Front Psychiatry. 2011 Jul 6; 2: 37. Doi: 10.3389 / fpsyt.2011.00037. eCollection 2011.
Helena JV Rutherford1*, Sarah K. Williams2*, Sheryl Moy2,3, Linda C. Mayes1 'me Josephine M. Johns2,3
- 1 Setsi sa ho Ithuta sa Bana sa Yale, Univesithi ea Yale, New Haven, CT, USA
- 2 Lefapha la Psychiatry, Univesithi ea North Carolina-Chapel Hill, Chapel Hill, NC, USA
- 3 Setsi sa Carolina sa Bokooa ba Ntlafatso, University of North Carolina-Chapel Hill, Chapel Hill, NC, USA
Tlatsetso e emetse tšebelisano e rarahaneng lipakeng tsa moputso le lipotoloho tsa neural tse nang le khatello ea maikutlo, le ts'ebeliso e matla ea lithethefatsi e bontšang phetoho ho tloha ho matlafatso e nepahetseng ho ea ho tse ling tse matlafatsang ts'ebetsong ho boloka ts'ebeliso ea lithethefatsi. Boithuto ba pele-pele bo bontšitse ho nka karolo ha libaka ka har'a amygdala e le ho boloka phetoho ena, haholoholo tlasa maemo a sithabetsang. Maemong a ho lemalla, sistimi ea moputso e sebeletsa ho boloka boits'oaro bo amanang le phokotso ea litšusumetso tse mpe, ka litšenyehelo tsa ho fumana boteng ba meputso e meng. Ka hona, temallo e bonts'a ho hloka taolo lipakeng tsa lits'ebetso tsa moputso oa mantlha, ho kenyelletsa sebaka sa pele (CC), sebaka sa karolelano ea sepheo (VPS), le li-axus tsa nucleus (NAc), hammoho le "hypothalamic-pituitary-adrenal axis" le "amhergdala" ea ts'ebetso ea khatello ea maikutlo. . Mona, re nahana ka litlamorao tsa liphetoho tsa ts'ebetso ea neural nakong ea kapa ho latela ts'ebeliso ea tlhekefetso ea botsoali, ts'ebetso e atlehileng e ka sitisoang ke bokhoba. Ka ho khetheha, re hlakisa lithuto tsa pele le tsa batho tse tšehetsang tšebeliso ea litheko tsa meputso le khatello ea maikutlo ka tlhekefetso le tlatsetso ea litsamaiso tsena ho botsoali. Bopaki bo ntseng bo eketseha bo bonts'a karolo ea bohlokoa bakeng sa hypothalamus, PFC, VTA, le NAc ho ba motsoali, ka libaka tse tšoanang e le tsona tse sa sebetseng ka ho kenella bokhobeng. Ho feta moo, ho batho ba baholo ba lemaletseng, re fana ka maikutlo a hore lintlha tsa ho ba motsoali li ka baka khatello ea maikutlo hape ho e-na le ho putsa, 'me sena se ka eketsa maemo a mabe, sa etsa hore ho be le boits'oaro bo bobe ba ho ba le monyetla oa ho lahla ngoana le ho hlekefetsoa.
Bokhoba ba tlhekefetso bo hlalositsoe e le mohato o potileng oa ho sitisoa ha melao ea tsamaiso. Ka bobeli mekhoa e metle ea ho matlafatsa e ka kenya letsoho ho bolokeng tšebeliso ea lithethefatsi; eo pele e neng e emela karabo ea moputso kamora ts'ebeliso ea pele, baemeli ba morao ba ne ba tsoela pele ho sebelisa ts'ebetso ea ho imolla boemo bo bobe ba ho tsieleha. Boemong ba methapo ea kutlo, ha ts'ebetso ea ts'ebetso ea moputso oa boko e hatisa ts'ebeliso ea lithethefatsi (ke hore, ts'ebetso e ntle), ts'ebetso ea ts'ebetso ea khatello ea kelello e ka laola mahlomola a amanang le ho tlohela (ke hore, ho matlafatsa hampe). Boemong ba bokhoba, phomolo ea khatello ea maikutlo ke moriana o lebisa tšebeliso e tloaetsoeng ea lithethefatsi, 'me sistimi ea meputso e "khethoa" ka sepheo sa ho boloka boits'oaro bo amanang le phomolo ea khatello ea maikutlo kapa maikutlo a fosahetseng. Ka kopanelo ena, meputso e meng e tlisoang ha e feto-fetoha hobane e se karolo ea sehokelo sa boemo ba moputso. Taba ea bohlokoa ke hore, meputso ena e kenyelletsa botsoalle sechabeng le likamano tsa batho, 'me mofuta ona oa kopanelo o na le litlamorao tse kholo ho boits'oaro ba botsoali hara batho ba baholo ba lemaletseng, hape le likamanong tsa bona ka kakaretso. Maemong ana, likamano li ka sithabetsa le ho feta ho motho ea lemaletseng lithethefatsi ka lebaka la litlhoko tsa tlhokomelo. Kahoo, ho fapana le moputso o tloaelehileng o fanoang ke kopanelo, kamano e ba e sithabetsang haholo 'me e sebetsa e le sesupo sa boitšoaro bo tsoelang pele bo qobelloang kapa ba bokhoba ha bo ntse bo ntšetsa pele boitšoaro bo botle ba thobalano kapa boimana.
Tokomaneng ena, re tla supa bopaki ho tsoa lithutong tsa pele le tsa batho ho rarolla litlhahiso tse latelang tse hlalositsoeng ho Setšoantšo. 1: (a) Lits'ebetso tsa tlatsetso ke sesupo sa ho thella ha tekanyo lipakeng tsa meputso le karabelo ea khatello; (b) ho ba motsoali ho kenyelletsa tloaelo e ikhethang ea moputso le taolo ea khatello ea maikutlo ho litloholo tse amehang tse tlohang ho motho e moholo, eo hona joale e leng motsoali; le (c) maemong a tlatsetsang, litloaelo tsa botsoali ha li na moputso empa li na le khatello ea maikutlo. Karabelo ena e matlafatsang ea khatello ea maikutlo e ka khothaletsa boitšoaro bo batlang lithethefatsi ho fapana le boits'oaro ba botsoali le ho hlokomela litlhoko tsa lesea. Re shebana le ts'ebeliso ea koae ea koae ho hlahloba bopaki ba mohlala ona.
Tšoantšiso 1
Setšoantšo sa 1. Kamano lipakeng tsa bokhoba ba ho lemalla le ho ba motsoali. Mohlaleng oa rona, bokhoba ba taolo bo emela khatello ea maikutlo le lits'enyehelo tsa moputso, tseo ka bobeli li fetohang ho tšehetsa motsoali. Boemong ba ho lemalla, re fana ka maikutlo a hore liphetoho tse fetotsoeng ke ts'ebeliso ea lithethefatsi li etsa hore boleng ba masea bo ntlafatsoe, 'me bo nkeloe karabelo e hatellehang ea methapo ea kutlo. Karabelo ena ea khatello ea maikutlo mekhoeng ea masea e ka holisa takatso ea lithethefatsi tsa tlhekefetso, ho khothaletsa ho batla lithethefatsi le ho khutlela ho bo-mme ba sa tsotelloeng, 'me ho ntšetsa pele mokoloko oa ho se tsotelle.
Cocaine e Senya Boitšoaro ba Boimana: Bopaki bo tsoang ho Rentent le Lintlha tsa Botho
Ts'ebeliso ea koae le tlhekefetso e emela bothata bo boholo ba bophelo bo botle ba sechaba. Ho 2008, Maamerika a limilione tse 1.4 a ile a kopana le litekanyetso tsa DSM-IV tsa ts'ebeliso e mpe ea lithethefatsi le ho itšepa ho lemalla joala ba tsona ba koae.Ts'ebeliso e Mpe ea Ts'ebeliso e Mpe ea Ts'ebeliso e Mpe ea Ts'ebeliso ea Litšebeletso tsa Bophelo ba kelello, 2008). Palo ea tšebeliso ea koae ho basali ba banyane e eketsehile, 'me liphuputso tse' maloa li tlaleha tšebeliso ea koae nakong ea kemolo (Kuczkowski, 2004). Ts'ebeliso ea "cocaine" e tsoelang pele nakong ea pelehi ha ea tloaeleha ebile e hlahisa mathata a bohlokoa bakeng sa tloaelo ea ho ba motsoali; ka kotloloho, tlhekefetso ea basali ea lithethefatsi e amahanngoa le keketseho e kholo ea tlhokomeliso ea bana (Cash le Wilke, 2003). Ho rarahana ha ho ithuta litlamorao tsa koae ho batho ba ho ba le bana maemong a boits'oaro le methapo ea kutlo ho kopantsoe ke bophahamo ba maikutlo le likokoana-hloko tse amanang le tšebeliso ea koae, ka hona, li hlahisa liphephetso tse ikhethang lipatlisisong tse matla. Ka hona, re nka mokhoa o tlatselletsang oa ho koala mosebetsi ho tsoa mahlale a motheo le lithuto tsa batho. Le ha mefuta e mengata ea liphoofolo tse anyesang e bonts'a mofuta o itseng oa tlhokomelo ea botsoali ho bana ba bona, litoeba e ba mohlala o motle bakeng sa ho ithuta le ho hlokomela boits'oaro ba bakhachane (MB) hobane, ho tšoana le batho, ba hlahisa masea a hlokang maiketsetso a hlokang boitlamo bo boholo ho netefatsa pholoho. 'Mangoane oa "rat" o qeta letsatsi lohle a na le malinyane bakeng sa libeke tsa 2 kamora ho hlaha, a siea sehlaha feela hore a fete lijo. Liroto li bonts'a boitšoaro bo makatsang ho masea (malinyane) a ka hlalosoang mme a ba le mekhoa ea boits'oaro ho batho, ho kenyelletsa ho anyesa le ho koetlisa lesea hape le ho lokisa tikoloho e sireletsehileng bakeng sa lesea (moaho oa sehlaha). Khalemelo ea 'm'ae kapa ts'ireletso e boetse e hlaha nakong ea pelehi, boitsoaro bo khahlisang “boiphihlelo” bo fumanoeng ke bo-mme ba batho. Re qala pampiri ena ka ho ts'ehetsa ts'ehetso e matla ea ts'oaetso ea koae ho li-MB lithutong tsa batho le litoeba.
Lithutong tsa batho, litlamorao tsa ho pepesetsoa ha koae le taolo ea ho ba motsoali li sibollotsoe haholo ka ho shebella ho sebelisana ha 'm'a le ngoana. Nakong ea 12-48 h kamorao ho nako, bo-mme ba neng ba sebelisa koae nakong ea kemolo ba ile ba araba ka mokhoa o ikhethileng mme ba ne ba khesoa ho masea a bona a macha ha ba bapisoa le bo-mme ba se nang lithethefatsi (Gottwald le Thurman, 1994). Mosebetsi oa pele ho bana o ile oa fana ka tlhahiso ea hore bo-'mè ba sebelisang koae nakong ea kemolo ba fane ka bopaki ba tšusumetso e ntle le kutloelo-bohloko ho lithahasello tsa masea, le bokhoni ba bofutsana le boiphihlelo nakong ea litšebelisano tsa dyadic (Burns et al., 1991, 1997). Ho na le bopaki bo bonts'ang hore le ha mathata a pelehi (mohlala, a fokotsa tlhokomelo ho lesea, a shebana le lesea) litšebelisanong tsa 'm'a le ngoana a bonahala a fetotsoe ke ho pepesetsoa ha koae, ho latela tlhahlobo e bonts'itse phokotso (Mayes et al., 1997) kapa ho ba sieo ha lits'ebetso tse tšoanangBall et al., 1997) ho fana ka maikutlo a litlamorao tsa ho pepesetsoa koae ho ka fetoha ha nako e ntse e tsamaea. Sena se entse hore lithuto li hlahlobe litlamorao tsa ho pepesetsoa ha koae nakong ea ts'ebelisano ea 'm'ae ho masea le bona, ho paka tšebelisano e mpe ea litheko tse mpe tsa lehloeo la cocaine (Johnson et al., 2002; Uhlhorn et al., 2005; Molitor le Mayes, 2010). Litlamorao tsa tšebeliso ea koae nakong ea kemolo le tsona li fuputsoe nakong ea lijo. Ho ile ha tlalehoa litšebelisano tse futsanehileng ho bo-'mè ba neng ba sebelisa koae nakong ea kemolo le ba ileng ba khutlela morao kamora ho bapisoa le bo-mme ba neng ba sa khutla.Blackwell et al., 1998). Mefuta ena ea phepo e bontšitse khohlano e kholo (Eiden, 2001) hammoho le ho hloka botsitso (Eiden et al., 2006) pakeng tsa mme le ngoana. Ho bohlokoa ho hlokomela hore mathata a ts'ebetsong ea 'm'a le ngoana a ka amana le palo ea koae e sebelisoang nakong ea kemolo (Tronick et al., 2005), hammoho le tšebeliso ea kamora 'matsatsi (Blackwell et al., 1998; Johnson et al., 2002; Eiden et al., 2006). Le ha ho le joale ho se liphuputso tse phatlalalitsoeng tsa bo-'mampuli ba lemaletseng koae, hona joale ho na le lipatlisiso tsa maiketsetso tse bontšang phapang e tlang pele ho basali ba basali bana ha ba bapisoa le bo-'mè ba se nang nalane ea tšebeliso ea koae.Strathearn le Kosten, 2008).
