Boko ba bocha (2008)

Dev Rev. 2008; 28(1): 62-77. doi:  10.1016 / j.dr.2007.08.003

PMCID: PMC2500212

inahaneloang

Nako ea bocha ke nako ea nts'etsopele e bonts'ang liqeto tse sa amaneng le liqeto tse etsang hore ho be le keketseho e kholo ea likotsi tsa ka boomo le pefo, tšebeliso e mpe ea joala le lithethefatsi, boimana bo sa reroang le mafu a tšoaetsanoang ka tsa thobalano. Litlhaloso tsa setso le tsa kelello tsa setso sa bocha li hlotsoe ho ikarabella bakeng sa liphetoho tse seng molaong tsa boitšoaro bo bonoang bongoaneng, ho amanang le bongoana le ho ba motho e moholo. Tlhahlobo ena e fana ka conceptualization ea biologically e hlakileng ea mekhoa ea neural e tlatselletsang liphetoho tsena tse sa amaneng le litloaelo, e le karabelo e phahamisitsoeng ho lits'usumetso ha taolo ea ts'usumetso e ntse e sa ts'oanehe nakong ena. Boithuto ba morao-rao ba batho le lithuto tsa liphoofolo li fana ka motheo oa tlhaho ea pono ena, tse fanang ka tlhahiso ea phapang e fapaneng ea litsamaiso tsa litefiso tsa litho tsa matsoho tse amanang le litsamaiso tse laolang taolo e tlase nakong ea bocha le ho ba motho e moholo. Mokhoa ona oa nts'etsopele o ka eketsoa le ho feta lilemong tsa bocha ba nang le tebello ea ho nka likotsi, o eketsa menyetla ea litholoana tse mpe.

Keywords: Bacha, Cortex ea pele, bokello ba nyutlelie, ho susumetsa, moputso, nts'etsopele, ho beha kotsi

Ho latela Setsi sa Naha sa Lipalopalo tsa Bophelo, ho na le lefu le fetang la 13,000 la bacha lilemong tsa United States selemo se seng le se seng. Hoo e ka bang 70% ea mafu ana a bakoa ke likotsi tsa makoloi, likotsi tse sa etsoang ka boomo, polao e mpe le ho ipolaea (Eaton et al., 2006). Liphetho tse tsoang ho 2005 National Youth Risk Behaeve Survey (YRBS) li bonts'a hore bacha ba etsa boits'oaro bo eketsang monyetla oa ho shoa kapa bokuli ka ho khanna koloi kamora ho noa kapa ntle le lebanta la setulo, ho jara libetsa, ho sebelisa lintho tse seng molaong, le ho etsa thobalano e sa sireletsehang. ho lebisa ho ima ho sa lebelloang le mafu a likobo, ho kenyeletsa le tšoaetso ea HIV (Eaton et al., 2006). Lipalo-palo tsena li hlakisa bohlokoa ba ho utloisisa likhetho le liketso tse kotsi ho bacha.

Bongata ba likhopolo tsa bongoana le "neurobiological" li ngotsoe lebaka leo bacha ba kenang boits'oarong ba boikhethelo. Thutong ea morao-rao ea lingoliloeng tse mabapi le nts'etsopele ea bongoana ea bongoana, Yurgelun-Todd (2007) e fana ka maikutlo a hore nts'etsopele ea kelello lilemong tsa bocha e amana le katleho e tsoetseng pele ea matla a taolo ea kelello. Ts'ebetso ena e hlalosoa e its'etleha holima kholo ea "cortex ea pele" joalo ka ha e netefalitsoe ke ts'ebetso e eketsehang libakeng tse ikhethileng (Rubia et al., 2000; Boko, Menon, & Reiss, 2002) le tšebetso e fokotsehileng libakeng tse sa sebetseng tsa boko (Brown et al., 2005; Durston et al., 2006).

