Tse ling tsa phekolo ea kelello ea lilemong tsa bocha: liphetoho tsa boqapi ba boko, liketso tse sebetsang, le mekhoa ea boitšoaro (2011)

1. Selelekela

Nako ea bocha ke nako eo ka eona batho ba bonang liphetoho tsa 'mele ea bona, ba nang le litabatabelo le litakatso tse ncha,' me ba iphumana ba e-na le tokoloho e kholo, boipuso le boikarabello. Le ha e hlalosoa ka tsela e fapaneng, bongoana ka kakaretso bo nkuoa e le ho qala ha ho kena bohlankaneng kapa boroetsaneng ebe bo fela ha motho a nka karolo mesebetsing ea batho ba baholo (Dahl, 2004; Spear, 2000). Nako ea ho kena bohlankaneng kapa boroetsaneng e kenyelletsa ho hola ho eketsehileng, liphetoho sebopeho sa 'mele, nts'etsopele ea manonyeletso le litho tsa botona le botšehali tsa botona le botšehali, le liphetoho tsa pelo le matšoafo-hangata li etsahala ho tloha lilemong tsa 10 ho isa ho 17 ho banana le 12 ho 18 ho bashanyana.Falkner le Tanner, 1986). Ha sena se etsahala mocha o feta maemong a fapaneng a kelello, boits'oaro, le kelello. Liphetoho tse fapaneng tsa bocha ha li qale hammoho ebile li fela hammoho, ka hona papali ea phetoho e mabapi le phetoho ea bongoana ea bocha e na le phephetso. Ho ithuta lilemo tsa bocha ho tšoana le ho thunya moo ho shebiloeng, 'me bafuputsi ba qapa lihlopha tsa "lilemo tsa bocha" tsa lilemo tse fapaneng le maemo a ntlafatso. Ho feta moo, ho tloha bohareng ba 19^th ka 20^th lekholong, lilemong tsa pele-pele tsa ho ea khoeling li bonoa lefats'eng la bophirima (Falkner le Tanner, 1986; Tanner, 1990). Ts'ebetso ea thuto e nka nako e telele ebile batho ba ikemiselitse ho ema nako e telele pele ba qala mesebetsi, ho nyaloa le ho ba le bana (Dahl, 2004). Kahoo, bolelele ba nako ea bocha ha bo tsitsitse (mme esale bo atoloha) mme nako e lumellana le lits'ebetso tse ngata tsa ntlafatso ea bioloji, e hlalosoa ka mokhoa o fapaneng ho latela maemo a kelello le a boits'oaro. Ka likhopolo tsena, likhatiso tse hlahlojoang mona li hlalositse boholo ba lilemo tsa batho e le lilemo tse mashome a mabeli tsa bophelo, ho litšoene e le lilemo tse peli ho isa ho tse 'ne, le likoranteng e le beke tse nne ho isa ho tse supileng.

Leha ho e-na le liphetoho tse hlakileng, ho tsebahala hore nakong ena liphetoho tse kholo lia etsahala, ho kenyelletsa le mefuta e fapaneng ea boits'oaro bo bonoang mefuta eohle ea mefuta. Ho na le boitšoaro bo eketsehileng sechabeng (Csikszentmihalyi et al., 1977), boqapi le maikutlo a ho batla (Adriani et al., 1998; Stansfield le Kirstein, 2006; Stansfield et al., 2004), mekhoa ea ho nka kotsiSpear, 2000; Steinberg, 2008), ho hloka botsitso maikutlong (Steinberg, 2005), le impulsivity (Adriani le Laviola, 2003; Litho tsa et al., 2003; Fairbanks et al., 2001; Vaidya et al., 2004). Likamano tsa lithaka lia ata, 'me ho ba le litšekamelo tse kholo tsa ho batla liphihlelo tse monate le tse monate (Nelson et al., 2005). Ho eketseha hoa bohlanya le ho batla lintho tse ncha ho ka feto-fetoha le maemo, kaha mekhoa ena e ka ntlafatsa menyetla ea mocha ea ikemetseng ea ho fumana lijo le molekane (Spear, 2010). Sechabeng sa mehleng ena, likarolo tsena li ka amahanngoa le ho beha likotsi tse sa hlokahaleng. Ka hona, lilemo tsa bocha li nkuoa e le nako ea boitšoaro bo kotsing: Bacha ba na le monyetla oa ho leka koae le lithethefatsi tse seng molaong le joala; khanna ka mokhoa o sa tsotelleng; etsa thobalano e sa sireletsehang; Ho ba le likhohlano pakeng tsa batho (Arnett, 1992; Arnett, 1999; Litho tsa et al., 2003; Spear, 2000). Ho ipeha kotsing ea bokhachane ho ka etsahala hangata ka lihlopha (mohlala, likotsi tsa makoloi), ha boitsoaro bo itseng bo nkoa bo amoheleha ho lithaka tsa hao (mohlala, thobalano e sa sireletsehang, ts'ebeliso ea lithethefatsi) (Steinberg, 2008), le maemong a qosoang ka maikutlo (Figner et al., 2009). Kahoo, leha bacha ba pholohile mathata a bophelo bo botle ba bongoana ho fokola le litekanyetso tsa ba shoang li imena habeli ho feta ho bana ba tlileng pele ho selemo (Dahl, 2004).

Ntle le likotsi tse eketsoeng tsa E tloaelehileng Kholo ea bocha, e boetse ke nako eo matšoao a mafu a fapaneng a kelello a hlahang hangata, ho kenyeletsa le mathata a ho feto-fetoha ha maikutlo, mathata a ho ja le mafu a kelello a kang schizophrenia (Paus et al., 2008; Pine, 2002; Sisk le Zehr, 2005; Volkmar, 1996). Nakong ena ho na le letoto la liphetoho tse ngata tse tsamaisang ntho e 'ngoe le e' ngoe ho tsoa liphatlalatsong tsa lipontšo tsa lihormone tse qalang ho kena bohlankaneng kapa boroetsaneng.Sisk le Zehr, 2005), ho eketsa bokhoni ba kutloisiso le liphetoho tse susumetsang (Doremus-Fitzwater et al., 2009; Luna et al., 2004). Ho utloisisa hantle hore na boko bo hola joang bongoaneng, le ho amana le liphetoho tse joalo litloaelong le maemong a bophelo, ho bohlokoa haholo bophelong ba sechaba. Mona re hlahloba tse ling tsa liphetoho tsa bocha, le li-neurodevelopment tsa bocha 'me re tšohla mefuta e mmaloa e li hokahanyang, ho kenyelletsa le monahano oa rona oa ts'ebetso e ntlafalitsoeng ea ts'ebetso.