Boithuto mabapi le matla a bo-'m'a lesea ka penteng e bonts'a hore ho pepesehela k'hok'heine ka lebaka la kalafo e matla, ea nakoana kapa ea kalafo e sa foleng, e sitisa likarolo tsa MB, moo boemo ba pherekano bo itšetlehileng ka tekanyetso, nako, tlhahlobo, le mofuta oa kalafo.Johns et al., 1994; Nelson et al., 1998). Ho "paradigm" e le 'ngoe e sebelisoang hangata, matamo a fumana koae ka linako tse ling matsatsing a ho emara (GD 1-20; 30 mg / kg) kapa ka ente e le' ngoe nakong ea kamora 'mele, ebe o ea hlahlojoa bakeng sa MB e laetsoeng ho latela karohano le ho kopana hape le malinyane. Mekhoa e meng le e 'ngoe e ka lebisa ho ho eketseheng ha nako le ho fokotsa nako ea booki, hammoho le ho sitisoa ha lick le moaho oa matlo. Matamo a phekotsoeng a Cocaine a boetse a bonts'a boitšoaro bo bobe ba ho ba makhopho a bo-mme a bonts'a hore likhaello tsa boitšoaro ba sechaba li ka holisa likamano tsa malinyane tse fetileng (Johns et al., 1997b; McMurray et al., 2008) .Linyeho tsena ha li hlahisoe ke ho hema kapa ho khaotsa koae (Johns et al., 1997b), e fana ka tlhahiso ea liphetoho lipakeng tsa litšebelisano tse khothatsang kapa tsa setsoalleng.
Ho fapana le kalafo nakong ea ho emara, ho pepesehela cocaine khafetsa pele ho ima ho eketsa ho khutla le boitšoaro bo hohelang lithetleng le litlohong tsa pele tsa mpa, ho fana ka maikutlo a hore ho pepesehela k'hok'heine ea batho ba baholo e fetoha sesosa sa ho hlohlona le boits'oaro bo tlang kamora nako, ka tsela ea tlhaho e khothatsang, joalo ka malinyane.Nephew le Febo, 2010). Le ha ho le joalo, le ha bo-'mè bana ba ne ba potlakela ho fumana malinyane, ba ile ba nka nako e telele ho qala li-MB tse ling, ba fana ka maikutlo a hore boteng ba tsona bo ka ba ka ho etsa hore malinyane a haufi empa a se ketsong ea ho a hlokomela. Mokhoa o sebetsang oa matla a matla a matla a khoheli (fMRI) ho liphoofolo tse falimehileng o lumelletse lipatlisiso tsa ts'ebetso ea libaka tsa boko kamora ho pepesehela tlhotlheletso e khotsofatsang. Kannete, ho pepesetsoa hoa koae ho boetse ho fokolitse ts'ebetso ea ho anyesa ngoana ka mokokotlong oa makhopho a methapo ea pele (mPFC), striatum, le auditory cortex, empa ha e a ka ea ama baster dopamine (DA) kapa keketseho ea liperesente tsa DA ha e pepesehela malinyane mPFC (Febo le Ferris, 2007), ho ts'ehetsa likarolo tsa bohlokoa ho MB le mekhoa ea bokhoba ba litereke tseo re nang le tsona, ke ntlha eo re tla khutlela ho eona bohatisong bona.
Ha li kopantsoe hammoho, liphumano tsena li bonts'a ts'ebeliso ea koae pele, nakong, le / kapa kamora ho ima e ka fetola haholo boemo ba pelehi. Kaha MB ha e felisoe ka botlalo, re fana ka maikutlo a hore liphetoho tsena tsa boits'oaro li ka bonts'a phapang mokhoeng oa moputso oa bana. Ho feta moo, phetoho ea ho tloha moimana ho ea pelehi e sithabetsa haholo ho bo-mme; leha ho le joalo, ho ikamahanya hantle le tikoloho ena e ncha ho ka nahanoa tlasa taolo ea methati e matla ka ho fetesisa. Allostasis e hlalositsoe e le ts'ebetso e sebetsang ea ho araba liphephetso tse tsoang tikolohong ho boloka homeostasis, hangata ka ts'ebetso ea likarabo tsa khatello ea maikutlo tsa li-hormone. Ho ikamahanya ka katleho le tikoloho ena e sithabetsang le bokhoni ba ho arabela litlhoko tsa lesea ka nepo ho bohlokoa bakeng sa bophelo ba lesea mme ka hona e bolokiloe ho iphetola ha lintho ka popo ea liphoofolo. Ho reriloe hore ho lemalla lithethefatsi ho ka senya tloaelo e tloaelehileng maemong a sithabetsang ao e seng a botsoali (Le Moal, 2009); leha ho le joalo, hore na ho joalo le ka nako ea pelehi e ntse e sa hlaka ebile e fana ka tlhaloso e ka bang bakeng sa bofokoli bo susumetsoang ke lithethefatsi bo hlalositsoeng ka holimo. Tlhahlobo e hlahisitsoeng mona e shebisisa ho kenya letsoho ha methapo ea neural tsamaisong ea moputso le ea khatello ea maikutlo molemong oa ho ba motsoali (e hlalositsoeng ho Setšoantšo. 2).
Tšoantšiso 2
Setšoantšo sa 2. Kamano lipakeng tsa methapo ea kutlo ea khatello ea maikutlo, moputso le botsoali. Rekethe ea botsoali (bofubelu) e arolelana libaka tse ngata ka khatello ea maikutlo (boputsoa) le moputso (o mosehla). Likarolo tse thathamisitsoeng setsing li bile le tšusumetso makaleng ohle a mararo, li fana ka maikutlo a hore pherekano libakeng tse potoloho e le 'ngoe e ka ba le tšusumetso e kholo ts'ebetsong ea lipotoloho tse ling tse hokahaneng. E ka bonoa ka palo ea libaka tse kenyellelitsoeng ho potoloho ea Botsoali hore ts'ebetso ea tlhokomelo e nepahetseng ea botsoali (le mefuta e mengata ea boitšoaro e oelang sehlopheng sena) e hloka ts'ebetso e tloaelehileng ea boholo ba boko. Ho kenya mebala letlalong ho bontša lits'ebetso tsa boko bo sebakeng seo karolo e 'ngoe le e' ngoe e leng ho sona.
Ho qala ts'ebetso ea bo-mme
Ntho ea bohlokoa ho tlhahlobo ena ke maikutlo a hore ho na le liphetoho tsa bohlokoa tsa neurobiological tse fanang ka phetoho ea botsoali. Tsena li kenyelletsa liphetoho tse fapaneng tsa sebopeho le methapo e bonts'a polasetiki ka bobeli maemong a taolo ea synaptic le transcriptal. Ho pholletsa le pampiri ena re bua ka liphetoho mesebetsing ea li-neuronal, polelo ea li-receptor kapa boemo ba peptide, ka lebaka la ho tsoa ha bo-mme kapa ho pepesehela lithethefatsi, joalo ka polasetiki ho bonts'a sebopeho sa "neurons" sa tlhaho. Bohlokoa phetohong ea ho ba bo-mme ke liphetoho tse kholo tšebetsong ea hypothalamus le tlhahiso ea neuropeptide oxetocin (OT), ka hona re qala ho nahana ka ho qalisoa ha MB le ho kenella ha hypothalamus le oxytocin pele ho hlahloba khatello le moputso oa neuraltocin potoloho le tloaelo ea tsona ea ho ba motsoali.
Hypothalamus
Sebaka sa medial preoptic (MPOA) le "ventral bed nucleus" ea stria terminalis (BNST) li bohlokoa bakeng sa ho qala le ho boloka MB, mme e emela sebaka se tataisang se fetolang phetoho ea boits'oaro ba botsoali (Numan, 2007). MPOA e na le khokahano e tobileng sebakeng sa "ventral tegmental sebakeng" (VTA), li-nucleus accumbens (NAc), mPFC, BNST le paraventricular nucleus (PVN), e e lumella hore e fane ka tšusumetso e matla nakong ea pelehi. Litho ho MPOA li felisa ho khutla le boits'oaro ba ho aha sehlaha litsing. Mooki o ntse a bonoa ka ho latela liso, leha e fokotsehile, mme e kanna ea ba sephetho sa malinyane a batlang le ho ts'oara letamo le sa arabeng. Liphetho tsena li bonts'a hore MPOA e bohlokoa bakeng sa liketso tsa ts'usumetso ho MB (Numan, 2007). Ho eketsa ts'ebetso ea methapo ea MPOA ea MPOA, joalo ka ha e lekantsoe ke c-FOS, CREB, le fMRI, e bonoa kamora ho pepeseha ha malinyane le likotsi tsa malinyane libekeng tsa pele tsa pele tsa 2 (Fleming le Korsmit, 1996; Febo et al., 2005; Jin et al., 2005). Haholo-holo, khaello ea CREB mefuteng ea phetoho ea panya e lebisa ho eketseho ha batho ba shoang le liphokotso tse kholo ho lateleng malinyane le palo ea malinyane a khutliselitsoeng sehlaheng (Jin et al., 2005). DA le serotonin (5-HT) li boloka maemo a motheo tikolohong ena ho pholletsa le kemolo, leha ho le joalo norepinephrine (NE) e fokotsehile nakong ena (Olazabal et al., 2004). Ka kakaretso, liphetho li bontša hore ts'ebetso ea CREB ho MPOA e ka ba bohlokoa haholo ho qala le ho hlahisa MB. Ho buisana le sena ho bo-'mè ba batho, phapang e fapaneng pakeng tsa maikutlo a bo-mme le bokhoni ba masea e senotse modumo e fapaneng ea hypothalamus, le libaka tsa pituitary le tsa PFC, ha bo-mme ba ntse ba shebile lifoto tsa masea a bona, bapisoa le masea a sa tsejoeng (Strathearn et al., 2009). Ho feta moo, thutong ena ea morao-rao, karabelo ea OT ea 'm'ae kamora tšebelisano e amanang le ts'ebetso ho hypothalamus (hammoho le pituitary le ventral striatum).
oxytocin
Oxetocin e bapala karolo ea mantlha ho qala ho qaleha ha MB liphoofolong tse 'maloa tsa liphoofolo tse anyesang, ho kenyelletsa rat le motho, empa karolo ea eona ea ho hlokomela MB ha e hlake haholo (Pedersen le Boccia, 2002; Feldman et al., 2007). Lehloeng, li-neuron tsa OT tse tsoang morerong oa PVN ho tloha bohareng ho ea ho olbatori ea bulb (MOB), MPOA, NAc, amygdala (AMY), hippocampus le VTA (bona Setšoantšo. 3). OT e tsoang morerong oa PVN le supraoptic nucleus (SON) ho ea sebakeng sa pituitary bakeng sa ho phunyeletsoa ka mothapelong oa mali ka lebaka la ts'usumetso e hlahisoang ke masea kapa e sithabetsang (Uvnas-Moberg et al., 2005; Hatton le Wang, 2008). Mosebetsi o sebelisang mela ea litoeba bakeng sa OT o bonts'itse hore OT e bohlokoa bakeng sa ho pholoha ha matlakala ka lebaka la litlamorao ho lesela la lebese, empa e kanna ea se be ea bohlokoa bakeng sa likarolo tse ling tsa likarabo tsa bo-mme.Lee le al., 2009). Le ha li-mutts tsa bana tsa OT li na le likarabelo tse tloaelehileng tsa ho ikatisa le ho holisa (Ferguson et al., 2000, 2001), matamo a mouse a nang le ts'oaetso e lebisitsoeng ea mofuta oa OT receptor a na le bofokoli bo boholo ho khutlisetsoeng popo, ho kenyelletsa le bolelele bo bolelele ba ho khutlisa ngoana ka mong le ho nahana ka boemo bo patehileng, le nako e khuts'oane e qetile e petetsane holim'a malinyane (Takayanagi et al., 2005). Ntle le moo, bo-'mè ba batho ba nang le ts'ebetso e tlase ea OT e sebetsang e tlase ba bonts'a kutloisiso e tlase ea bo-mme nakong ea pelehi (Bakermans-Kranenburg le van Ijzendoorn, 2008). Liphetoho tšebetsong ea OT li khothalelitsoe ho pepesa liphokojoe tsa bo-mme ho litoeba le phetoho ho Peg3 (Paternally e bontšitse gene 3). Mosali Peg3 litoeba tse feto-fetohang li na le mathata a tebileng ho MB le ho hloka lebese le fokolang (Champagne et al., 2009). Lits'oaetso tsena li hokahane le phokotso ho li-neurons tsa OT ho hypothalamus mme li fokotse ho tlama ha OT ho MPOA le septum ea morao-rao ho Peg3 mutants (Champagne et al., 2009).