Mokhoa ona o akaretsang, oa taolo e ntlafalitsoeng ea temohisiso le ho holofala ha karolo ea pele ea cortex, o fana ka maikutlo a keketseho ea morao-rao ho tloha bongoaneng ho ea ho motho e moholo. Leha ho le joalo liqeto le liketso tse nyane tse bonoang lilemong tsa bocha li emela phetoho e seng molaong ea boits'oaro e ka khetholloang ho tloha bongoaneng le ho ba motho e moholo, joalo ka ha ho bonahala ke Setsi sa Naha sa Lipalopalo tsa Bophelo boits'oarong ba bocha le ho shoa. Haeba taolo ea kelello le "cortex" e tlang pele ho nako e ne e le motheo oa boitšoaro bo ikhethileng, joale bana ba lokela ho shebahala hantle haholo kapa ba be mpe haholo ho feta bacha, ha ba fuoa bokhoni ba bona bo sa ntlafalitsoeng ba bokhoni ba pele. Kahoo, ts'ebetso ea pele ho nako feela, e ke ke ea ikarabella bakeng sa boitšoaro ba bocha.

Khakanyo e nepahetseng ea liphetoho tsa kelello le tsa ramatiki nakong ea bocha li lokela ho nka nako ea bocha e le nako ea phetoho ea nako ea bocha (Spear, 2000), ho fapana le ho thubisoa ka nako e le 'ngoe ka nako (Casey, Tottenham, Liston, & Durston, 2005). Ka mantsoe a mang, ho utloisisa nako ena ea nts'etsopele, phetoho ho tsoa le ho tsoa bongoaneng e hlokahala bakeng sa ho khetholla litšobotsi tse ikhethileng mohatong ona oa ntlafatso. Ho theha li-trajectories tsa nts'etsopele bakeng sa ts'ebetso ea kutloisiso le ea neural li bohlokoa ho tšoaea liphetoho tsena le ho hatella litlhaloso ka liphetoho tsa boitšoaro nakong ena. Boemong ba bonohe kapa bo boits'oaro, bacha ba tšoauoa ka mokhoa o sa susumetsang (ke hore, ba haelloa ke taolo ea kelello) le ho ipeha kotsing ka litlatsetso tsena tse sebelisitsoeng ka mokhoa o ts'oanang ntle le kananelo bakeng sa tšekamelo e ikhethang ea nts'etsopele ea eona. Boemong ba methapo ea kutlo, likhopolo tsa batho le lithuto tsa liphoofolo li fana ka maikutlo a hore na methapo ea kutlo e arohaneng le litekanyetso tsa nts'etsopele ea methapo ea kutlo e ka tlasa melaoana ena ea taolo e matla le liqeto tse kotsi.