2. Boitšoaro ba bocha

Boithuto ho litoeba le batho li bonts'itse hore bacha ba bontša "khetho e potlakileng" e hlalosoang e le khetho ea moputso o nyane o etsahalang kapele ha moputso o moholo o lieha, joalo ka ha o lekantsoe le mesebetsi ea theolelo ea ho lieha (Adriani le Laviola, 2003; Steinberg et al., 2009). Hoa hlokomeleha hore lithutong tsa batho feela bacha ba lilemong tse tlase ba bonts'a phapang ena; ka ho lieha ho theola litheko tsa batho ba baholo ka lilemo tsa 16-17 (Steinberg et al., 2009). Bacha ba lilemong tsa bocha le bona ba iponahatsa ba le maemong a phahameng ho feta batho ba baholo, ba batona ba bonts'a maemo a holimo ho feta a basali.Zuckerman et al., 1978). Ho batla ho utloisoa bohloko ke "tlhoko ea maikutlo, lipono tse fapaneng, le maikutlo a rarahaneng ..." (Zuckerman et al., 1979, leq. 10), tse ka hlahang ka boikemelo, kapa hammoho le ho ts'oaroa. Ho batla ho khothatsoa ho hoholo nakong ea bongoana ho isa bohareng ba lilemo tsa bocha, le ha taolo ea tšusumetso e bonahala e ntlafala hanyane ka hanyane lilemong tsa bocha, a fana ka maikutlo a hore ba tšehelitsoe ke mekhoa e fapaneng ea likokoana-hloko (Steinberg et al., 2008). E lumellana le bopaki ba motho ba ho batla ho teba ha bacha lilemong tsa bocha, litoeba tsa bongoaneng li rata ho hlaka (Adriani et al., 1998; Douglas et al., 2003; Stansfield et al., 2004), bonts'a mofuta o mocha o bululetsoeng ka mokhoa o mocha (Stansfield le Kirstein, 2006; Sturman et al., 2010), 'me u qete nako e ngata ho phenya matsoho a bulehileng ka mochini o moholo o moholo ho feta batho ba baholo (Adriani et al., 2004; Macrì et al., 2002).

Litloaelo tsa bacha tsa ho batla boiphihlelo ba bona, leha li le kotsing ea ho lemala kapa ho senyeha sechabeng, li ka lebelloa haeba matla a bona a ho lekola menyetla kapa sephetho sa sephetho a ntlafala. Bokhoni ba kelello bo ntse bo tsoela pele ho ntlafala ka nako ena (Luna et al., 2004; Spear, 2000). Ho ea ka Piaget, nako ea ts'ebetso e hlophisitsoeng, e amanang le mabaka a sa utloahaleng, e fihla khōlong e felletseng nakong ea bocha (Schuster le Ashburn, 1992,. Ho phehella ha mohlala, ho tsoella ha bacha ho ba le 'bamameli ba nahanang' hammoho le 'pale e iqapetsoeng' ea maikutlo a ikhethang, ho ka etsa hore ba lumele hore ba ikhethile 'me ho ba fa maikutlo a ho hlaseloa (Arnett, 1992; El mosa, 1967). Leha ho le joalo, ke ntlafatso e hlakileng ea kelello e hlahang ho tloha lilemong tsa bocha ho ea pele (Luna et al., 2004; Spear, 2000), esita le bana ba banyenyane ba bonts'a kutloisiso e nepahetseng e hlakileng ea monyetla (Acredolo et al., 1989). Ho feta moo, ho na le bopaki bo fokolang ba hore bacha ba ipona e le kotsi e ka kenang kapa ea e nyenyefatsa; Ebile, hangata ba kotsing e kholo, joalo ka monyetla oa hore ba tla ima pele selemo se fela, ba kenele teronkong, kapa ba bolaoe ba le banyenyane (de Bruin et al., 2007). Kamora nako, tlhaloso e ngoe le e ngoe e lemohileng bakeng sa ho beha bacha kotsing ea ho kena bohlankaneng e tlameha ho ikarabella bakeng sa hore bana ba beha likotsi tse fokolang mme leha ho le joalo ba holile tsebong ea bongoaneng.

Ntle le moo, karohano ea boitšoaro ba bacha e ka amana le liphapang tsa maano a kelello. Khopolo-taba e 'ngoe e bitsoang "fantize the trace theory," e re ho fapana le ho haelloa ke bokhoni ba ho lemoha, bacha ba sebetsana le maemo a kotsi / melemo ea liqeto tse hlakileng ho feta batho ba baholo. Ka mokhoa o makatsang, bacha ba ka phela ka mokhoa o hlonephileng ho feta batho ba baholo ka ho etsa likhakanyo tse hlakileng tse fapaneng tsa khetho e fapaneng, empa sena se ka lebisa ho ipeheng kotsing e kholo (Meroalo et al., 2008). Ho latela Melao le basebetsi-'moho (2008), ka nts'etsopele re hatela pele ho tsoa ho ho sebelisa “mantsoe a mangata” ho ea ho "fantisi" ea boemo bo phahameng ba "grafic" e ts'oereng lintlha le mohala o tlase ntle le lintlha. Sena se ntlafatsa bokhoni ba ho nka liqeto ebile se re khetha ho khetha liqeto tse kotsi kaha re tla qoba sephetho se sebe ntle le ho lekola menyetla ea seabo. Mohlala, ho fapana le bacha, batho ba baholo ba khetha khetho e tlatselletsang phaello kapa ho fokotsa tahlehelo ka lebaka la litekanyetso tse tšoanang tse lebelletsoeng (Meroalo et al., 2008). Ka kakaretso, khopolo ea hore khetho ea bacha e ka bonts'a liphapang tsa leqheka la ho lemoha - empa e se bofokoli polelong ea pheletso-ea makatsa. Boithuto ba morao-rao ba 'mele ba nakong e tlang le boithuto ba' mele ba ho etsa liqeto tsa bocha bo ka una molemo ka ho nahana ka monyetla oa hore phapang ka tatellano e nepahetseng ea tšebetso ea neural, esita le ka har'a likarolo tse tšoanang tsa boko, hammoho le boemo ba kopanyo pakeng tsa libaka tse fapaneng, li ka tsamaisa mefuta e meng ea tlhahlobo ea kelello.