Tšoantšiso 3
Setšoantšo sa 3. Projeke ea Oxetocinergic bokong ba methapo. Ho lokolloa ha Oxetocin ho loketse ka mokhoa o hlakileng ho fana ka maikutlo a moputso, khatello le lits'oaetso tsa bo mme. Li-neurocellular neurons tsa PVN (oval e khubelu) le SON (oval e putsoa) ho morero oa morao-rao ho lokolla OT ka mokhoa o ts'oanang ha o arabela molemong oa ho anyesa kapa o sithabetsang. Li-neurons tsa parvocellular tsa PVN li boetse li kenella ho li-circry tsa moputso (VTA le NAc), khatello ea khatello (hippocampus, AMY, le tokollo ea intra-PVN), le potoloho ea basali (MPOA / BNST le OB), 'me ho lumeloa hore e bohlokoa bakeng sa litšebelisano tse nepahetseng tsa sechaba. PVN, nucleus ea li-hypothalamus; SON, nucleus ea superthalamus; VTA, sebaka sa ho kenella hare; NAc, li-bokelle tsa nucleus; AMY, amygdala; MPOA, sebaka sa kalafo se atamelaneng sa hypothalamus; OB, tleloubu ea ho etsa lintho. Brain schematic e fetotsoe ho tloha Paxinos le Watson (1997).
Pampiri ea moraorao ke Yoshida et al. (2009) o khothalelitse hore mochine o ka sehloohong oa litlamorao tsa OT litloaelanong tsa bophelo le matšoenyeho o ntlafalitsoe ka serotonergic neurotransmission. Tsamaiso e bohareng ea OT ho "median raphe" e lebisitse ho lokollotsoeng ea 5-HT ka litoeba (Yoshida et al., 2009). Tloso ea 5-HT1A or 1B receptor e ka fetola nako e sebelisitsoeng e patetse sehlaheng, ho khutlisetsa popo kapa mehato e meng ea ho holisa, 'me e ka fetola litlamorao ho boits'oaro ba boitšoaro ba bana (van Velzen le Toth, 2010). Liphetoho tsa transporter ea serotonin (5-HTT) li ka etsa hore menyetla ea ho ba le mathata a ho ba le bana le mathata a mang a tikoloho e be le litlamorao tsa nako e telele tsa ho mamella maemo a mabe (Bakermans-Kranenburg le van Ijzendoorn, 2008; Kinnally et al., 2009; Heiming le Sachser, 2010). Matla a ntlafatsang serotonergic neurotransication ke OT a ka ba le katleho e ts'oanang taolong ea ts'ebetso ea CREB e amanang le ho arabela khatello ea maikutlo, ho pepesetsoa koae le lithethefatsi tse ling, le ho MB. Litšebelisano tse khutlang lipakeng tsa OT le 5-HT li-signways litseleng tsa midbrain le hypothalamus li ka senyeha haholo ka mor'a ts'ebeliso ea lithethefatsi mme li ka kenya letsoho khaellong ea MB.
Ka bobeli plasma le boko OT e ka sebelisana le hypothalamic-pituitary-adrenal (HPA) axis (Uvnas-Moberg et al., 2005; Slattery le Neumann, 2008). Tlhahisoleseling ena e kanna ea ama lipatlisiso tsa bongaka ba batho, joalo ka ha phuputso e 'ngoe e tlaleha hore bo-mme ba batho ba sebelisang koae nakong ea boimana ba fokotse maemo a OT ea plasma le khatello e kholo ea kelello (Leseli et al., 2004). Sistimi ea OT e sitisoa ke koae libakeng tse 'maloa, ka ho tšoana le tšitiso ea boitšoaro ea MB (Johns et al., 1997a, 2005). Mohlala, kalafo e sa foleng ea matamo a rat le cocaine e ka fokotsa maemo a OT haholo ho MPOA, hippocampus, le VTA ka har'a 24 h ea ho tsamaisoa (Johns et al., 1997a). Acoca cocaine nakong ea pelehi e ka ama maemo a OT le li-receptor tsa OT libakeng tse 'maloa tsa boko, ho kenyelletsa le maemo a tlase ho MPOA ka letsatsi la pelehi (PPD) 1, le maemo a phahamisang AMY ho PPD 6 (Jarrett et al., 2006; McMurray et al., 2008). Ho khahlisang, OT e ka bapala karolo ho litlamorao tsa meputso ea lithethefatsi. Boithuto ba morao-rao mehlaleng ea litoeba bo bonts'itse hore tsamaiso ea OT e ka fokotsa kapa ea thibela likarabo tse amanang le psychostimulant litekong tsa boikhethelo ba sebaka (CPP), boitaolo, le ho khutlisetsa mekhoa ea ho batla lithethefatsi (Yang et al., 2010). Lintlha tsena li fana ka maikutlo a karolo ea bohlokoa bakeng sa OT kopanong ea taolo, khatello ea maikutlo le MB.
Ho fetohela ha Ts'ebetso ea Meputso bakeng sa Kholo le Kameho ea Cocaine
Bo-makhopho le litheko tsa bana li na le boleng bo khotsofatsang (bo susumetsang) ba ho fokotsa matamo, bo bontšitsoeng ka bobeli ba CPP le li-paradigms tse sebetsang.Lee le al., 2000; Mattson le Morrell, 2005). Taba ea bohlokoa ke hore, malinyane hangata a hohela CPP ho feta phello ea koae, ho bonts'a matla a ho hlohlona ha litho tsa (Seip le Morrell, 2009). Leha ho le joalo, lithutong tsa litoeba, lipalo tse ngata tsa 30% ea matamo li rata kamore e sa amaneng le pop, mme matamo ana a bonts'a sensitization e kholo ea cocaine, a bonts'a setjhaba sa ka tlase se ka ba kotsing ea ho ba le tšusumetso ea koae ka ho potoloha ha meputso. ka MB (Mattson le Morrell, 2005; Seip le Morrell, 2007). Boleng ba tšusumetso ea "circry" e khothalletsang litšitiso tse matlafatsang bo bonahala bo feto-fetoha nakoana kamora hore matamo a se nang likarabo tse arabang (joalo ka ha a lekantsoe ke CPP) kamora ho amohela liente tsa "cocaine" tse tsamaeang le koae ha a bapisoa le likhoto tsa basali tse sa reng letho, a fana ka maikutlo a hore physiology e ka hare e ka bapala karolo e kholo litlamorao tsa maemo lithethefatsi (Seip et al., 2008).
Maikutlo a mangata a potolohileng moputso a khothalelitse hore DA e phethe karolo ea bohlokoa moputsong oa lithethefatsi le hore "potoloho ea moputso," joalo ka ha e hlahisitsoe ho Setšoantšo. 4, e na le tsela ea "midbrain-forebrain" e hokahanyang VTA le NAc le mPFC, e nang le leseli le fetohang ho NAc ho khanna likarabo tsa locomotor mabapi le ho batla moputso (Koob le Volkow, 2010; Sesack le Grace, 2010). Ts'ebeliso e pharaletseng ea fMRI e boetse e nolofalelitse lipatlisiso tsa lefutso ho batho ho ka etsoa lipatlisiso, ka libaka tse bapisoang, ho kenyeletsoa orbitofrontal cortex (OFC), amygdala, NAc, hammoho le PFC le anterior cingrate cortex (ACC; McClure le al., 2004). Ho ntse ho hlaha sengoloa se hanyetsang litloaelo tsa ho batla moputso mekhatlong e ikemetseng ea kelello: “ho batla,” “ho rata” le “ho ithuta” (Berridge, 2004; Berridge le al., 2009). Ho rata ho hlalosoa e le karabelo ea mokokotlo e tlisoang ke ho amohela moputso hanghang. Ka lehlakoreng le leng, "ho batla" ho hlalosoa e le boleng bo susumetsang ba moputso o tšoanang. Ho phatlalalitsoe hore dopaminergic signaling e hlalositsoeng ka holimo (VTA / Nac / mPFC) e bohlokoa haholo ho likarolo tsa "ho batla" tsa boitšoaro. Taba ena e ka thusa ho hlalosa hore na hobaneng, le ha likarolo tse ling tsa tlhokomelo ea bo-mme li sa khone ho ba le tšusumetso e matla (ke hore, mooki oa pele, tlhaselo ea basali, ho sitisoa ha boroko), bo-mme ba ntse ba "batla" ho fana ka tlhokomelo ea bo-mme.
Tšoantšiso 4
Setšoantšo sa 4. Moputso circry bokong ba methapo ea letsoho. Letlapa lena le bohareng ba "rodent" bohareng le bontša likarolo tsa mantlha tsa boko tse reretsoeng ho arabela ka mokhoa o khotsofatsang. Karolo ea "ventral tegmental" (VTA) ea sebopeho se tsamaisang methapo ea methapo e romela dopaminergic (metsu e putsoa) ho li-nucleus accumbens (NAc) le medial prefrontal cortex (mPFC). NAc (hammoho le habenula ea morao-rao, LHb) e romela likhakanyo tsa GABAergic (metsu e khubelu) ho VTA, ha mPFC e romella merero ea glutamatergic (metsu e tala) ho VTA. MPFC le NAc li na le menahano ea ho khutlisa glutamatergic. Liphetoho tsa ts'ebetso ho NAc li fella ka ho batla moputso kapa boitšoaro bo khothalletsang (motsu o motšo). Brain schematic e fetotsoe ho tloha Paxinos le Watson (1997).
Sebaka sa Tefo ea Ventral
Ventral tegmental sebakeng sa tšebetso ea "neuron" ea boikarabello ba boikarabello ba ho ts'oaroa ha nakoana le phasic DA ho kenya NAc (Sombers et al., 2009) hammoho le potolohong ea moputso, le ts'ebetso ea VTA ho lumeloa hore e bohlokoa bakeng sa boleng ba tšusumetso le boits'oaro bo shebileng sepheo (Koob le Volkow, 2010). VTA e na le li-neurons tsa projeke tsa DA, tse pota-potiloeng ke li-interneurons tsa GABAergic (bakeng sa tlhahlobo Adell le Artigas, 2004). Phatlalatso ea cocoaine e ama ts'ebetso ea VTA ka mekhoa e ka fetolang bokhoni ba eona ba ho arabela ts'usumetso e ntle ea tlhaho. Maemo a Synaptic a 5-HT, NE, le DA a eketseha haholo ka koae ka bokhoni ba eona ba ho thibela ba tsamaisang, e leng se bakang litheko tse ngata tsa ho thunya (Thomas le Malenka, 2003). Cocaine e boetse e kenya letsoho ho eketseng ho thunya ka ho fokotsa litšitiso, ka ho fokotseha ha VTA GABAergic interneuron ho thunya (Steffensen et al., 2008). Liphetoho tsena tse susumetsoang ke koae ha li luleng khafetsa; ka hona, cocaine ka mokhoa o hlakileng le e sa foleng e eketsa molumo o thabisang ho VTA, e hlahisang litheko tse phahameng (Thomas le Malenka, 2003), ka mokhoa o nang le monyetla oa ho chechisa mohopolo o mocha (oa lesea) o ke ke oa fihla. Lihormone tsa mahe a linotši li matlafatsa phello ea koae ho ho thunya ha VTA ho litoeba tsa basali (Zhang et al., 2008), ho fana ka tlhahiso ea hore liphetoho li ka ba ngata nakong ea kemolo ha lihormone tsa mahe li phahamisoa haholo. Habohlokoa, tsamaiso ea koae ea koae e baka ts'ebetso ea nako e telele ho VTA hore e se ke ea itaola.Chen le al., 2008), ho fana ka tlhahiso ea hore cocaine e ka thibela polasetiki e eketsehileng ho VTA e hlokahalang nakong ea pelehi.