Re hlahisitse mofuta oa methapo ea kholo ea kholo ea bongoaneng ka har'a moralo ona o thehiloeng ho mehlala ea methapo (Laviola, Adriani, Terranova, & Gerra, 1999; Spear, 2000) le lithuto tsa morao-rao tsa ho nahana ka bongoana (Ernst et al., 2005; Galvan, Hare, Voss, Glover, le Casey, 2007; Galvan et al., 2006). Feie. 1 ka tlase e supa mohlala ona. Ka letsohong le letšehali ke tšobotsi ea setso sa bocha joalokaha e amana hantle feela le ho se sireletsehe ha cortex ea pele. Ka ho le letona ke mohlala oa rona oa "neurobiological" o bonts'ang tsela e bonts'ang likarolo tsa taolo ea maoto le taolo pele ho taolo e tlase li lokela ho nkuoa hammoho. Cartoon e bonts'a li-trajectories tse fapaneng tsa nts'etsopele ea lits'ebetso tsena, ka lits'ebetso tsa maoto le nts'etsopele pejana ho libaka tsa taolo ea pele. Ho ea ka mohlala ona, motho o khetholloe haholo ke libaka tse holofetseng tsa bocha nakong ea bohlankana (ke hore, ho se lekane ha litho tsa maoto le taolo ea pele) ha li bapisoa le bana, bao litsamaiso tsena (ke hore, li-legic le pele) li ntseng li tsoela pele le tsona; le ho bapisoa le batho ba baholo, bao lits'ebetso tsena li holileng ka botlalo ho bona. Pono ena e fana ka motheo oa phetoho e sa amaneng le litheko ho ntlafatseng, ka lebaka la ho hola ha nako ea setho sena sa maoto le taolo ka tlase ho taolo pele. Ka nts'etsopele le boiphihlelo, khokahanyo e sebetsang pakeng tsa libaka tsena e fana ka mokhoa oa taolo e holimo-tlase ea libaka tsena (Hare, Voss, Glover, le Casey, 2007a). Ho feta moo, mohlala o bapisa ho hanana le lipalo tsa bophelo bo botle ba bophelo bo botle nakong ea bocha, le pono e makatsang ea Reyna le Farley (2006) hore bacha ba khona ho beha mabaka le ho utloisisa likotsi tsa boitšoaro boo ba kenang ho bona. Ho latela mohlala oa rona, maemong a hlobaetsang a maikutlo, tsamaiso ea maoto le matsoho e tla atleha ho laola litsamaiso tsa taolo ha e fuoa bokhoni ba eona ba ho hola tsamaisong ea taolo ea pele. Bopaki bo tsoang boithuto ba boits'oaro le boits'oaro ba batho ba ho ts'ehetsa mofuta ona bo fanoa molemong oa liketso maemong a monate le a maikutlo (Galvan et al., 2006, 2007; Hare, Voss, Glover, le Casey, 2007b; Hare et al., 2007a). Ntle le moo, re nahana ka hore na ke hobaneng ha boko bo ka hlaha ka tsela ena le hore na ke hobaneng ha bacha ba bang ba ka ba kotsing e kholo ea ho etsa liqeto tse sa lebiseng liphellong tse mpe tsa nako e telele (Galvan et al., 2007; Hare et al., 2007b).

 

Tlhaloso ea moetlo ea boits'oaro ba bongoana ho khothalelitsoe hore e bakoa ke nts'etsopele ea protortal cortex (A) ea pele. Mohlala oa rona o nahana ka nts'etsopele ea "cortex ea pele" hammoho le likarolo tsa subcortical limbic (mohlala, li-nucleus accumbens) tse kentsoeng ts'ebetsong ea likhetho le liketso tse kotsi (B).

 

Nts'etsopele ea boitšoaro bo lebisoang ho sepheo

Sefika sa nts'etsopele ea kelello ke bokhoni ba ho hatella menahano le liketso tse sa lokang bakeng sa ba tataisoang ke sepheo, haholoholo boteng ba tšusumetso e khothalletsang (Casey, Galvan, & Hare, 2005; Casey et al., 2000b; Casey, Thomas, David-mora, Kunz, & Franzen, 2002a; Casey, Tottenham le Fossella, 2002b). Lithuto tse 'maloa tsa ntlafatso ea khale li bonts'itse hore bokhoni bona bo hlaha bongoaneng le boseeng (Nyeoe, 1972; Flavell, Feach, & Chinsky, 1966; Keating & Bobbitt, 1978; Pascual-Leone, 1970). Litsebi tse ngata tsa thuto li boletse hore nts'etsopele ea kelello e bakoa ke keketseho ea lebelo la ts'ebetso le ho sebetsa hantle eseng ka lebaka la keketseho ea matla a kelello (mohlala, Bjorkland, 1985; Bjorkland, 1987; Nyeoe, 1985). Litsebi tse ling tsa khopolo-taba li kenyellelitse ho theha mekhoa ea "inhibitory" tlalehong ea bona ea nts'etsopele ea kelello (Harnishfeger le Bjorkland, 1993). Ho latela tlaleho ena, tlhokomeliso ea bokhachane e tšoauoa ka ho ts'oaroa ke ts'itiso ho tsoa mehloling ea tlholisano e lokelang ho hatelloa (mohlala, Brainerd & Reyna, 1993; Casey, Thomas, Davidson, Kunz, & Franzen, 2002a; Dempster, 1993; Taemane, 1985; Munakata & Yerys, 2001). Kahoo boitšoaro bo tataisoang ke sepheo bo hloka taolo ea litlamorao kapa ho lieha ho khotsofatsa bakeng sa sephetho mme bokhoni bona bo bonahala bo hola bongoaneng le bonyaneng.