Ho hloka masene ha bacha ho ka bakoa ke ho fapana ha bona hore na ba kotsing le moputso joang. Tlhaloso e ngoe ke ea hore bacha ba lilemong tsa bocha ba ba le tšusumetso e mpe e fetisisang le maikutlo a sithabetsang, mme ba ka ikutloa ba sa natefeloe ke ho susumetsoa ke boleng bo tlase kapa bo itekanetseng ba tšusumetso. Bacha ba ka hona ba batla sepheo sa ho ba le matla a maholo a ho khotsofatsa tlhoko ea bona ea moputso (bona Spear, 2000). Sena se tšehetsoa ke lithuto tse bonts'ang phapang ea boleng ba hedonic ba likarabo tsa sucrose ho batho ba baholo ba bapileng le bacha. Hang ha mohopolo oa sucrose o feta ntlha ea bohlokoa, boleng ba hedonic bo fokotseha haholo; leha ho le joalo litheko tse joalo ha li hlahisoe haholo kapa ha li teng ho bana le lilemong tsa bocha.De Graaf le Zandstra, 1999; Vaidya et al., 2004). Tlhaloso e 'ngoe ke hore bacha ba na le kutloisiso e kholo ea thepa e matlafatsang ea mohopolo o monate. Mohlomong monyetla ona o lumellana le mehlala ea liphoofolo moo bacha ba sebelisang tharollo ea ho ikemela (Vaidya et al., 2004), khetha likamore tse neng li amana le tšebelisano ea sechaba (Douglas et al., 2004), mme o bonts'e bopaki ba boleng bo phahameng ba khothatso bakeng sa lithethefatsi tse kang nicotine, joala, amphetamine le cocaine ho feta batho ba baholo (Badanich et al., 2006; Brenhouse le Andersen, 2008; Shram et al., 2006; Spear le Varlinskaya, 2010; Vastola et al., 2002). Sena ha se bonoe kamehla, leha ho le joalo, (Frantz et al., 2007; Mathews le McCormick, 2007; Shram et al., 2008), le ho eketseha ha khetho ea lithethefatsi tsa bongoana ho ka amana le ho fokotsa maikutlo le likotsi tsa morao-rao le ho tlohela (Little et al., 1996; Moy et al., 1998; Schramm-Sapyta et al., 2007; Schramm-Sapyta et al., 2009). Ka mokhoa o ts'oanang, bacha ba ka etsa boits'oaro bo kotsi ha tlhahlobo ea bona ea litlamorao e sa susumetse kapa e sa nyenyefatse (kapa haeba thabo ea ho ipeha kotsing e beha boitšoaro bo joalo).

Ntho e 'ngoe e ka bakang karohano ea boitšoaro ba bacha ba lilemong tsa bocha ke phello ea maikutlo (maikutlo a phahameng, maikutlo, maikutlo le maikutlo a itseng a itseng) ka boitšoaro. Ho se lumellane ka boitšoaro ho ka hlaha haeba bacha ba e-ba le maikutlo a fapaneng ka tsela e fapaneng, kapa haeba maikutlo a fapaneng a susumetsa ho etsa liqeto nakong ena ea maikutlo le boits'oaro bo matla (1).Arnett, 1999; Buchanan et al., 1992). Hangata maikutlo a batho a nka hore ho etsa liqeto hoa utloahala. Le ha hona ho ka ba joalo maemong a mang (haholo-holo ha litaba tsa maikutlo li sa tsamaellane kapa li sa amane le maemo a qeto), mosebetsi oa morao-rao o lekotse hore na maikutlo a ka ntlafatsa liqeto tse itseng joang. Mohlala, "somatic marker hypothesis" e re maemong a makatsang, ts'ebetso ea maikutlo e ka tataisa boitsoaro (Damasio, 1994). Mosebetsi oa ho becha oa Iowa o ne o etselitsoe ho leka ho etsa liqeto tlas'a maemo a ho hloka bonnete (Bechara et al., 1994). Batho ba nang le marapo a PFC kapa li-amygdala ba thatafalloa ke ho amohela leano la ho qoba kotsi, ba fana ka maikutlo a hore khaello ea ho kenyelletsa tlhahisoleseling ea maikutlo e ka lebisa liqetong tse mpe (Bechara et al., 1999; Bechara et al., 1996). Bacha le batho ba baholo ba ka fapana ka tsela eo ba kopanyang tlhahisoleseling ea maikutlo le liqeto: Bacha ba ka se tsebe ho fetolela kapa ho kopanya litaba tsa maikutlo tse amehang, kapa ba sa sebetse hantle ho theheng mekhatlo e joalo. Cauffman et al. (2010) haufinyane tjena ho ile ha lekoa bana, bacha le batho ba baholo tsebong e fetotsoeng ea Iowa Gossip Task; ba hlokometse hore ha bacha le batho ba baholo ba ntlafatsa liqeto tsa bona tsa ho etsa liqeto ha nako e ntse e feta, batho ba baholo ba entse sena ka potlako e kholo. Phuputso e 'ngoe e bonts'itse hore ke feela ho tloha lilemong tsa bocha ho fihlela bonyenyaneng bao lithuto li ntlafalitseng ts'ebetso ea bona ea papali ea chelete, le hore ntlafatso ena e tsamaisana le ponahalo ea likarolo tsa mmele tse hlasimolohileng.Crone le van der Molen, 2007). Liphetho tsena li bontša hore bacha ba ka se sebetse hantle ho theha kapa ho toloka mofuta oa tlhaiso-leseling e nepahetseng e hlokahalang ho qoba liqeto tse kotsi.