Bopaki bo tsoang lithutong tsa litoeba bo tšehetsa karolo ea li-neurons tsa proion tsa DA tse tsoang VTA likarolong tse khothalletsang tsa MB (Numan, 2007). Meroallo ea VTA, kapa li-axons tse arohaneng ho tloha MPOA ho ea VTA, li sitisa MB (Numan, 2007). Ho qaptjoa ha nakoana ea VTA, haholo-holo ka GABAA Li-receptors, e sitisa ho khutlisoa ha bo-mme, ho ba mooki le CPP bakeng sa malinyane (Numan et al., 2009; Seip le Morrell, 2009), ha e ntse e sa sitise cocaine CPP, ho fana ka maikutlo a hore ts'ebetso ea VTA e bohlokoa bakeng sa likarabo ho susumetsoa ke popo. Litsuonyana tseo ka tlhaho li hlokomolohileng malinyane a tsona li bonts'a ts'ebetso e phahameng ea bas-c-FOS, leha ho sena phapang lipalo tsa lisele tsa DA ho VTA (Gammie et al., 2008a). Sena se fana ka maikutlo a hore ho hlokomoloha malinyane ho ka nkuoa ho le molemo bakeng sa matamo ana, mohlomong ka lebaka la phokotso ea khatello ea maikutlo. Leha ho le joalo, hore na ho tlosoa ha bana ho fokotsa khatello ea maikutlo le ho nyolla letšoao la moputso matamong a mang empa ha ho joalo ka ba bang ba ntseng ba lokela ho hlahlojoa ka kotloloho. Likarabo tsa Electroencephalogram (EEG) le fMRI ho VTA li eketseha ka ho pepesetsoa lipuisano le lipopao tsa likokoanyana tsa bana (Febo et al., 2005; Hernandez-Gonzalez et al., 2005). Ts'ebeliso e tobileng ea OT kapa opioids ho VTA e ka tsamaisa MB (Pedersen et al., 1994; Thompson le Kristal, 1996). Bopaki ba morao-rao bo bonts'a karolo e tobileng bakeng sa OT molomong oa ho thunngoa ha lisele tsa VTA le ho lokolloa ha DA ho hlaheng.Shahrokh et al., 2010). Ho khahlisang, bahanyetsi ba OT ba fokotsa ho lokolloa ha DA matamong a basali haholo, empa eseng matamong a tlase a bo-mme, ho fana ka maikutlo a hore ho ka ba le phello e "matla" ho bokhoni ba OT ba ho tsamaisa DA ho thunya lisele. Ha li kopantsoe hammoho, liphumano tsena li bonts'a tšitiso ea ts'ebetso ea VTA e ka ama haholo MB le boleng bo khotsofatsang ba malinyane ho matamo. Phekolo ea koae ea Gestational ha e ame maemo a basal a NE, DA, kapa DA metabolites ho VTA nakong ea pelehi; leha ho le joalo, basal 5-HT le OT li fokotsehile (Johns et al., 1997a; Lubin et al., 2003). Lintlha tsena, hammoho le liphetoho tse boletsoeng esale pele tsa koae ts'ebetsong ea VTA, li fana ka maikutlo a hore li-neurons tsa VTA li ka 'na tsa se ke tsa arabela haholo ho OT mme ka hona, li fokotseha haholo, leha sena se sa ntse se lokela ho hlahlojoa ka kotloloho.
Nucleus Accumbens
NAc, e arotsoe mokokotlong le khetla e sebeletsa mesebetsi e mengata ea boits'oaro. Khetla e kenelletse mehahong e hlohlelletsang ea ho susumetsa ea moputso ka ntlafatso ea mekhatlo ea meputso, ha karolo ea mantlha e kenella likarolong tsa ts'ebetso ea ho batla meputso (Di Chiara, 2002; Russo et al., 2010). Ka ho batho, NAc e loants'ara tebello le ponelopele ea moputso (Knutson le Cooper, 2005), ka mesebetsi ea tikoloho e fapaneng ho latela boholo ba moputso (Haber le Knutson, 2009). NAc e na le li-neuron le li-interneuron tse hlahelletseng tsa Gabaergic. Morero o ka sehloohong oa NAc GABAeric efferents ho lateral hypothalamus, VTA, substantia nigra, brainstem, le ventral pallidum, tseo ts'ebetso ea eona e amanang le ho batla moputso (Koob le Volkow, 2010). Ts'ebetso ea phetisetso ea DA ea thibelo ho tsoa ho VTA ho NAc ho arabela taolong e matla ea lithethefatsi tsohle tse kholo tsa tlhekefetso e hlokometsoe (Koob le Volkow, 2010). Cocaine haholo e eketsa maemo a inhibitory 5-HT le NE joalo (Li et al., 1996). Ho pepesetsoa ha koae ho sa feleng ho etsa hore D1 li-receptors striatum (Ben Shahar et al., 2007), e ts'oanang le polelo ho matamo a bo-mme haholo (Champagne et al., 2004), ho fana ka maikutlo a hore ho pepesehela maikutlo a susumetsang a nang le moputso ho ka ba le litlamorao tse tšoanang. Le ha polelo ea DA ea transporter e sa fetohe, ts'ebetso ea eona e eketseha ka tšebeliso ea koae (Oleson et al., 2009). Phatlalatso ea koae e sa foleng e hlakisa polelo ea transporter ea 5-HT, leha e sa ame polelo ea transporter ea NE (Belej et al., 1996). Keketseho ea ts'ebetso ea transporter e ka bonts'a teko ea li-neurons ho khutlela boemong ba ho thunya ho tšoanang le ho boneng pele ho ts'ebeliso ea lithethefatsi. Ho pepesetsoa ha cocaine ho fokotseha matla a synaptic pakeng tsa maikemisetso a mofuthu a PFC ho khetla ea NAc, mme ho baka ho fokotseha ha nako e telele hoa mantlha (Russo et al., 2010).
NAc e bohlokoa bakeng sa ho qala le ho hlokomela MB. Tlatsetso ea NAc e fokotseha haholo MB, 'me ka ho khetheha, lesapo la NAc le senya mokhoa oa ho khutlisa ntle le ho sitisa ho tsukutlana, ho nyeka kapa moaho oa sehlaha (Li le Fleming, 2003). NAc e bonts'a ts'ebetso e eketsehileng ea neuronal, ka polelo ea c-FOS le fMRI, ho arabela malinyane ho beke ea pele ea pelehi (Fleming le Korsmit, 1996; Febo et al., 2005). Leha ho le joalo, ha ho karabelo e bonoang feela ho lithaka tsa banaFleming le Korsmit, 1996). Ho fapana le moo, ts'ebetso e arabelang ho lla hoa masea (e amanang le lerata le tšoeu) e bonoe ho bo-mme ba batho libakeng tse potolohileng NAc, hammoho le libaka tse atolositsoeng tsa cortical le subcortial, haholo-holo tse ngotsoeng ke dopamine (Lorberbaum et al., 2002).
Likhoto tse nyehelang li na le DA ea basal e tlase ho NAc ha e bapisoa le likhoto tsa boroetsana (Olazabal et al., 2004), mohlomong ho lumella kutloisiso e kholo ho liphetoho maemong. Hona joale Ka vivo litekanyetso tsa voltammetry le micodialysis li bonts'itse kholo ea NAc shell ea DA nakong ea booki, le khokahano e tobileng ea mahloriso a DA ka nako e telele ea boits'oaro bo bobe (Champagne et al., 2004; Afonso et al., 2008). Phekolo le ba-agonists ba DA e eketsa lick ka kotloloho ho matamo ao pele a neng a tsebahala e le "ho theola tlase" (Champagne et al., 2004), ho fana ka maikutlo a hore mohato o mong o tlameha ho fihlelleha ho fihlela litheko tse phahameng. Mekhoa e tšoanang ea kalafo ea litlama ho NAc e bonts'itsoe ho phehella "ho hloka" meputso (Berridge le al., 2009). Haufinyane, ho bontšitsoe hore D2 ts'ebetso ea li-receptor e bohlokoa bakeng sa MB e tloaelehileng (Zhao le Li, 2010). Leha D2 li-receptor binding ha li tšoane pakeng tsa matamo a nang le bo-'mè ba phahameng haholo le a matangoane, a matamo a matšehali a nang le bo-'mè ba nang le D.1 le D3 li-receptors le li-transporter tse tlase tsa DA li tlama ka hara khetla ea NAc ha li bapisoa le matamo a tlase-a nyehelang / boikoetliso (Champagne et al., 2004). Leneng la toeba le etselitsoeng ho hlokomolohuoa ha bo-mme, polelo e phahameng haholo ea C-FOS e bonoa hang kamora ho qaleha ha ho qheleloa thoko ha ho bapisoa le matamo a taolo.Gammie et al., 2008a), ka phello e kholo bohareng ho feta khetla. Ha ho tsamaisoa hammoho, ho supa dopaminergic ho NAc ho bohlokoa bakeng sa MB, 'me kaha tšebeliso ea lithethefatsi e ka fetola haholo ho supa, sena se ka lebisa ho makhopho a MB. Leha ho le joalo, ha ho tsejoe haholo ka hore na ho pepesetsoa ha lithethefatsi ho ka ama polasetiki ka hara NAc nakong ea kemolo, ho arolelana, le lebollo mme sena e tla ba sepheo sa lipatlisiso tsa nako e tlang.
Pele Cortex
PFC e kenya letsoho mesebetsing e mengata e fapaneng ea kelello, eo kaofela e leng likarolo tsa bohlokoa phetohong ea ts'ebeliso ea lithethefatsi, bokhoba ba tahi kapa MB. Ho utloisisa karolo ea PFC ho rarahane, haholo ha ho fanoa ka phapang e kholo litekanyetsong tsa cortical parcellation lithutong tsa batho le tsa liphoofolo (ho kenyeletsoa ACC, OFC, mPFC, le infralimbic) e hlokometsoeng lithutong tsohle (Dalley et al., 2004). ACC e amahantsoe le ho kenyeletsoa le ho ananeloa ha tlhaiso-leseling ea sechaba ka lebaka la ho ikamahanya le eona ka kotloloho le AMY, ventral striatum, hypothalamus, periaqueductal grey (PAG), le cortex ea auditory (Dalley et al., 2004), hammoho le tatellano ea boits'oaro ba nakoana ea boits'oaro. OFC, e nang le karolo e thehiloeng hantle boikarabelong ba ho etsa liqeto le ho tsosa likamano tsa meputso, e amohela mohopolo ho tsoa mekhoeng eohle ea kutlo le ho kenella ka hare ho terriatum le amygdala. Tšenyo ea OFC e ka baka phetoho ea ho tšoenyeha / tšabo le boits'oaro bo mabifi, ho fana ka maikutlo a bohlokoa ba eona litšebelisanong tsa sechaba. Li-cortices tsa prelimbic le infralimbic, tseo e leng karohano ea mPFC e lutseng ka kotloloho le ka tsela e fapaneng, li amana le ts'ebetso ea memori le ho sebetsa. Leha ho le joalo, ba bapala likarolo tse fapaneng tsa ho ithuta, ka cortex ea prelimbic e kenyang letsoho ho mekhatlo ea sephetho sa sephetho le infralimbic e kenyang letsoho ho theheng mekhoa ea litoeba. Ka bobeli mPFC le OFC li bile le tšusumetso ho laola khatello ea maikutlo (Dalley et al., 2004), ntlha eo re tla khutlela ho eona hamorao. Ha ho nahanoa hore mesebetsi ena kaofela e bohlokoa bakeng sa litšebelisano tse ntle tsa sechaba, ho utloisisa hore na tšebeliso ea lithethefatsi e fetola tšebetso ea bona joang ho tla hlakisa hore na libaka tsena li ka nka karolo joang literekeng tse fetotsoeng tsa kamoso.