Boitšoaro ba bocha bo hlalositsoe e le bo sa qobellohe ebile bo le kotsi, hoo e ka bang ka mokhoa o tšoanang, leha ho le joalo, tsena li itšetleha ka lits'ebetso tse fapaneng tsa kutloisiso le li-neural, tse fanang ka maikutlo a ikhethang a ikhethang le li-trajectories tse fapaneng tsa nts'etsopele. Haholo-holo, tlhahlobo ea lingoliloeng e bonts'a hore maikutlo a tlatselletsang a fokotseha ka lilemo ho fihlela bongoaneng le bohlankana (Casey et al., 2002a; Casey, Galvan et al., 2005; Galvan et al., 2007) mme e amana le nts'etsopele e tsoetseng pele ea cortex ea pele (Casey, Galvan et al., 2005), leha ho e-na le phapang ea hore na motho ea fuoeng o susumetsoa kapa che, ho sa tsotelehe hore na o lilemo li kae.

Ho fapana le hoo, ho susumetsa / taolo ea kelello, ho nka likotsi ho bonahala ho eketseha nakong ea bohlankana le boroetsana le ho ba motho e moholo ebile ho amana le litsamaiso tse tlase tse tsejoang ho nka karolo ho lekola moputso. Lithuto tsa ho nahana ka batho tse tla hlahlojoa, li fana ka maikutlo a keketseho ea ts'ebetso ea subcortical (mohlala, li-kukusanya) ha ho etsoa liqeto tse kotsi (Kuhnen le Knutson, 2005; Matthews & et al., 2004; Montague le Berns, 2002) e feteletsang ho bacha, e amanang le bana le batho ba baholo (Ernst et al., 2005; Galvan et al., 2006). Liphumano tsena li bonts'a li-trajectories tse fapaneng bakeng sa boitšoaro bo thehiloeng moputsong kapa khokahanong, ka nts'etsopele ea mantlha ea lits'ebetso tsena tse amanang le litsamaiso tse bonts'ang netefatso ea nts'etsopele, le mabapi le ho phahamisa likhetho le liketso tse sa lokang molemong oa bao ba lebisitsoeng ho sepheo.

Bopaki bo tsoang lipatlisisong tsa bongoana tsa kholo ea kholo ea motho

Patlisiso ea morao-rao ea nts'etsopele ea bongoana e thehiloe holima tsoelo-pele ea mekhoa ea neuroimaging e ka sebelisoang habonolo ka ho nts'etsopele ea batho. Mekgwa ena e itshetlehile hodima mekgwa ya matla a ho thuisa a maqhubu a matla a maqhubu a maqhubu a maqhubu a maqhubu a maqhubu a marang-rang (bona Feie. 2) hape e kenyelletsa: MRI ea sebopeho, e sebelisetsoang ho lekanya boholo le sebopeho sa meaho; MRI e sebetsang e sebelisetsoang ho lekanya lipaterone tsa tšebetso ea boko; le infusion tensor imaging (DTI) e sebelisetsoang ho supa khokahano ea lipampitšana tsa fiber tse tšoeu. Bopaki ba moralo oa rona oa nts'etsopele oa tlholisano lipakeng tsa libaka tsa cortical le subcortical bo tšehetsoa ke khokahano ea mesebetsi le tšebetso e lekantsoeng ke DTI le fMRI, ka ho latellana.

Feie. 2    

Mekhoa e tloahelehileng ea ho sebelisa maqhubu a sebelisang boithuto ba nts'etsopele ea batho e bontšitsoe kaholimo. Structural magnetic resonance imaging (MRI) ho hlahisa litšoantšo tsa sebopeho sa boko tse bohlokoa bakeng sa lithuto tsa anatomical le morphometric (A), tlhahiso ...