Ho latela Melao le basebetsi-'moho (2008) ho se tšoane ka tšebetso e sebetsang ea likheo tsa botšehali ho etsa hore bacha ba kenelle habonolo liphellong tse ka fokolisang tsa ho etsa liqeto. Maemong a ho tsosa takatso e phahameng, ho fokotseha hoa boitšoaro ho ka etsa hore motho a tlohele ho "beha mabaka" ho fetoha "ntho e sebetsang" kapa e sa tsitsang. Ba boetse ba pheha khang ea hore tloaelo ea bongoaneng ea ho etsa lits'oants'o tse ngata tsa tlhaiso-leseling e etsa hore sena se be bonolo, le ha litekanyetso le litheko tsa ts'ebetso ea "gist" ea motho e moholo e le leseli ka ho fetelletseng ho mmuso o hlasimolohileng (Meroalo et al., 2008). Ba bang ba boetse ba pheha khang ea hore boits'oaro ba bongoana bo ka ama ka ho khetheha maemo a bohloko bo phahameng ba maikutlo (Dahl, 2001; Spear, 2010). Phuputso ea morao tjena ke Figner le basebetsi-'moho (2009) o lekile ka ho otloloha mohopolo ona a sebelisa mosebetsi o lekantseng kotsi e ka tlisoang maemong a fapaneng a tumello. Bacha le batho ba baholo ba ile ba etsa mosebetsi oa "Card Card Task", moo boemo ba kotsi bo mamelloang bo neng bo hlahlojoa tlasa maemo a makhopho a maholo le a fokolang mme ba ntse ba sebelisa lintlha tse fapaneng tse joalo ka boholo ba phaello / tahlehelo le monyetla oa bona. ). Bacha ba ile ba beha likotsi tse ngata ho feta batho ba baholo feela ba maemong a phahameng a takatso, 'me maemong ana, bacha ba ne ba sa angoe ke ho phaellana le monyetla oa ho lahleheloa ke monyetla, ba fana ka maikutlo a hore ts'ebeliso ea tlhaiso-leseling e entsoe ke bacha lilemong tsa maemo a phahameng a mofets'e (Figner et al., 2009).

Ka kakaretso lithuto tsena li bonts'a hore le ha bacha ba nahana le ho itshwara joalo ka batho ba baholo, maemong a itseng ho na le phapang leano la bona la ho lemoha le / kapa karabelong ea bona kotsing le moputsong, haholoholo tlasa maemo a phahamisang maikutlo. Liphetoho tsena tsa boits'oaro li kanna tsa bonts'a kholo e kholo ea khokahano ea boko - ho kenyeletsoa likarolo ho PFC, basal ganglia, le li-system tsa neuromodulatory (mohlala dopamine) - tseo e leng tsa bohlokoa boits'ebetsong bo susumetsoang (Lethathamo 1).

  

Lethathamo 1

Phapang mokhoeng oa boitšoaro ba bocha le "neurodevelopment" ea methapo

3. Adachecent organic neurodevelopment

Boko ba bongoana bo fetoha haholo lithutong tse tebileng tsa morphology. Boithuto ba ho nahana ka sebopeho sa batho bo bonts'itse hore ho pholletsa le bongoana ho na le tahlehelo ea lintho tse bosoeu lilemong tsa bocha, ka litheko tsa lintho tse tsamaeang le bohlooho ka likarolo tsa sekhukhu sa nakoana le PFors ea dorsolateral e etsahalang nakong ea bocha.Gogtay et al., 2004; Sowell et al., 2003; Sowell et al., 2001; Sowell et al., 2002). Phokotso ea litaba tsa Grey e boetse e bonahala ka har'a li-striatum le likarolo tse ling tsa subcortical (Sowell et al., 1999; Sowell et al., 2002). Liphetoho tsena li ka amana le ho pongoa ha li-synapses tse ngata nakong ena ho tsoa lithutong tsa liphoofolo (Rakic ​​et al., 1986; Rakic ​​et al., 1994), leha ba bang ba belaella khokahano ena e le bouton ea synaptic e etsang karolo e fokolang feela ea bophahamo ba cortical (Paus et al., 2008). Menahano ea batho e boetse e pepesitse hore taba e tšoeu e eketseha ka ho ba mocha lilemong tsa lipampitšana tsa cortical le subcortical fiber (Asato et al., 2010; Benes et al., 1994; Paus et al., 2001; Paus et al., 1999), e hlahisoang ke myelination e eketsehileng, axon caliber, kapa ka bobeli (Paus, 2010). Liphetoho mekhoeng ea khokahano le tsona li etsahala lilemong tsa bocha. Mohlala, ho mela le ho hola hoa axonal ho 'nile ha bonoa makaleng a hokahanyang amygdala le sepheo sa cortical (Cunningham et al., 2002), le mehato e ntseng e eketseha ea litaba tse tšoeu e bonoa lipakeng tsa PFC le striatum le libakeng tse ling (Asato et al., 2010; Giedd, 2004; Gogtay et al., 2004; Liston et al., 2006; Paus et al., 2001; Sowell et al., 1999).

Ka tekanyo e ntle, litekanyetso le lipatlisiso tsa morao-rao li bontšitse liphapang tse ngata tsamaisong ea methapo ea bongoana. Bacha ba tloaetse ho hlahisa haholo dopaminergic, adrenergic, serotonergic le endocannabinoid receptors ho pholletsa le libaka tse ngata tse lateloang ke ho roka maemong a batho ba baholo (Lidow le Rakic, 1992; Rodriguez de Fonseca et al., 1993). Ba hlahisa D1 le D2 dopamine receptors ka maemo a hloahloa ka lipheo tsa subcortical tse kang dorsal striatum le nucleus accumbens, leha ba bang ba sa fumane polelo e fokolitsoeng ea batho ba baholo sebakeng sena sa morao-rao (Gelbard et al., 1989; Tarazi le Bal)arini, 2000; Tarazi et al., 1999; Teicher et al., 1995). Nakong ea bocha, ho boetse ho na le liphetoho tlhahisong ea dopamine le phetiso ea mali, hammoho le bopaki ba liphetoho bakeng sa litlamorao tse theohelang tsa receptor-ligand binding (Badanich et al., 2006; Cao et al., 2007; Coulter et al., 1996; Laviola et al., 2001; Tarazi et al., 1998). Ka ts'ebetso, ho na le bopaki bo tsoang ho likhoto tse sa anthieng hore mosebetsi o ikatisang oa li-neuron tsa dopamine neurons ka nako ea bocha ebe oa fokotseha (McCutcheon le Marinelli, 2009). Liphetoho tsa nts'etsopele ea mesocorticolimbic dopamine circry le ts'ebetso li ka baka liphapang tse ling tsa boitšoaro bo susumetsoang ka kakaretso, hammoho le ho ipeha kotsing ea ho lemalla le ho lemalla. Liphuputso tse 'maloa li hlokometse litla-morao tsa psychomotor tsa lithethefatsi tse matlafatsang liphoofolong tsa bocha empa li matlafalitse kapa li khothaletsa tse tšoanang (Adriani et al., 1998; Adriani le Laviola, 2000; Badanich et al., 2006; Bolanos et al., 1998; Frantz et al., 2007; Laviola et al., 1999; Mathews le McCormick, 2007; Spear le Brake, 1983). Ho fapana le seo, bacha ba amehile haholo ka litla-morao tsa cataleptic tsa li-neuroleptics (mohlala, haloperidol), e leng bahanyetsi ba li-receptor tsa dopamine (Spear le Brake, 1983; Spear et al., 1980; Teicher et al., 1993). Ba bang ba re sebopeho sena, hammoho le tlhahlobo e ntseng e eketseha ea ho batla lintho tse ncha, li bontša hore sistimi ea dopamine ea lilemong tsa bocha e haufi le "siling e sebetsang" qalong.Litho tsa et al., 2003).