Ts'ebeliso e mpe ea lithethefatsi e tlamahantsoe le bofokoli ba ho saena hoa monoamine ho PFC; Leha ho le joalo ho ntse ho sa hlake hore na likhaello tsena li bakoa kapa ke biketsiso (litekolo: Dalley et al., 2008; Koob le Volkow, 2010; Sesack le Grace, 2010). Bopaki ba morao-rao bo bonts'a ts'ebetso ea limolek'hule e etsang hore ho se sebetse hantle ho nang le ts'ebeliso ea lithethefatsi. Glucose metabolism e eketsa ho latela k'hok'heine e mpe haholo, empa e fokotseha ka mor'a ho itaola k'hok'heine, e bonts'ang karabelo e fetohileng ea neuronal (Hammer Jr. le Cooke, 1994). Cocaine e eketsa phallo ea mali lithutong tsa fMRI tse kenyeletsang likhoto tsa banna, tsa basali le tsa lactating (Febo et al., 2004; Ferris et al., 2005). "Ts'ebetso" ena e eketsehileng ea mPFC, e lekantsoeng ke phallo ea mali e eketsehileng, e ka hlaha mesebetsing e eketsehang ea li-interneurons tsa GABAergic ho e-na le li-neurons tsa projamestgic tsa projeke, hobane cocaine e baka thibelo e kholo ea VTA ea li-neuron tsa mPFC (Peterson et al., 1990), ho fana ka tlhahiso ea ts'ebetso e eketsehang ea li-neuron tsa projeke sebakeng sena. Ts'ebeliso ea cocaine e baka phetoho ea mofuta oa li-genaptic tsa cellapt tse makholo joalo ka ha li lekantsoe ke Microarray (Freeman et al., 2010) hammoho le ho nyolleloa ha D1 receptor le corticotropin releasing factor (CRF) shughuli (Ben Shahar et al., 2007; Corominas et al., 2010). Ha li kopantsoe hammoho, lintlha tsena li hlahisa mekhoa e mengata ea ho theoha ha ts'ebetso ea mPFC ka ts'ebeliso ea koae. The ACC, OFC, infralimbic, le prelimbic cortex tsohle li bonts'a polelo e eketsehang ea c-FOS ha e araba mekhoa e amanang le koae ha e bapisoa le taolo ea saline (Ciccocioppo et al., 2001), ho supa karolo ea bohlokoa bakeng sa ho ithuta ka lithethefatsi libakeng tsena. Kaha libaka tsena li kanna tsa arabela ho pepesetsoa hoa koae ka phetoho ea polasetiki ho polelo ea liphatsa tsa lefutso, li-neuron tsena li kanna tsa se ke tsa arabela ka tekanyo e nepahetseng ea neuroplasticity e hlokahalang bakeng sa phetoho ho etsa MB. Phuputso e le 'ngoe ea tsamaiso ea koae pele ho ima e bontsitse ho fokotseha ha ts'ebetso ea ho anyesa mPFC, empa ha e a ka ea ama motheo oa DA kapa keketseho ea karolo ea DA ha e pepesehela malinyane mPFC (Febo le Ferris, 2007). Leha ho le joalo, hore na cocaine nakong ea kemolo e ama nts'etsopele ea liphetoho tsa polasetiki ho PFC nakong ea bokhachane le pelehi, e ts'oanang le e bonoang ho banna ha e so lokela ho hlahlojoa ka kotloloho.
Karolo ea PFC ea ho hlophisa boits'oaro e bohlokoa bakeng sa phetoho ho MB, le litšitiso tsa ts'ebetso ea PFC tse bakang khaello ea MB. Khatello ea litheko tsa lithethefatsi tsa methapo ea sodium kapa ts'ebetso ea GABA mPFC e bonts'itse hore sebaka sena sea hlokahala bakeng sa ho khutlisa boits'oaro ba matamo a rat (Febo et al., 2010). Liteko tsena ha lia ka tsa fetola boitšoaro ba batho ho malinyane, empa feela qeto ea ho ba khutlisetsa sehlaheng, e bonts'a phetoho ea boits'oaro e seng boits'oaro ba lipatlisiso. Lesitotoxicion ho mPFC le eona e sitisa ho khutlisetsa bana, ho nyeka, le paterone kapa tatellano ea li-MB, ho bonts'a bohlokoa ba sebaka sena molemong oa ho hopola le ho ela hloko nakong ea pelehi (Afonso et al., 2007). Ho ts'oara ha Pup ho phahamisa karabelo ea fMRI ho PFC ea morusu le ea morao-rao le infort cortex ea lactating rats, phello e itšetlehileng ka OT (Febo et al., 2005). Lintlha tsa EEG li fana ka tlhahiso ea hore tšebetso ea mPFC e fetohe ho arabeleng monko oa bana (Hernandez-Gonzalez et al., 2005). Joalokaha ho boletsoe ka holimo, DA e kenya letsoho tšebetsong ea PFC. Maemo a DA a tlase ho litoebeng kamora nako ea bokhachane ha a bapisoa le litoeba tsa basali ba batšehali (Olazabal et al., 2004), e ka hlahisang ts'ebetso e phahameng ka ho fetesisa e fuoeng DA e thibela ts'ebetso ho mPFC (Peterson et al., 1990). Haufinyane, ts'usumetso e phahameng e hokahane le bofokoli ho MB, e ka amanang le liphetoho tsa tšebetso ea mPFC (Lovic et al., 2010). Kaha mPFC DA ke mosireletsi oa bohlokoa oa khatello (Dalley et al., 2008) mme e ka sitisoa ke ts'ebeliso e mpe ea lithethefatsi, liphapang tse teng mokhatlong oa boitšoaro nakong ea MB li ka hlaha kamora ts'ebeliso ea lithethefatsi (leha sena se sa ntse se lokela ho hlahlojoa ka kotloloho). Haholo-holo, pejana ho PPD1, cingxe cortex e bonts'a polelo e eketsehileng ea c-FOS ha e araba malinyane, mme e ntse e tsoela pele ho arabela ka li-cues bekeng ea pele (Fleming le Korsmit, 1996). Ntle le moo, cortex ea infralimbic e arabela ho cues ha prelimbic cortex e sa etse (Fleming le Korsmit, 1996), e fana ka tlhahiso ea bohlokoa ba tikoloho e ikhethang ea potoloho.
Ka ho bo-'mè ba batho, leha mosebetsi o atileng ka bokong o bonoa ha ho pepesoa mekhoa ea masea, OFC e hlaha e le sebaka sa mantlha ho potolohong ea ho ba motsoali, ka ho ts'ehetsa lithuto ka botlalo le ka mekhoa e fapaneng. Ts'ebetso ka letsohong le letona la OFC e ne e le kholo ha bo-'mè ba mamela likeleli tsa masea ha li bapisoa le lerata le tšoeu (Lorberbaum et al., 1999, 2002), le tšebetso ea naha ea OFC e eketsehile ha bo-mme ba shebella lifoto tsa ngoana oa bona ha ba bapisoa le ngoana ea sa tsejoeng (Nitschke et al., 2004). Ho bokellana ha kelello le maikutlo a ho itlaleha, ts'ebetso ena e kopanetsoeng ho OFC e kopantsoe haholo le lintlha tse ntle tsa maikutlo ha u ntse u shebile lifahleho tsa masea (Nitschke et al., 2004), ka ts'ebetso ea kushoto ea OFC e tsamaellana le mohopolo o motle le tšebetso e nepahetseng ea OFC e lumellanang le lintlha tse mpe tsa mohopolo thutong e latelang (Noriuchi et al., 2008). Tlalehong ena ea morao-rao, libaka tse ling li bontšitse kutloelo-bohloko ho tloaelo ea masea, ho kenyeletsoa le dColoral PFC, insula, putamen, le PAG. Mosebetsing oa tlhahlobo ea pele, PAG e nahanoa hore e kopanya boemo ba maiketsetso ba bookameli ho tloha ha ho pepesehela malinyane a anyang ka mokhoa o ikhethileng ho kenya ts'ebetsong PAG ka tekanyo e kholo ho feta ho pepesehela malinyane a sa anyang (Lonstein le Stern, 1997). PAG e boetse ea kenella haholo ho laola boitšoaro bo mabifi nakong ea pelehi le ho tsamaisa tšabo kapa matšoenyeho (Lonstein et al., 1998). Ho bohlokoa ho hlokomela hore OFC ea morao-rao (le PAG) e arabela ka mokhoa o ikhethileng litloaelanong tsa basali, tse nang le libaka tse akaretsang tse kenyeletsang striatum, insula, le dorsal ACC e arabelang mekhoeng ea ho ikamahanya le basali le maikutlo a lerato (Bartels le Zeki, 2004). Magnetoencephalography e boetse e bonts'a karolo ea OFC ho hlokofatseng kutloelo-bohloko ea masea mme e boetse e fana ka maikutlo a hore OFC e ka fana ka karolo e holimolimo ho pono ea sefahleho sa masea (Kringelbach et al., 2008). Mohlala o nang le batsoali le bao e seng batsoali, ho qalisoa ha 130 ms ka mor'a ho khothaletsoa ho bile le keketseho e kholo ea ts'ebetso MOFC ho arabela ho shebella lifahleho tsa masea empa eseng sefahleho sa batho ba baholo. Ntle le moo, maikutlo ana a khale a lifahleho tsa masea a ne a sa bonoe libakeng tseo ka tloaelo li amanang le ts'ebetso ea sefahleho (ke hore, fusiform cortex). Leha ho le joalo, kamora ho 165 ms ho tloha ponong ea sefahleho, phapang e ka bapisoang ea mesebetsi e arabelang lifahleho tsa masea le tsa batho ba baholo ho fusiform cortex e ile ea bonoa. Liphumano tsena li fana ka maikutlo a hore mOFC ha e amehe feela ka mekhoa ea masea, empa e ka boela ea lekanya liketsahalo tse latelang libakeng tsa fusiform bakeng sa ts'usumetso e ntle ea ts'usumetso ea sefahleho sa masea.
Hamorao Habenula
Sebaka se seng sa bokapele se ka tlatsetsang ho MB ke habenula ea morao-rao (LHb; Geisler le Trimble, 2008). Ts'ebetso ea LHb e hokahane le ho haella ha moputso o lebelletsoeng hammoho le ts'usumetso e sithabetsang, e fana ka maikutlo a karolo ea ts'ebetso ea ho thella le boleng ba meputso e khothatsang le e sithabetsang. LHb e bonts'a c-FOS e eketsehang ea li-cocaine tse mpe haholo le tse amanang le koae empa karabelo ena e fokotse ho senoloa khafetsa (Franklin le Druhan, 2000), ho fana ka tlhahiso ea hore ho pepesetsoa ha koae ho sitisa bokhoni ba LHb ba ho fokotsa mesebetsi ea VTA. Sena se ka ba bohlokoa ka ho khetheha haeba li-neurons tsa VTA li se li fihlile boemong ba ho thunya bo ke keng ba fetoha ka ho fetesisa ha ho angoa le ts'usumetso ea masea. Sebopeho sena se entsoe ka mokhoa oa ho araba malinyane a PPD7 mme se kopana le lithapo tsa bana ho PPD 10 (Felton et al., 1998). Ho khahlisang, karabelo ea c-FOS e fokotseha matamo a bonts'a CPP e matla bakeng sa malinyane. Sena se ka hlalosoa ke karolo ea LHb maemong a sa thabiseng a moputso (Mattson le Morrell, 2005). LHb e boetse e bontšitsoe hore e na le maikutlo le menahano e thabisang ea bobeli (MB)Geisler le Trimble, 2008). Ena ke sebaka se khahlisang sa lipatlisiso tsa nako e tlang kaha se ka bapala karolo ea bohlokoa ho khetholleng boroko ba mehopolo e fapaneng nakong ea ho beleha.
Ka kakaretso, meaho e mengata ea mantlha e fumanoe ho li-neural circry mme re hlalositse bopaki ho fana ka maikutlo a ho ikamahanya le boemo ba bona ba botsoali. Ho feto-fetoha ha methapo ea kutlo ka cocaine ho kenya letsoho tseleng eo tšebeliso ea lithethefatsi e ka amang boitšoaro ba botsoali. Karolo ea li-circry tsa moputso, haholo mesocorticolimbic DA, le eona e bile le tšusumetso khokahanong ea sechaba ka bophara ho feta lithutong tsa preclinical tsa MB le li-bonding (Insel, 2003). Sena e tla ba mokhoa oa bohlokoa oa ho etsa lipatlisiso tsa nako e tlang ho utloisisa hore na cocaine e susumetsa MB joang le ho theha le ho boloka kamano ea bo-mme.
Phetoho ea tsamaiso ea khatello ea maikutlo bakeng sa botsoali le litlamorao tsa Cocaine
Ntle le moputso oa li-circuits tsa neural, ho boetse ho na le mosebetsi oa bohlokoa oa ho hokahanya li-neurocircuits ho MB ho batho le litoeba. Ho feta moo, leruo la boits'oaro le nang le boits'oaro le fana ka khatello ea maikutlo ho batleng le lithethefatsing (Corominas et al., 2010; Koob le Volkow, 2010). Ka hona re lebisa tlhokomelo ea rona ho potoloha ea neural ea sistimi ea khatello ea maikutlo, ho kenella ha eona ho MB, le ho feto-fetoha ha molumo oa koae.