Lithuto tsa MRI tsa nts'etsopele ea boko ba motho

Boithuto bo 'maloa bo sebelisitse MRI ea sebopeho ho etsa' mapa oa kholo ea kholo ea kholo e tloaelehileng (sheba tlhaiso ka Durston et al., 2001). Leha boholo ba boholo ba boko bo batla bo lekana le 90% ea boholo ba eona ka lilemo tse ts'eletseng, karolo e putsoa le e tšoeu ea kelello e ntse e tsoela pele ho ba le liphetoho tse matla nakong ea bocha. Boitsebiso bo tsoang boithuto ba morao-rao ba MRI bo bonts'a hore bophahamo ba litaba tsa bohlooho bo na le sebopeho se senngoeng sa U, ka phapang e kholo ho feta taba e tšoeu (Giedd, 2004; Gogtay et al., 2004; Sowell et al, 2003; Sowell, Thompson, & Toga, 2004). Ka kakaretso, libaka tse ts'ehetsang mesebetsi ea mantlha, e kang tsamaiso ea makoloi le ea sensory, e holileng tsebong; libaka tse hloahloa tsa kopano e hokahanyang mesebetsi ena ea mantlha, e holileng tsebong hamorao (Gogtay et al., 2004; Sowell, Thompson, & Toga, 2004). Mohlala, lithuto tse sebelisang mehato e thehiloeng ho MRI li bonts'a hore tahlehelo ea boea ba cortical gray e etsahala pele libakeng tsa sensorimotor tsa mantlha le tsa morao-rao litabeng tsa dorsolateral pele le tsa morao-rao tsa nakoana (Gogtay et al., 2004). Sebopeho sena se lumellana le lithuto tsa batho ba morao-rao eseng tsa motho le tsa postmortem tse bonts'ang hore preortalal cortex ke e 'ngoe ea likarolo tsa ho qetela tsa boko ho hola (Bourgeois, Goldman-Rakic, & Rakic, 1994; Huttenlocher, 1979). Ha e bapisoa le taba e putsoa, ​​bophahamo ba litaba tse tšoeu bo eketseha ka tsela e batlang e tšoana, e ntse e hola ho hola ho fihlela motho e moholo.Gogtay et al., 2004). Liphetoho tsena li bonts'a tloaelo ea li-axon tse tsoelang pele ke oligodendrocyte tse ntlafatsang tlhahiso ea methapo le puisano.

Leha tlhokomelo e sa tsotelleloe ho libaka tse tlasa maemo a tlase ha ho hlahlojoa liphetoho tsa sebopeho, liphetoho tse ling tse kholo ka ho fetisisa bokong ho feta ntlafatsong li bonoa libakeng tsena, haholo-holo ho basal ganglia (Sowell et al., 1999, bona Feie. 3) mme haholoholo ho banna (Giedd et al., 1996). Liphetoho tsa nts'etsopele ea bophahamo ba sebopeho kahara basal ganglia le libaka tse tlang pele li khahla molemong oa lits'ebetso tse tsebahalang tsa nts'etsopele (mohlala, dendritic arborization, lefu la cell, pruning ea synaptic, myelination) tse etsahalang nakong ea bongoana le bohlankana. Ts'ebetso tsena li lumella tokiso e ntle le matlafatso ea khokahano lipakeng tsa libaka tsa pele le tse tlasa tlhabollo le thuto tse ka tsamaeang le taolo e kholo ea kelello. Liphetoho tsee tsa sebopeho li amana joang le phetoho ea kelello? Boithuto bo bongata bo na le ts'ebetso ea pele ea lobe sebopeho le ts'ebetso ea kelello e sebelisang mehato ea neuropsychological le temohisiso (mohlala, Sowell et al., 2003). Ka ho khetheha, mekhatlo e tlalehiloe pakeng tsa MRI e thehiloeng pele ea cortical le basal ganglia lebelo le mehato ea taolo ea kelello (ke hore, bokhoni ba ho fetisa karabelo e sa lokang molemong oa e mong kapa ho hatella tlhokomelo molemong oa tšusumetso e sa sebetseng molemong oa semelo se khothalletsang tšusumetso (Casey, Mokoetlisi et al., 1997). Liphumano tsena li fana ka maikutlo a hore liphetoho tse nang le kelello li bonahala literekeng tsa phetoho ea boko le ho hatisa bohlokoa ba nts'etsopele ea subcortical (basal ganglia) le cortical (mohlala, pele ho cortex).