Ho na le bopaki bo bongata bo bontšang hore tekanyo ea thabo le khatello ea methapo ea kutlo e fapane haholo lilemong tsa bocha ha li bapisoa le batho ba baholo. Mehato ea GABA, e ka sehloohong inhibitory neurotransmitter bokong, e eketseha ka mokhoa o hlakileng ka nako ea bocha ho qoqothong la rat (Hedner et al., 1984). Polelo ea li-receptor tse sebetsang tsa NMDA tse sebetsang ka li-neurons tse potlakileng (tse nahanoang hore ke li-inhibitory interneurons) li fetoha haholo ho PFC ea bacha. Ka nako ena boholo ba li-interneurons tse potlakileng haholo ha li bontše maqhubu a li-synaptic NMDA receptor-mediated (Wang le Gao, 2009). Ho kenyelletsa phello ea modulatory ea liphetoho tse tlamang tsa dopamine-receptor nakong ea bocha (O'Donnell le Tseng, 2010). Ke ka nako ena feela moo ts'ebetso ea dopamine D2 receptors e eketsang tšebetso ea li-interneuron (Tseng le O'Donnell, 2007). Ntle le moo, tšebelisano pakeng tsa dopamine D1 receptor activation le phetoho ea li-receptor tsa NMDA nakong ea bocha, li lumella depolarizations tsa Plata tse ka tsamaisang polasetiki e itšetlehileng ka maemo (synaptic plasticity)O'Donnell le Tseng, 2010; Wang le O'Donnell, 2001). Liphetoho tsena tsa bocha tsa dopamine, glutamate, le GABA li bontša phapang e kholo ea tšebetso ea neural bokong ba mocha. Ts'ebetso tsena kaofela li bohlokoa lits'ebetsong tsa kelello le maikutlo. Ho se sebetse ha bona ho amehile ho mafu a mangata a kelello a amanang le mathata a ho feto-fetoha le maikutlo le bokhoba ba lefu la kelello.

4. Adachecent functional neurodevelopment

Liphuputso tse entsoeng ka neuroimaging li bontšitse phapang e teng tšebetsong ea bongoana ea batho ba sebetsang libakeng tse 'maloa tsa bokapele. Liphapang tsena li shebiloe haholo-holo libakeng tsa boko tse kenyeletsang bohlokoa ba maikutlo (mohlala, amygdala) li kopanya leseli la maikutlo le maikutlo bakeng sa ho kopanya litebello tsa boleng (mohlala, orbitofrontal cortex), 'me li bapala likarolo tse fapaneng ho susumetsa, ho khetha liketso, le ho ithuta mohoanto (mohlala. striatum). Ha ho bapisoa le batho ba baholo, bacha ba na le karabelo e fokotsehileng ea hemodynamic ka cortex ea morao-rao ea morao-rao le tšebetso e eketsehang molemong oa ho fumana moputso (Ernst et al., 2005; Galvan et al., 2006). Ba bang ba fumane tšebetso e fokotsehileng maemong a nepahetseng a nepahetseng le litheolelo tse atolositsoeng ka nako ea tebello ea moputso, ho se na phapang e amanang le lilemo tse amanang le lilemo ka mor'a ho fumana sephetho (Bjork et al., 2004). Mosebetsing oa ho etsa liqeto, bacha ba fokotse ts'ebetso ea kahare ea letsoho mme ba tlohella li-orbitof Pambal / ventrolateral PFC bapisoa le batho ba baholo nakong ea likhetho tse kotsi (Eshel et al., 2007). Bacha ba boetse ba kenya tšebetsong matla a bona a ho kenella kahare ho feta batho ba baholo joalo ka ha ba ne ba ipeha kotsing e kholo nakong ea papali ea Stoplight - e leng se susumetsoang ke khatello ea lithaka e sa bonahaleng (Chein et al., 2011).

Boithuto bo 'maloa bo hlokometse ho se sireletsehe ha litsamaiso tsa taolo ea lilemong tsa bocha, hammoho le ts'ebetso e mpe ea boits'oaro (Luna et al., 2010). Mohlala, nakong ea mesebetsi e hlokang ho thibeloa ha karabelo ea pele (tšebetso ea eona e ntlafatsang le lilemo), bacha ba ekelitse ts'ebetso ea PFC maemong a mang le ho fokotsa mosebetsi ho ba bang (Bunge et al., 2002; Rubia et al., 2000; Tamm et al., 2002). Nakong ea ts'ebetso ea taolo ea antisaccade ea tlhokomeliso ea kelello, lilemong tsa bocha (empa eseng batho ba baholo) ts'ebetso ea boemo ba moea ea bongoana e ile ea fokotsoa ha ba ntse ba shebile setšoantšo se bonts'ang haeba moputso o le teng nakong ea nyeoe e fanoeng, empa e ne e ts'ebetsoe ho feta molekane e moholo nakong ea tebello ea moputso (Geier et al., 2009). Kahoo, bacha ba kenya tšebetsong likarolo tse tšoanang tsa tlhalohanyo le tse amohelehang joalo ka batho ba baholo, leha ho le joalo hangata ka bongata bo fapaneng kapa sebopeho sa maemo le sa nakoana, kapa maemo a kopaneng a kopaneng (Hwang et al., 2010).