HPA Axis
Ts'ebetso ea khatello ea kelello ea "HPA axis" e bonahala e bapala karolo ea bohlokoa ho ntlafatseng tlhekefetso ea lithethefatsi, athe extrahypothalamic stress circry [BNST, hippocampus, karolo ea medial ea NAc, le amydala (CeA) e bonahala e na le karolo ea bohlokoa ho feta ts'usumetso. litlamorao tsa ho ikhula hape ho matla le khatello ea maikutlo e khutlisetsang morao (Aston-Jones le Harris, 2004; Corominas et al., 2010; Koob le Volkow, 2010). Ho nahanoa hore ts'ebeliso ea tlhekefetso e tsoa ho phetoho ea neuroadaptational mabapi le hore na meputso e sebetsoa joang, haholo-holo tahlehelo e ntle ea ts'ebetso le phetiso e 'ngoe ka ts'usumetso e mpe ka hare ho potoloho ea basal e reng amygdala e holisitsoeng (Koob le Volkow, 2010). Phetoho ena ea nako e telele ea kamoo methapo ea khatello ea kelello e sebetsanang le lits'oants'o tse tšoanang le tikoloho (allostasis) kamora ho pepesetsoa lithethefatsi kapa liketsahalo tse nyahamisang tse hlalositsoeng e hlalositsoe e le mojaro o matla haholo (McEwen le Gianaros, 2011). Liphetoho maemong ohle a tlisoang ke ho pepesetsoa khatello ea maikutlo le 'mele.
Ehlile, axis ea HPA e hlahisoa ke liketsahalo tse fapaneng tsa kantle le tsa kahare (bona setšoantšo 5). PVN ho hypothalamus e lokolla CRF ka har'a phepelo ea mali ea hypophyseal, e susumetsang e lokolloang ha adrenocorticotropic hormone (ACTH) ho tsoa gland ea pituitary ho kenella phepelong ea mali. ACTH e sebetsa ho adrenal medulla ho lokolla glucocorticoids (GC), ke hore, cortisol (batho) kapa corticosterone (CORT; litoeba) ho potoloha moo e fanang ka litlamorao tse ngata tsa 'mele. Habohlokoa, CORT e fana ka maikutlo a fosahetseng ka ts'ebetso ea GC ho pituitary, PVN, le hippocampus, e khutlisetsa tsamaiso ho homeostasis. Ntle le ho lokolloa ha CRF maling, li-neurons tsa PVN li ea libakeng tse ling tsa methapo ea methapo ea kutlo, tse kang BNST, CeA, le VTA (Palkovits et al., 1998; Rodaros et al., 2007), ho fella ka likarabo tse fapaneng tsa li-neuronal libakeng tseo tsa boko (bakeng sa tlhahlobo bona Corominas et al., 2010). PVN e khutla ka mokhoa o feto-fetohang ka mokhoa oa kemolo nakong ea kemaro le ho lactation ho lumella tlatsetso e kholo ea thabo (Panatier le Oliet, 2006), e fanang ka maikutlo a hore ena ke nako e matla haholo bakeng sa liphetoho bakeng sa lits'ebetso tsa khatello ea kelello. Haeba karabelo ea tšebeliso ea lithethefatsi PVN e arabeloe nakong ea kamora ho pepa, sena se ka ba le litlamorao tse tlisoang ke hore PVN e na le lisele tse hlahisang OT (Slattery le Neumann, 2008), mme PVN e fumanoe e sebetsa ho kenya ts'ebetso ho arabela lipotsana ke PPD7 (Fleming le Korsmit, 1996; Febo et al., 2005).
Tšoantšiso 5
Setšoantšo sa 5. Ho sithabela ha khatello ea kelello kelellong ea methapo. Neroase ea "patventricular nucleus" (PVN) ho hypothalamus e romela likhakanyo tsa corticotrophin releasing factor (CRF) (metsu e tala) ho amygdala (CeA), mookotaba oa bethe ea stria terminalis (BNST), le gland ea pituitary. Tšoelesa ea “pituitary gland” e ntsa hormone ea adrenocorticotrophic (ACTH) molomong oa mali o eang gland ea adrenal. Adrenal gland e ntsha corticosterone (CORT) maling. CORT e sebetsa e le lets'oao la maikutlo a mabe (metsu e khubelu) ho pituitary, PVN, le hippocampus, e romellang maikutlo a nyakallo ho PVN (metsu ea mola o motala). PVN e fumana litlatsetso tse eketsehileng tse tsoang ho CeA le BNST. Brain schematic e fetotsoe ho tloha Paxinos le Watson (1997).
Ho na le kamano e thehiloeng tlatsong pakeng tsa tšebeliso e mpe ea lithethefatsi le tšenyo e amanang le khatello ea maikutlo ho batho le mefuta ea liphoofolo (Sinha, 2001; Goeders, 2002; Koob le Volkow, 2010). Cocaine e sebetsa hantle axis ea HPA (Goeders, 2002), karabelo e hlahisoang ke lihormone tsa botona ba botšehali (Russo et al., 2003), ho fana ka maikutlo a hore bokhachane, le lihormone tsa basali tse ngata tse potolohang tsa basali, e ka ba nako e tebileng haholo bakeng sa litlamorao tsa khatello ea maikutlo ea koae. Litlamorao tse sa feleng li itšetleha ka 'muso oa kalafo; mohlala, HPA ha e arabe ebile ha e tsotelle ho tsamaiso ea koae ea letsatsi le letsatsi, le ha likarabo tsa ACTH le CORT litekong tse tlisoang ke ho itlopa lihlahisoa li sa luleng ka makhetlo a mangata (Goeders, 2002). Leha ho le joalo, ho itšeha ha k'hok'heine ho baka karabelo e atileng ea CORT mme e fokotse karabelo e mpe e tsamaellanang le li-receptor tsa GC tse tlase ho PVN empa eseng likarolo tse ling tsa forebrain (Rodaros et al., 2007), ho supa libaka tse ling tsa boko ho ka bonts'a karabo e tsoelang pele. HPA reactivity ea matlafatsoa nakong ea ha motho a khaotsa haholo, 'me khatello ea mali e phehella nakong ea ho itšoaea sefuba (Goeders, 2002; Corominas et al., 2010). Ho bohlokoa, koae e sa foleng e ka phahamisa maemo a CORT haholo nakong ea kemolo (Quinones-Jenab et al., 2000), leha e le hore tshusumetso ho molao oa tsamaiso ea litaba e hlakile. Tlhahisoleseling e tlatselletsang e bonts'a hore khatello ea maikutlo le ho supa ha HPA ho ka tsamaisa tsamaiso ea kelello ea psychostimulant (Goeders, 2002), e bonts'a mokhoa oo khatello ea maikutlo e ka bang le ona ho batleng lithethefatsi kamora ho pepa.
Karolo ea khatello ea khatello ea HPA ho MB e sa tsoa qala ho utloisisoa, 'me ho hlakile hore molao o tiile o nka karolo phetohong eohle ho tloha moimaneng, ho losis le ho khoesoa. Joalokaha ho boletsoe kaholimo, allostasis kapa karabelo e matla ea HPA le lits'ebetso tsa khatello ea kelello maemong a fetohang a fetohang, mohlomong e bapala karolo ea bohlokoa, leha ho le joalo, karolo ea mekhoa ea allostatic e tlhokeho e kholo ea ho ithuta. Nako ea boikoetliso le nako ea kamora ho pepa e tšoauoa ka maemo a phahameng a CORT ea basotho, karabelo ea lihomone tsa khatello ea maikutlo ho khatello ea maikutlo, le maemo a tlase a ho tšoenyeha (Slattery le Neumann, 2008). Liphetoho tsa karabelo ea khatello ea maikutlo ea mme li tsamaellane le bofokoli ba tlhokomelo ea bo-mme (Smith le al., 2004; Bosch et al., 2007; Chen le al., 2010). Matšoenyeho nakong ea kemolo a ka fokotsa MB ka litoeba, leha ho le joalo, haeba likhoto li ne li ba le MB e tlase, khatello ea maikutlo ha ea ka ea ba ama, a fana ka maikutlo a hore tlhokomelo e nepahetseng e ka fokotsoa ho isa bohōleng bo itseng feela (Champagne le Meaney, 2006). Ho tsamaisa CORT ho likhama tse emereng kapa tse anyesang ho fokotse booki mme ho eketsa boits'oaro bo sa tsotelleng (Bosch et al., 2007; Brummelte le Galea, 2010). Matšoenyeho a phetoang nakong ea pelehi e ka emisa ho lactation ka litoeba, ho fana ka maikutlo a litlamorao tse tobileng tsa lihormone (Lau le Simpson, 2004). Ka lehlakoreng le leng, ho tlosa li-hormone tsa khatello ea maikutlo ho fokotsa empa ha ho felise MB (Rees et al., 2004). Lactation e ipapisitse le maemo a poteletseng a OT mme OT e tsejoa ho sebelisana hantle le ts'ebetso ea HPA, ka kalafo e sa foleng ea OT e lebisang ho karabelo tse fokolisang tsa khatello ea maikutlo (Uvnas-Moberg et al., 2005), ho fana ka maikutlo a hore OT e ka thusa ho thusa ho fokotsa khatello ea maikutlo nakong ea pelehi (Slattery le Neumann, 2008).
Li-neurotransmitters tse ngata tse kenyelletsang ho taolo ea khatello ea maikutlo li fetoloa nakong ea pelehi ea pelehi, ho kenyelletsa 5-HT, DA, NE, vasopressin, OT, le CRF (Slattery le Neumann, 2008). Lipontšo tsena li sebetsa haholo kahare ho PVN ho tsamaisa karabelo ea khatello ea maikutlo, haholoholo ho lokolloa ha CRF le OT. CRF e sebetsa e le sesupo sa "khatello ea maikutlo" eseng feela ka ho sebetsa axis ea HPA, empa hape le ka ho supa ho amygdala e atolositsoeng le VTA, e lebisang ho eketseha ha litšebeletso tse potileng ketsahalo e sithabetsang (Gulpinar le Yegen, 2004; Corominas et al., 2010). Ho bile le tlhahiso ea hore liphetoho tsa kamora ho sebetsana le khatello ea maikutlo li bakoa ke phokotso ea tlhahiso ea CRF ho PVN (Slattery le Neumann, 2008), mohlomong ka maemo a phahameng a OT, a ka fanang ka tlhahiso ea CRF mRNA ha a sebetsana le khatello ea maikutlo (Lightman et al., 2001; Windle et al., 2004). Letotong la lithuto tse sebelisang mela e potolohileng ea mouse, Gammie le basebetsi-mmoho ba bonts'itse hore CRF signature modulates likarolo tsa MB (Gammie et al., 2007, 2008b; D'Anna le Gammie, 2009). Ts'ebetso e lebelletsoeng ea CRFR1 e fokolitse haholo booki, ha CRFR2 Matamo a Knoutout a bontša ho fokotsa mabifi a 'm'ae tlhahlobong ea batho ba kenang sekolo. Kaha ho pepesehela motho ea kenang ntle le tumello ho ka u sithabetsa haholo bakeng sa letamo, ho ka etsahala hore ts'ebetso ea CRF e bohlokoa haholo bakeng sa MB e amanang le maemo a mabe kapa a ho tšoenyeha. Liphetoho ho lets'oao la CRF-Mediated, joalo ka ha le bonoa ka kalafo ea koae (khafetsa)Corominas et al., 2010), ka hona e ka senya ts'ireletso e tloaelehileng ea bana. Joale re tla sheba likarolo tsa bohlokoa tsa neural tse amehang karabong ea khatello ea maikutlo, bokhoba le motsoali; ka ho khetheha hippocampus le amygdala e atolositsoeng, pele a hlahloba ts'ebelisano ea bohlokoa lipakeng tsa khatello ea maikutlo le li-circuits tsa moputso.
Hippocampus
Ts'ebetso ea Hippocampal e fana ka tšusumetso e thibelang, ka khokahano e otlolohileng ea "neurons" e amanang ka kotloloho le PVN, hape e laola tokollo ea lihormone tsa khatello (Herman et al., 2005). Hippocampus e na le likhokahano tse khahlang tsa ho hlasimoloha, ka "intorhinal cortex", le mPFC, ACC, insular, le li-cortices tse ling tsa mokhatlo, e fana ka maikutlo a karolo ea ona ho hokahaneng le lintlha tsa nako le tsa boiketlo ba sechaba, hammoho le ho kenya letsoho karabong ea khatello nakong ea kemolo le ho nyeka. Ho pepesetsoa hoa koae e sa fetoleng boemo bo hlalosang li-signature tsa monoamine hammoho le litsela tse 'maloa tsa kinase tse bontšang (Dworkin et al., 1995; Freeman et al., 2001), ho fana ka maikutlo a cocaine ho ka fokotsa taolo ea bokhoni ba sebopeho sa hippocampal ho kokobetsa khatello ea khatello ea PVN.
