 2004; 5: 483-494. [E fetotsoe]
255. RA ea bohlale. Sebelisa likarolo tse ling tsa moputso le tšusumetso. Lingoloa tsa neurology e bapisang. 2005; 493: 115-121. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
256. Bohlale SP. Masimo a ka pele: phylogeny le ts'ebetso ea mantlha. Mekhoa ea li-neuroscience. 2008; 31: 599-608. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
257. Yamada H, Matsumoto N, Kimura M. Li-neuron tse sebetsang ka mokhoa o hlakileng ka bokong ba "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" "" ". J Neurosci. 2004; 24: 3500-3510. [E fetotsoe]
258. Yamada H, Matsumoto N, Kimura M. Nalane- le litekanyetso tsa morao-rao tse thehiloeng ho litekanyetso tsa litlamorao tsa boits'oaro bo botle lithutong tsa morao-rao. J Neurophysiol. 2007; 98: 3557-3567. [E fetotsoe]
259. Yin HH, Knowlton BJ. Karolo ea sehlopha sa basal ganglia ho theheng tloaelo. Nat Rev Neurosci. 2006; 7: 464-476. [E fetotsoe]
260. MOTHO e monyane, Moran PM, Joseph MH. Karolo ea dopamine maemong le lithibelo tsa morao-rao: keng, neng, neng, kae, joang? Litlhahlobo tsa bophelo ba morao-rao ka morao-rao. 2005; 29: 963-976. [E fetotsoe]
261. Zaghloul KA, Blanco JA, Weidemann CT, McGill K, Jaggi JL, Baltuch GH, Kahana MJ. Human substantia nigra neurons e kenyelletsa moputso o sa lebelloang oa lichelete. Mahlale. 2009; 323: 1496-1499. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
262. Zahniser NR, Sorkin A. Melao e potlakileng ea sephethephethe sa dopamine: karolo ea bokhoba bo matlafatsang? Neuropharmacology. 2004; 47 Suppl 1: 80-91. [E fetotsoe]
263. Zhang L, Doyon WM, Clark JJ, Phillips PE, Dani JA. Taolo ea phetiso ea tonic le phasic dopamine phetisong ea dorsal le ventral striatum. Thutela ea limolek'hule. 2009; 76: 396-404. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
264. Zink CF, Pagnoni G, Martin ME, Dhamala M, Berns GS. Karabelo e potlakileng ea motho litlamong tse ling tse sa fetoheng. Tlaleho ea Neuroscience. 2003; 23: 8092-8097. [E fetotsoe]
265. Zweifel LS, Argilli E, Bonci A, Palmiter RD. Karolo ea li-receptor tsa NMDA ho li-dopamine neurons bakeng sa boits'oaro ba semmuso. Neuron. 2008; 59: 486-496. [Tlhahiso ea mahala ea PMC] [E fetotsoe]
266. Zweifel LS, Parker JG, Lobb CJ, Metsi a pula A, Wall VZ, Fadok JP, Darvas M, Kim MJ, Mizumori SJ, Paladini CA, et al. Ho fokotseha ha ho phatloha ho thehiloeng ke NMDAR ho tsebisoang ke dopamine neuron ho fana ka tlhahlobo e khethiloeng ea boitšoaro bo itšetlehileng ka phasic dopamine. Ts'ebetso ea Setsi sa Naha sa Mahlale sa United States of America. 2009; 106: 7281-7288. [Tlhahiso ea mahala ea PMC] [E fetotsoe]