Boholo ba puisano ea pakeng le ea lipakeng tsa selehae le kopano ea methapo ea methapo li ka bapala karolo ea mantlha kholong ea boitšoaro ba bacha. Ho na le kamano e tobileng lipakeng tsa methati ea taba e tšoeu ea pele-pele, e ntseng e eketseha ka nako ea bocha, le ts'ebetso ea taolo ea inhibitory (Liston et al., 2006). Nts'etsopele ea litaba tse tšoeu e amana ka kotloloho le kopanyo e ntlafalitsoeng ea tšebetso ea libaka tse tsoetsoeng ka bohlooho, ho fana ka maikutlo a ts'ebetso ea marang-rang e phatlalalitsoeng ka ho fetisaStevens et al., 2009). Sena se netefatsoa ke boithuto ba hore, ba sebelisang khokahano ea ts'ebetso ea mmuso e phomotsang mmuso hammoho le tlhahlobo ea litšoantšo, ba hlokometse phetoho e tsoang khokahanong e kholo le linthong tse nang le tšebeliso ea methapo ho tsa marang-rang tse neng li hokahane haholo ho feta libaka tsohle ho ba motho e moholo ho sa tsotelehe hore na sebaka ke eng (Fair et al., 2009). Ka mokhoa o ts'oanang, keketseho e amanang le lilemo bakeng sa ts'ebetso e kopantsoeng ea libaka tsa pele le tsa parietali e ts'ehetsa ts'ebetso e ntlafalitsoeng ea taolo ea thibelo ea thibelo e holimo-tlase mosebetsing o thibelitsoeng pele.Hwang et al., 2010). Nts'etsopele ea litaba tse tšoeu, ho kuta ha li-synapses ka potlako (tseo boholo ba tsona e leng khokahano ea thabo ea lehae), le liphetoho tsa tšebetso ea mahareng a lehae li ka tsamaisa tšebelisano-kholo e eketsehileng lipakeng tsa likarolo tsa boko ka nts'etsopele. Ketsahalo e sa abeloang haholo ho bacha e bontšitsoe ketsahalong e 'ngoe ea taolo ea kelello (Velanova et al., 2008). Ka nako e ts'oanang, senya pontšo ea ts'ebetso e sa amaneng le ts'ebetso ea mosebetsi e fokotseha ka nts'etsopele (Durston et al., 2006). Kahoo, mokhoa oa batho ba baholo oa ho sebelisa marang-rang a abeloang haholo o tsamaisana le ts'ebetso e fokotsehileng e sa sebetseng, e bonts'ang ts'ebetso e kholoanyane mohlaleng le bohareng ba ts'ebetso ea cortical.

Lithuto tsa electrophysiological li boetse li fumane bopaki ba nts'etsopele ea likarabo tsa neuronal le tšebetso e hloahloa ea lehae le ea nako e telele ka nako ea bocha. Mohlala, Contingent Negative Variation, e leng ketsahalo e mpe e amanang le boemo ba motlakase nakong ea litokiso tsa karabelo, e hlaha bongoaneng mme e tsoela pele ho ba kholo ka bocha.Bender et al., 2005; Segalowitz le Davies, 2004). Sena se nahanoa hore se bonts'a liphapang tse amanang le lilemo ho phatlalatso ea ts'ebetso ea PFC ea tlhokomelo le taolo ea makoloi a phahameng (Segalowitz et al., 2010). Phetoho e 'ngoe e amanang le lilemo tsa elektroniki ke tsoelo-pele ea tlhoro e matla (P300) e batlang e le 300 ms kamora ho ea ho khothatsoa. Pato ea P300 e holileng ha e hlahe ho fihlela lilemo tse ka bang 13 (Segalowitz le Davies, 2004). Kamora nako, Negativity e amanang le phoso ke motlakase o fosahetseng o kenang ka holim'a cortex ea anterior nakong ea liteko tsa mesebetsi e fapaneng. Le ha ho e-na le phapang e itseng lilemong tsa ponahalo ea eona, e bonahala e fihla lilemong tsa bohareng ba bocha (Segalowitz le Davies, 2004). Liphetho tsena li fana ka bopaki bo eketsehileng bakeng sa ntlafatso ea ts'ebetso ea pele ho cortical nakong ea bohlankana. Segalowitz le basebetsi mmoho le bona ba fumane hore tekanyo ea molumo oa lipontšo tsa motlakase oa bana le bacha hangata e ne e le tlase ho feta ea batho ba baholo. Sena se ka bakoa ke ho se sireletsehe hoa mosifa kapa ho se tsitse ha motho ka mong libakeng tsa boko tse hlahisang matšoao ana (Segalowitz et al., 2010). E ka boela ea bonts'a tšebelisano ea methapo ea kutlo ea bongoana kahare le lipakeng tsa boko. Tlhaloso ena e tsamaisana le mosebetsi o etsoang ke Uhlhaas le basebetsi-'moho (2009b), eo ho eona ho neng ho tlalehiloe li-electroencephalograms (EEGs) baneng, lilemong tsa bocha le ho batho ba baholo nakong ea mosebetsi oa ho amohela sefahleho. Ba bone ba fokotse matla a "theta" (4-7 Hz) le "gamma band" (30-50 Hz) matla a ho hlasimolla bocheng ha a bapisoa le batho ba baholo. Ntle le moo ho ne ho e-na le phase-sychrony ea nako e telele ho theta, beta (13-30 Hz), le lihlopha tsa gamma, hammoho le ts'ebetso e ntlafalitsoeng ea mosebetsi ho batho ba baholo. Li-oscillations tsa EEG li bakoa ke ho fetoha ha maemo ho thabo ea thabo 'me ho nahanoa hore li hlakola nako ea tlhahiso ea seiketsehi (Meketso, 2005). Mehato ea synchrony lihlopha tse ikhethileng tsa maqhubu li nolofalletsa puisano lipakeng tsa lihlopha tsa methapo, 'me e ka ba bohlokoa lits'ebetsong tse ngata tsa pono le temohisiso (Uhlhaas et al., 2009a). Ka hona, liphumano tsena ke bopaki ba ntlafatso e ntlafalitsoeng ea tšebetso ea lehae le puisano e ntlafalitsoeng ea libaka ho tloha bocha ho ba motho e moholo (Uhlhaas et al., 2009b).