Hippocampus e bonts'a lets'oa la BOLD ha le araba ngoana ea anyang (Febo et al., 2005), mme liso tsa sebaka sena li tla senya botata ka kotloloho (Kimble et al., 1967), fana ka tlhahiso ea mohlomong karolo ea ho ithuta libaka tse sireletsehileng tsa booki. Cortex ea entorhinal, e bapileng ka kotloloho le hippocampus e bonts'a karabelo e ntle ea BANG ho ngoana ea anyang (Febo et al., 2005), e bonts'ang karolo ea mohopolo oa sechaba. Batho ba baholo ba neurogeneis ka hippocampus e fokotseha ho litoeba tse khahlisang basali, phello e tlameletsoeng maemong a CORT a potolohileng.Pawluski le Galea, 2007), mme e ts'oana le se bonoang kamora ts'ebeliso ea koae (Venkatesan et al., 2007), ho fana ka tlhahiso ea hore CORT e eketsehileng ho tsoa ponelisong ea koae e ka fokotsa neurogeneis le ho feta, leha sena se ntse se lokela ho lekoa. Maemo a monoamine a Hippocampal ha a fetohe nakong ea bokhachane kapa ea ho pepesoa ha koae (Lubin et al., 2003; Olazabal et al., 2004), e bonts'a hore liphetoho tse ka bang teng ts'ebetsong li ka its'etleha ho tšoaea CRF le CORT. Ntle le moo, litekanyetso tsa OT li fokotsehile hippocampus ho litoeba tsa moroetsana le kamora ho hlaha kamora ho pepeseha ho sa sebetseng hoa gestational cocaine (Johns et al., 1997a; Lubin et al., 2001), e ka fanang ka tlhahiso ea ts'ebelisano le CRF le CORT.
Amygdala e holisitsoeng
Amygdala e atolositsoeng e kenya letsoho ho sebetsanang le maikutlo (haholo tšabo le matšoenyeho), e nchafatsa ho kenya letsoho hoa litho tsa motor motor (Alheid, 2003; Koob le Volkow, 2010) hape o ka nka karolo ho kopantseng tlhaiso-leseling ea cortical le ts'ebetso ea axis ea HPA. Amygdala e atolositsoeng e na le CeA, medial amygdala (MeA), li-amygdala tse atolositsoeng, BNST le likarolo tsa NAc (tsa medial le caudal)Alheid, 2003). CeA le BNST li na le khokahano e kopanyang le PVN mme ke mohloli o ikemetseng oa CRF (Alheid, 2003). Ho pepesetsoa ha Cocaine ho hlahisa phetoho ea nako e telele mesebetsing ea CRF libakeng tsena (Corominas et al., 2010). Phekolo ea cocaine e sa foleng e na le litlamorao tse khutšoane le tsa nako e telele karabong ea methapo ea khatello ea maikutlo ka ho eketsa ts'ebetso e itšetlehileng ka CRF ho amygdala le BNST ho arabela khatello ea maikutlo ho banna (Kash et al., 2008); leha ho le joalo, litlamorao tsa eona ho basali ha li hlake hantle. Ho sebetsa lipakeng tsa CRF ho bile le tšusumetso ho neuroadaptation nakong ea mofuta o sa feleng oa koae le ho khutlisoa ha moputso oa koae (Corominas et al., 2010). Le ha boholo ba mosebetsi ona bo tsepamisitse maikutlo ho tlotseng koae, ho fana ka maikutlo a hore tsela e sa lekanyetsoang ea ho pepesa li-CRF e tšoaea. Ntle le moo, tokollo e fanoeng ea NE, e ka fetoloang ke ho pepesetsoa koae, ho BNST ho arabela khatello ea maikutlo e ka phahamisa matšoenyeho a eketsang boleng ba moputso oa lithethefatsi ka ts'ebeliso e mpe (Aston-Jones le Harris, 2004; Koob le Volkow, 2010). Ka kakaretso, lintlha tsena tsa bopaki li tšehetsa karolo ea sebaka sena e le ntlha ea bohlokoa e kopanyang pakeng tsa moputso le khatello ea khatello kelellong. Acoca cocaine e ngata e ka eketsa OT ho amygdala (Elliott et al., 2001), ha kalafo ea koae e sa foleng nakong ea kemolo e fokotsa ho amohela ha OT ho BNST le amygdala kamora ho qala (Johns et al., 2004; Jarrett et al., 2006).
Ho ferekana mesebetsing e atolositsoeng ea amygdala ho ka ba le litlamorao tse kholo ho MB. Ho kenngwa tshebetsong ha libaka tsa amygdala le BNST ho ka lebisa ho fokotseha ha MB (Rasia-Filho et al., 2000; Walker et al., 2003; Bosch et al., 2005). Haholo-holo, ts'ebetso ea MeA e ka thibela matamo ho atamela malinyane. Ho feta moo, matamo a mouse a tšoauoa ka ho se tsotelle ha bo-mme a na le polelo e phahameng ea c-FOS ho MeA le CeA ha e bapisoa le matamo a taolo (Numan, 2007; Gammie et al., 2008a). MeA le cortical amygdala (CeA) li hlahisoa ka ho pepesehela malinyane nakong ea beke ea pele ho pelehi, empa eseng ka ho pepesehela lithahasello tsa malinyane.Fleming et al., 1994a; Fleming le Walsh, 1994b; Stack et al., 2002). Basyati ea khale ea basolateral (BLA) ha e sebetse ho fihlela PPD3 mme e arabela litšupiso ho PPD10, e lumellana le karolo ea eona ho ithuteng cue (Pego et al., 2008). OT ho AMY e bohlokoa bakeng sa ho laola ho tšoenyeha le boits'oaro bo bobe ba ho mmeela, mme e eketseha kamora ho pepesetsoa ha koae e sa foleng (Bosch et al., 2005; McMurray et al., 2008). Ka lebaka la liphetoho tse rarahaneng tse etsahalang amygdala e atolositsoeng nakong ea pelehi, ho ka etsahala hore tšebeliso ea lithethefatsi e fetileng e ka sitisa tsela e tloaelehileng ea ts'ebetso ea polasetiki.
Tšebelisano ea khatello ea maikutlo le likarete tsa moputso
Ho bohlokoa, khatello ea maikutlo e fetola mokoloto oa moputso. Leha thekiso ea khatello ea maikutlo mosebetsing oa potoloho e le eona e shebileng ho tšoaea hoa CRF ho amygdala e atolositsoeng, ts'ebetso ea GC le eona e bohlokoa. Ho sithabela maikutlo ho sa feleng ho eketsa ho pepeseha hoa glutamatergic le ho sebetsa ha synaptic khetla ea NAc le VTA e ts'oanang le se bonoang ka mor'a ho pepesetsoa ha psychostimulant (Meshul et al., 1998; Campioni et al., 2009; Lodge le Mohau, 2005). Liphetoho tse bakoang ke Cocaine ts'ebetsong ea VTA le tokollo ea NAc DA li its'etleha ho CRF le CORT (ka bobeli)Cleck et al., 2008; Kash et al., 2008). Li-GC li ka fetolela kutloelo-bohloko ho DA ho li-neuron tsa NAc, haholo-holo likotong tsa lactating (Der-Avakian et al., 2006; Byrnes et al., 2007). Karolo ea li-GCs ho hlokofatseng NAc ho li-psychostimulants e kanna ea ba ea bohlokoa haholo, ha ho fanoa ka palo e kholo ea GC e potolohang nakong ea kemolo le ho beleha.Byrnes et al., 2007). CREB e hatisitsoeng e bile le tšusumetso liphetoho tse phehellang bokong kamora ho pepesetsoa ke lithethefatsi tsa lithethefatsi kapa liketsahalo tse sithabetsang tsa tikoloho, 'me li hlahisoa ho potoloha moputso (Briand le Blendy, 2010). Maemo a eketsehang a CREB ea phosphorylated e ka ba mochine oa bohlokoa litlamorao tse mpe le tse sa feleng tsa taolo ea koae le sensitization (Briand le Blendy, 2010), le ha khatello ea maikutlo e khutlisetsoa ho likarabo tse nang le boemo ba koae (cocaine)Kreibich le Blendy, 2004). Tšitiso ea TREEPO ts'ebetso e ka lebisa tlhokomelisong e phahameng ea litlamorao tsa cocaine, empa e sitisa monyetla oa boits'oaro bo amanang le lithethefatsi ho latela likarolo tsa khatello ea kelello (Dinieri et al., 2009), ha a ntse TREEPO ho tepella maikutlo ho ka ntlafatsa litlamorao tsa koae (Kreibich et al., 2009; Briand le Blendy, 2010). Tšitiso ea ho saena ka CRF receptor 1 e ka thibela ntlafatso e matlafatsang ea likarabo tse nang le boemo ba koae, hammoho le keketseho e nyarosang ea phallo ea phosphorylated CREB (Kreibich et al., 2009). Ha li kopantsoe hammoho, lintlha tsena li fana ka maikutlo a hore liphetoho tse bakoang ke koae ka khatello ea kelello li ka sebelisana 'moho le liphetoho lipuisanong tsa meputso ho ama karabelo ea' m'ae.
Kamora nako ho bohlokoa ho hlokomela tlhahiso ea hore ts'ebetso ea ts'ebetso ea allostatic e hloka tumellano e kopanetsoeng pakeng tsa hippocampus, amygdala, le PFC (McEwen le Gianaros, 2011). Kaha ho hlakile hore libaka tsena li bohlokoa bakeng sa karabelo ea khatello ea maikutlo le ts'ebetso le tlhokomelo ea MB, mme li anngoe hampe ke ho pepesoa ha koae, ba totobatsa libaka tse lokelitsoeng ho etsa lipatlisiso tse eketsehileng mehlaleng e pepesitsoeng ea botsoali.
Ho ba motsoali ke batsoali ba nang le khatello ea maikutlo
Joalokaha re hlokometse mona, ho potoloha ha neano ea meputso le lits'ebeletso tsa khatello li kenya letsoho ho qalisong ea tšebeliso ea lithethefatsi, hammoho le ts'ebeliso e tsoelang pele le boits'oaro bo latelang. Lisebelisoa tse ngata tsa bohlokoa tsa neural ka har'a lipotoloho tsena le tsona li bonoa lithutong tsa ho ba motsoali, li fana ka maikutlo a hore li-circuits tsena tse fetang holimo tsa neural li teng e le mokhoa oo lithethefatsi tsa tlhekefetso li ka fetolang boitšoaro ba botsoali. Liphetho tsena tse amanang le mohlala o hlahisitsoeng kenyelletsong ea tekolo ena li hlahisoa ho setšoantšo 6. Karolo ea ho qetela ea mohlala oa rona e tiisa hore maemong a lemallo, litokelo tsa masea lia sithabetsa ho e-na le moputso, le hore maemo a phahameng a khatello ea maikutlo a eketseha a lakatsa lintho tsa tlhekefetso tseo ka boiphihlelo bo fetileng bo amanang le phokotso ea litšusumetso tse mpe. Ka hona, ketso ea ho hlokomela lesea e ka khothaletsa mekhoa ea ho batla lithethefatsi hajoale e sebelisa bo-mme, le ho baka bocha bo-mme.
Tšoantšiso 6
Setšoantšo sa 6. Karolo ea phetoho ea likamano kamanong ea tšebeliso ea lithethefatsi le ho ba motsoali. Ts'ebeliso ea lithethefatsi e tsejoa ho baka liphetoho tse ngata bokong (mabokose a teal le metsu). Liphetoho tsena li ka susumetsa e mong le e mong (metsu e khubelu e phehiloeng habeli) ka ho holisa kapa ho fokotsa liphetoho ho latela maemo a bophelo le a tlhaho. Habohlokoa, liphetoho tsena li bontšitsoe ka boikemelo ho kenya letsoho boits'oarong ba tlhokomelo ea botsoali mme ha e sitisoa ke tšebeliso ea lithethefatsi, ho fella ka maikutlo a fokotsehileng a boleng bo putsa ba masea le khatello e matla ea maikutlo. Karabelo ea khatello ea maikutlo e ka ba e lekaneng ho tsosa takatso ea lithethefatsi e lebisang ho ts'ebeliso e tsoetseng pele ea lithethefatsi le ho khutlela ho bo-'mè ba khaotsoeng. Ntle le moo, lithethefatsi tse batlang ho fokotsa khatello ea maikutlo le tsona li ka holisa leqhubu la ho se tsotelle.
Boemong ba methapo, kamano pakeng tsa ho ba motsoali, bokhoba ba khatello ea maikutlo le khatello ea maikutlo li bongoaneng. Leha ho le joalo, ts'ebeliso ea lithethefatsi e kopantsoe hantle le dalili tse amanang le khatello ea maikutlo (Sinha, 2001), mme ho tloha kelellong ho totobalitsoe e le boits'oaro bo fetisang ba motsoali (Webster-Stratton, 1990). Maemo a ntseng a eketseha a khatello ea maikutlo ho ba motsoali ho lumeloa hore a amana le lisebelisoa tse sa lekaneng (mohlala, chelete, botsitso ba maikutlo) ho sebetsana le litlhoko tsa ho hlokomela ngoana, le hore sena se matlafatsoa ho bo-mme ba lemaletseng, ba tlalehang maemo a phahameng a khatello ho feta ao e seng ntho e sebelisoang ke bo-'mè (Kelley, 1998). Tlhahisoleseling ena e fana ka maikutlo a hore bo-'mè ba lemaletseng lithethefatsi ba ka bontša phetoho e mpe taolong ea khatello ea maikutlo nakong ea pelehi. Patlisiso e eketsehileng e bonahetse khatello ea khatello ea botsoali joalo ka mokena-lipakeng oa lintho tse ka bakang likotsi tsa bakhachane le tšusumetso ea bona boits'ebetsong ba botsoali.Suchman le Luthar, 2001). Lithuto tsena tsa mantlha li tšehetsa mohopolo oa ho ba motsoali e le khatello ea maikutlo, 'me joale re tla nka kamano lipakeng tsa khatello ea maikutlo le ho lakatsa tseo e leng karolo ea mohlala oa rona.
Bopaki bo akaretsang bo bonts'itse hore batho ba nang le takatso e matla haholo ha ba pepesitsoe khatello ea maikutlo ba ka khona ho khutla hape, le hore tšebeliso ea lithethefatsi e fana ka mokhoa o le mong oa taolo ea khatello ea maikutlo, leha e le ho ba le ts'ebetso e mpe, e itšehlang thajana (Sinha le Li, 2007). Lithutong tsena, barupeluoa ba pepesetsoa khatello ea maikutlo le liphetoho phetohong ea hemodynamic li bapisoa le ho pepesetsoa boemo bo sa nkeng khatello ea maikutlo. Ho batho ba sa sebeliseng thepa, ho pepesetsoa khatello ea maikutlo ho bontšitsoe ho eketsa karabelo ea hemodynamic libakeng tsa ka pele (1), ho kenyeletsoa le MPFC e nepahetseng le ventral ACC; le (2) maoto le matsoai a mahareng a mats'oafo, ho kenyelletsa le sebaka sa kamore ea ho phomola, le lesehali, thalamus, patudate le putamen, mme a tloha libakeng tsa hippocampal le parahippocampal (Sinha et al., 2004). Mosebetsi o mong o etsoa ke sehlopha se le seng sa lipatlisiso (Sinha et al., 2005) e bonts'itse hore le ha liphetoho tse ling ho karabo ea hemodynamic li tloaelehile ho batho ba tloaelehileng le lithuto tse itšetlehileng ka koae, taolo e phetseng hantle e bonts'a ts'ebetso e kholo ho ACC, ha barupeluoa ba sebelisang k'hok'heine ba ntse ba na le ho fokotseha hoa ts'ebetso sebakeng sena se atolositsoeng ho ea morao. bokapele ba ka pele. Bangoli ba fetolela phapang ena ho ACC e sebetsang mabapi le phapang ea taolo ea maikutlo le taolo ea kelello pakeng tsa lihlopha tsena tse peli le kamano ea mesebetsi ena ho litloaelo tse tlatsetsang. Ho pheta seo ba se fumaneng pejana, ho pepesa khatello ea maikutlo ho matlafalitse ts'ebetso libakeng tsa hippocampal le tsa parahippocampal ka har'a taolo e phetseng hantle, empa karabelo ena e ne e le sieo ho barupeluoa ba ts'ebeliso ea koae, eo ho fapana le moo ba bonts'itseng karabelo e matla seterekeng se kopaneng sa dorsal striatum le caudate. Ts'ebetso sebakeng sena sa morao-rao e lumellana hantle le lintlha tsa litlaleho tse labalabelang, tse lumellanang le karolo ea sebopeho sena sa bokhoba. Keketseho ea tšebetso ho PFC e nepahetseng ea dorsolateral, hammoho le sebaka se ka morao sa ho sesa le se phahameng ka ho fetesisa, se ikopantseng le litlaleho tse eketsehang tsa tlaleho ea boithati le khatello ea karolo ho barupeluoa ba sebelisang koae. Litlamorao tsa liphumano tsena ke hore, ha bokhoba bo fetolela karabelo ea khatello ea maikutlo, ts'ebetso ena e tsamaisana le litlaleho tsa boithati ba litakatso tsa lithethefatsi, e fana ka maikutlo a kamano e pakeng tsa ho lakatsa, khatello ea maikutlo le bokhoba ba tahi. Sena se boetse se totobatsoa ke ho fumana hore ts'ebetso e ntseng e eketseha libakeng tse kenyelletsang PFC ea morafo, kamora ho imeloa kelellong, ho bolela nako ea ho khutla hape, e tsamaellanang le tšebeliso ea lithethefatsi ketsahalong ka 'ngoe, hammoho le palo ea matsatsi a tšebeliso ea lithethefatsi e etsahetseng kamora ho oela hape (Sinha le Li, 2007). Letlotlo le lengata, le leholo haholo ho qaqisa mona, le se le qalile ho fumana mekhoa ea limolek'hule le mekhoa ea ts'ebetso ea ts'ebetso ea ts'ebetso ea boits'oaro bo tšoanang le bo tsosang khatello ea maikutlo mehlaleng ea liphoofolo. Bohlokoa ho li-hypotheses tsa rona ke lintlha tse fanang ka maikutlo a hore DA le CRF ke limolek'hule tse bonts'ang matla ho VTA, amygdala e atolositsoeng, le PFC (Erb, 2010; Van den Oever et al., 2010; Bohlale le Morales, 2010), hammoho le ho hokahana le liphetoho lipakeng tsa moroalo o matla haholo.
Ho fumana ho pepesetsoa khatello ea maikutlo ka bobeli ho baka karabelo ea boko tse ka khethollang batho ba lemalloang ho batho ba sa lemalleng le hore mesebetsi ea neural e lumellana le litakatso le ho khutlela morao e bontša bohlokoa ba ho ba tlokotsing ea khatello ea maikutlo molemong oa ho lemalla bokhoba. Haholo-holo, lithuto tsena li bonts'a hore ho pepesetsoa khatello ea maikutlo ho holisa takatso ea litakatso tse hlahisang boits'oaro ba ho batla lithethefatsi le ho oela hape. Ha re kenyelletsa sephetho sena tlhahlobisong ea joale, re fana ka maikutlo a hore mekhoa ea ho ba motsoali e tla etsa hore khatello e tšoanang e be teng (mohlala, Kelley, 1998) e ka eketsang boits'ebetso bo batlang lithethefatsi ho 'm'a ea lemaletseng lithethefatsi, ka hona e ka kenya letsoho boits'oarong bo sa tsotelleng bo amanang haholo le ts'ebeliso ea lithethefatsi ho bo-mme (mohlala, Cash le Wilke, 2003). Ke sepheo sa lithuto tsa rona tsa thuto ea mantlha tsa bophelo ba mantlha le ba batho ho hlahloba sena ka mokhoa o matla.
fihlela qeto e
Thutong e hlahisitsoeng mona re supile tlatsetso ea moputso le tsela ea khatello ea maikutlo ho potolohong ea neural ea ho ba motsoali, re totobatsa ho feto-fetoha ha litsela tsena ka bokhoba. Re hlalositse bokhoba e le khatello ea moputso le khatello ea methapo, lits'ebetso tse ts'oanang tse fetotsoeng bakeng sa botsoali ho eketsa ho nyenyefatsa ha masea. Re fana ka maikutlo a hore maemong a tlatsetsang, mekhoa ea ho ba motsoali ha e na moputso ka tsela eo e neng e ka ba ka eona mme e ka ba khatello ea maikutlo, e ka bakang tšebeliso e mpe ea litekanyetso tsa ho feto-fetoha ha khatello ea maikutlo e ka lebisang ho eketseheng ha lithethefatsi le boits'oaro bo sa tsotelleng ba ho ba motsoali. Ha re ntse re shebile haholo ts'ebeliso ea bokhoba ba koae, melaoana ea mohlala ona e tla ts'oara mekhoa e meng e tlatsetsang, ka lebaka la likarolo tse tloaelehileng tsa khatello le ts'ebetso ea moputso ho qaleng le ho boloka ts'ebeliso ea lithethefatsi. Ho feta moo, ho lemoha likamano tsa 'm'a le ngoana esale pele e le mohloli oa khatello ea maikutlo ho tla ba bohlokoa ha ho nahanoa mekhoa e nepahetseng ea kalafo bakeng sa thibelo le kalafo ea tlhekefetso ea lithethefatsi tsa motsoetse (mohlala, Pajulo et al., 2006; Asman et al., 2008). Sena se totobatsoa ke litheko tse phahameng tsa ho khutla pele ho nako ke bo-mme ba tlohelang lintho tse sebelisang tlhekefetso nakong ea bokhachane, ba ts'ehetsang taba ea hore nako ea ho beleha e hlahisa e le nako e itseng ea tlokotsi ea khatello ea maikutlo bo-mme ba morao tjena. Ho joalo, puisano e hlahisitsoeng mona e fana ka maikutlo a hore mekhoa ea phekolo e lebisang taolo ea khatello ea maikutlo e ka ba ea bohlokoa bakeng sa bokhoni ba ho ba motsoali, ho boloka bokhoba ba tahi le ho fokotsa kotsi ea tlhekefetso ea bana le ho se tsotelloe. Lipheo tse ka bang teng tsa neurobiological li ka kenyelletsa CRF le OT kaha li bonts'itsoe e le litsamaiso tsa bohlokoa tsa ho supa khatello ea maikutlo, bokhoba le motsoali.
Tlhōlisano ea Tlhaloso
Bangoli ba bolela hore lipatlisiso li ne li etsoa ka ho se be le likamano leha e le life tsa khoebo kapa tsa lichelete tse ka nkoang e le khohlano e ka 'nang ea e-ba le thahasello.
lumela hore baa fokola
Bangoli ba ne ba tšehelitsoe ke Award Number P01DA022446 (Josephine M. Johns) oa Setsi sa Naha sa Tlhekefetso ea Lithethefatsi. Litaba ke eona feela boikarabello ba bangoli 'me ha ho hlile ha e emetse maikutlo a semmuso a Setsi sa Naha sa Ts'ebeliso ea Lithethefatsi kapa Likolo tsa Naha tsa Bophelo.
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Mantsoe a bohlokoa: bokhoba, moputso, khatello ea maikutlo, botsoali, preclinical, botho
Caring: Rutherford HJV, Williams SK, Moy S, Mayes LC le Johns JM (2011) Ho ferekana ha potoloho ea motsoali ea motsoali e motšehali ka ts'ebetso ea tlatsetso: ho ts'oaroa ha meputso le lits'ebetso tsa khatello ea maikutlo. Ka pele. Psychiki 2: 37. Doi: 10.3389 / fpsyt.2011.00037
E amohetse: 22 February 2011; Papali e ntseng e emetse e hatisitsoeng: 06 April 2011;
E amohetsoe: 09 June 2011; E phatlalalitsoe marang-rang: 06 Julayi 2011.
E hlophisitsoeng ke:
Rina Eiden, Univesithing e Buffalo, USA
Hlahloba by:
Lisa Carter, Univesithi ea Illinois e Chicago, USA
Kelly Lambert, Koleji ea Randolph-Macon, USA
Copyright: © 2011 Rutherford, Williams, Moy, Mayes le Johns. Ena ke sengolo se bulehileng se fihlellang laesense e seng e ikhethileng pakeng tsa bangoli le Frontiers Media SA, e lumellang tšebeliso, phetiso le tlhahiso liphatlalatsong tse ling, ha feela bangoli ba mantlha le mohloli ba ngolisitsoe 'me maemo a mang a Frontiers a tsamaisoa.
* Ngoliso: Helena JV Rutherford, Setsi sa Thupelo sa Bana sa Yale, Univesithi ea Yale, 230 South Frontage Road, New Haven, CT 06520, USA. lengolo-tsoibila: [imeile e sirelelitsoe]; Sarah K. Williams, Lefapha la Psychiatry, Univesithi ea North Carolina-Chapel Hill, 436 Taylor Hall, CB # 7096, Chapel Hill, NC 27599, USA. lengolo-tsoibila: [imeile e sirelelitsoe]
;