Mokhoa o mong o sebetsang oa ho hlahloba liphetoho tsa tšebetso ea maqhubu a lilemong tsa bocha ke oa Ka vivo ho rekota hoa motlakase ho tsoa ho li-arrrode tse kentsoeng tsa elektrose ho liphoofolo tse ts'oereng tse tsohileng. Mokhoa ona o thusa motho hore a tlalehe tšebetso ea li-neuron ka bomong hammoho le menyetla e meholo ea tšimo. Re sa tsoa etsa thuto e joalo, eo ho eona likhoto tsa bocha le ba baholo ba ileng ba etsa lintho tse bonolo tse tataisang sepheo sa bona (Setšoantšo sa 1a) ha lirekoto li nkuoe ho orbitofrontal cortex. Ha bacha le batho ba baholo ba ne ba etsa boitšoaro bo tšoanang, ho ne ho bonoa liphapang tse amanang le lilemo tse amanang le neural, haholoholo ho putsa (Sturman le Moghaddam, 2011). Sena se bontša hore leha boitšoaro bo ka bonahala bo ts'oana, "cortex ea bongoana" e maemong a fapaneng ho feta ea batho ba baholo. Ka ho khetheha, li-neurons tsa bongoaneng tsa orbitofrontal cortex li ile tsa nyakalla haholo moputsong, ha karolo ea li-neurons tsa bongoana e ne e le nyane haholo ka nako eo le linthong tse ling mosebetsing (Setšoantšo sa 1b). Joalo ka neural inhibition e bohlokoa bakeng sa taolo ea nako e nepahetseng ea li-spikes le ho kenyelletsa tšebetso e kopantsoeng ea oscillatory (Cardin et al., 2009; Fries et al., 2007; Sohal et al., 2009), thibelo e amanang le lilemo tsa bongoana e amanang le cortex neuronal inhibition e amanang le mosebetsi e ka amana ka kotloloho le phapang e kholo ea neural encoding e boneng thutong ena mme e hlalositsoe ke ba bang. Kamora nako, boholo ba mosebetsi bacha ba bonts'a phapang e kholo ea nako ea teko ea maiketsetso, e ka bonts'ang lerata le tlase la lerata ho cortex ea lilemong tsa bocha. Ka hona, ha cortex ea pele e nts'etsopele, tšitiso e eketsang phasic maemong a yuniti e le 'ngoe e ka ts'ehetsa ho hokahano ho hoholo ha hare le lipakeng tsa selekane le ho sebetsa hantle.

  

Setšoantšo sa 1

A) Schematic ea mosebetsi oa boits'oaro. Lirosa li sebelitse ka hare ho kamore e sebetsang e tloaelehileng. Nyeoe e 'ngoe le e' ngoe e qalile ka ho qala ha leseli la cue ka sekoting sa nosetso (Cue). Haeba lehare le kenelletse ka sekoting sena lebone le ntse le le (Poke) ...

5. Neurobehaivereal Hypotheses

Ka liphetoho tsohle tsa bocha tsa bongoana, ke eng e etsang hore ho be le phapang e atileng nakong ee? Likarolo tse fetileng li bonts'a bopaki ba liphetoho tse fapaneng tsa bongoaneng tsa bocha tsa bongoana le liphapang tse amanang le lilemo tsa boits'oaro le bofokoli. Mona re hlahisa li-hypotheses tse 'maloa kapa mehlala e hokahanyang ka ho hlaka phapang ea bongoana boitshwarong bo susumetsoang, nts'etsopele ea sechaba, le boitšoaro bo sitisang le kholo ea li-circuits tse khethehileng tsa neural (Lethathamo 2).

  

Lethathamo 2

Li-hypotheses tsa Neurobehaivereal tse kopanyang liphetoho tsa boitšoaro tsa bongoana le kholo ea kholo ea kelello

Ntlafatso ea bocha ea marang-rang a ntlafatso ea tlhahisoleseling ea sechaba ke mohlala o le mong o hokahanyang nts'etsopele ea bacha le liphetoho tsa boko (Nelson et al., 2005). Moralo ona o hlalosa likarolo tse tharo tse hokahantsoeng tse sebetsang ka mokhoa o ikhethileng oa sebopeho se hlophisitsoeng: sebaka sa ho iphumanela (inferior occipital cortex, inferior le anterior temporion cortex, intraparietal sulcus, fusiform gyrus, le supignal tempcus), sethala se sebetsang. mookotaba oa bethe ea stria terminalis, hypothalamus, le orbitofrontal cortex maemong a mang), le node ea taolo ea kelello (likarolo tsa cortex ea pele). Mokhoa oa ho lemoha o khetholla hore na ts'usumetso e na le tlhaiso-leseling ea sechaba, e ntlafalitsoeng hape ke semelo se amehang se susumetsang tšusumetso joalo ka bohlokoa ba maikutlo. Node e tataisang kelello e tsoela pele ho sebetsa tlhahisoleseling ena, e etsa mesebetsi e rarahaneng e amanang le ho lemoha maemo a kelello a batho ba bang, e thibela likarabo tsa pele, le boitšoaro bo tataisoang ke sepheo (Nelson et al., 2005). Liphetoho tsa bocha maikutlong le tšebelisanong ea methapo ena li susumetsoa ka botebo ho matlafatsa liphihlelo tsa sechaba le maikutlo, li susumetsa ka matla liqeto tsa bocha, hape li kenya letsoho ho hlaha ha psychopathologies nakong ena (Nelson et al., 2005).

Mohlala oa "node"Ernst et al., 2006) e fana ka maikutlo a hore mokhoa o tsoetseng pele oa nts'etsopele ea libaka tsa boko o tšehetsang ts'ebetso e sebetsang le taolo ea kelello, le botsitso lipakeng tsa bona, li ka fokotsa tekano ea bacha ba lilemong tsa bocha. Moetso ona o boetse o ipapisitse le tšebetso ea likarolo tse tharo tse tsamaellanang le likarolo tse itseng tsa boko. Maemong ana node e ikarabellang bakeng sa mokhoa oa moputso (ventral striatum) e lekana le node ea thibelo ea kotlo (amygdala). Mokhoa o feto-fetohang oa molumo (preortalal cortex) o ama tsusumetso e lekanyelitsoeng ea matla ana a hlaselang, mme boits'oaro bo kotsi bo tla hlaha ka mokhoa oa ho qetela oa kamohelo ea lipalo. Ho ea ka mohlala ona, maemong a kenyelletsang khoebo e pakeng tsa takatso ea lijo le boiphetetso, mokhoa ona o atile haholo ho bacha. Hyperacaction kapa hypersensitivity ea mokhoa oa ho fumana moputso e kanna ea fetoloa ke ts'ebetso likarolo tse ling tsa kort ea pele, leha ho le joalo tsoelo-pele ea eona ea lilemong tsa bocha ha e lumelle ho lekola ho lekola le ho laola taolo (Ernst le Fudge, 2009).

Casey le basebetsi mmoho ba tiisa hore ho se tšoane ho ntlafatseng hoa bongoana bo tlang pele ho cortex le lintho tse sa bonahaleng (mohlala, ventral striatum le amygdala), le khokahano lipakeng tsa tsona, li ka ikarabella bakeng sa boitšoaro ba bocha ba boitšoaro (Casey et al., 2008; Somerville le Casey, 2010; Somerville et al., 2010). Nakong ea mosebetsi o kenyelletsang ho amohela litheolelo tse fapaneng tsa meputso, boholo ba tšebetso ea bongoana bokong ba nko e ne e ts'oana le ea batho ba baholo (leha e le ka bongata bo boholo) athe mokhoa oa ts'ebetso ea bongoaneng o ne o shebahala o tšoana le oa bana ho feta batho ba baholo.Galvan et al., 2006). Kholo e tsitsitseng ea lits'ebetso tsa subcortical le ho se sireletsehe ha cortex ea pele, e bohlokoa ho taolo ea kelello, e ka lebisa ho kholo le ho feta tekong ea bohlankana mabapi le maikutlo a ho batla le ho ipeha kotsing. Senotlolo mona, joalo ka mohlala oa node e khahlisang, ke mohopolo oa ho se lekane ho pakeng tsa tikoloho nakong ea bohlankana, ho fapana le bongoana ha libaka tsena kaofela li sa le hole le ho ba motho e moholo ha tsona tsohle li se li holile.Somerville et al., 2010). Moetso ona o ts'oana le sebopeho sa Steinberg, moo ho fokotsoa ha kotsi ea ho ba motho ea lilemong tsa bocha ho ba motho e moholo ka lebaka la nts'etsopele ea litsamaiso tsa taolo ea kelello, likhokahanyo li thusa ho kopanya maikutlo le ho ama libakeng tsa cortical le subcortical, le phapang ea ho se lekane ka moputso kapa kutloelo-bohloko (Steinberg, 2008).

Sehlooho sa mantlha sa mefuta ena ke hore ho bacha, ho fapana ka maikutlo, boemo, kapa phello ea lintho tse etsoang libakeng tse fapaneng le tsa taolo ea kelello. Ho ipapisitsoe le datha le bopaki bo bong, re tiisa hore liphapang tse joalo e ka ba sephetho sa ts'ebelisano ea "neuronal" le ts'ebetso ea ts'ebetso ho bana ba lilemong tsa bocha tse bonts'ang ka lebaka la phetisetso ea tlhahisoleseling e sa sebetseng lipakeng tsa libaka le ho se leka-lekane ha tlhahiso ea neuronal le tšitiso lipakeng tsa libaka tsa bohlokoa tsa boko. , joalo ka orbitof Pambal cortex le likarolo tsa sehlopha sa basal ganglia. Joalokaha ho hlalositsoe pejana, a vitro mosebetsi o bonts'itse liphetoho tse kholo mokhoeng oa polelo oa li-receptor tse fapaneng, le litlamorao tsa ts'ebetso ea li-receptor, ho kenyelletsa le karabelo ea li-innethons tse potlakileng tsa spiking tsa dopamine le ts'usumetso ea li-receptor tsa NMDA. Liphetoho tse joalo li ka lebelloa ho ama botsitso ba ho nyatsa le ho sitisa le ho hokahana ha lihlopha tsa neuronal. Ha ts'ebetso ea "interneuron" e potlakileng e potlakisa ho laola nako e nepahetseng ea ts'ebetso ea neural le ho kenella ha oscillations, liphetoho tsa nts'etsopele tsa ts'ebetso ea bongoana le karabelo ea tsona ho li-neuromodulators tse kang dopamine e ka ba karolo ea tse ling tsa liphapang tsena tse amanang le lilemo. Ka lebaka la sena, ts'ebetso ea neano ea bongoana e kanna ea hlophiseha hantle, ea sa tsotelleng, le ea lehae, mme hape e ka tsepamisa maikutlo ho litlamorao tsa ts'ebeliso ea meputso, bocha, kapa tse ling tse susumetsang. Phokotso ea kopanyo e pakeng tsa libaka le tikoloho, e sitisoang ke ho fetella hoa tumellano, e ka ikarabella bakeng sa ts'ebetso e sa sebetseng haholo e bonoang lithutong tsa monahano. Tloaelo e boletsoeng esale pele ea bacha ea ho khetha khetho e kotsi maemong a mahlonoko a maikutlo e ka amana le puisano e fokotsang ea lipakeng tsa setereke (mohlala, ho se atlehe ha "cortex ea pele" ho hlakola matšoao a "ho ea" ka hara basal ganglia), le ho fetelletsa litaba ts'ebetso le / kapa ho fokotsa ts'itiso ho litloaelo tse poteletseng maemong a boitšoaro bo susumetsoang, joalo ka ha re bone nakong ea tebello ea moputso mokokotlong oa orbitofrontal.

6. Summary

Ha re ntse re ithutile haholoanyane ka liphetoho tse ikhethileng tsa bocha le boits'oaro tsa bongoana li se li khothalelitsoe. Ntho e bohareng ho boholo ba tsena ke maikutlo a hore ts'ebetso ea "neuronal process" e kenang pele ho cortex le likarolo tse ling tsa cortical le subcortical, hammoho le tšebelisano ea bona, e lebisa ho boitšoaro bo khetholloang ho fihlela menyetla, moputso, le ho nkeha maikutlo maikutlong nakong ea bohlankana. Mosebetsi oa morao-rao oa nts'etsopele ea lipotoloho tsa methapo ea methapo le tšebetso ea tsona e fetolang le li-system tsa neuromodulatory nakong ea bocha li kanna tsa fana ka leseli la hore na ke hobaneng ha maloetse a kang schizophrenia a bonahala ka nako ena. Re sebelisa mekhoa e kang fMRI ho batho le lirekoto tsa motlakase liphoofolong tsa batho ba sebetsang, re se re qala ho tseba hantle hore na bacha ba etsa joang moputso le likarolo tse ling tsa boits'oaro bo fapaneng le batho ba baholo. Ho etsa joalo ke bohato ba bohlokoa ba ho netefatsa likotsi tsa ts'ebetso ea bongoana le ho utloisisa mafu a kelello a mafu a kelello a hlahang nakong ena.

Mongolo o botlaaseng ba leqephe

References