Feie. 3    

Papiso ea libaka tsa boko e bonts'a liphetoho tse kholo ho feta tsa bohlankana kapa boroetsana (ho tloha Sowell et al., 1999).

DTI lithuto tsa kholo ea kholo ea motho

Boithuto bo thehiloeng ho moralo oa MRI bo hlahlojoang bo fana ka maikutlo a hore khokahano ea cortical e ntse e hlophisitsoe hantle ka ho felisoa ha li-synapses tse ngata le ho matlafatsa likhokelo tse amanang le nts'etsopele le boiphihlelo. Tsoelo-pele ea morao-rao theknoloji ea MRI, joalo ka DTI e fana ka sesebelisoa sa ho lekola karolo ea lipampitšana tsa litaba tse tšoeu ho nts'etsopele ea kelello le boits'oaro ka lintlha tse ngata haholoanyane. Se loketseng pampiri ena ke lithuto tse natefisang tse amanang le nts'etsopele ea lipampitšana tsa fiber le ntlafatso ea bokhoni ba kelello. Haholo-holo, ho bontšitsoe likamano pakeng tsa mehato e thehiloeng ho DTI ea nts'etsopele ea litaba tse tšoeu pele le taolo ea kelello baneng. Phuputsong e le 'ngoe, nts'etsopele ea matla ana e ne e hokahane hantle le lipampitšana tsa pele-parietal fiber (Nagy, Westerberg, & Klingberg, 2004) e lumellanang le lithuto tse sebetsang tsa neuroimaging tse bonts'ang ho hira libaka tse fapaneng ho bana ho amanang le batho ba baholo.

U sebelisa mokhoa o tšoanang. Liston et al. (2005) e bonts'itse hore lipampitšana tsa litaba tse tšoeu lipakeng tsa lipampitšana tsa fiber-pre-basal ganglia le -posterior fiber li ntse li tsoela pele ho hola bongoaneng ho fihlela e le motho e moholo, empa ke lipampitšana tseo feela pakeng tsa cortex ea pele le gangal tse amanang le taolo ea tšusumetso, joalo ka ha ho lekantsoe ke ts'ebetso ea ho ea / nogo mosebetsi. Lipampitšana tsa pele tsa fiber li ne li hlalosoa ke likarolo tsa lithahasello tse khethiloeng thutong ea fMRI ba sebelisa mosebetsi o tšoanang. Hohle lithuto tsa DTI ka bobeli tsa nts'etsopele, mehato ea pampitšana ea fiber e ne e hokahane le nts'etsopele, empa ho nepahala ha lipampitšana tse ling tsa fiber le ts'ebetso ea kelello li ile tsa bonts'oa ka ho khetholla pampitšana e itseng (Liston et al., 2005) kapa bokhoni ba ho lemoha (Nagy et al., 2004). Liphumano tsena li hatisa bohlokoa ba ho hlahloba eseng feela tsa tikoloho, empa liphetoho tse amanang le potoloho ha li etsa liqeto mabapi le liphetoho tse itšetlehileng ka lilemo bakeng sa nts'etsopele ea kelello.

Lithuto tsa MRI tse sebetsang tsa ntlafatso ea boitšoaro le bokong

Leha liphetoho tsa sebopeho tse lekantsoeng ke MRI le DTI li amahantsoe le liphetoho tsa boits'oaro nakong ea nts'etsopele, mokhoa o otlolohileng haholoanyane oa ho hlahloba sebopeho sa tšebetso-karolo ke ho lekola liphetoho bokong le liketsong ka nako e le ngoe, joalo ka fMRI. Bokhoni ba ho lekanya liphetoho tse sebetsang bokong bo ntseng bo eketseha le MRI bo na le monyetla o moholo bakeng sa lefapha la saense ea nts'etsopele. Moelelong oa pampiri ea hajoale, fMRI e fana ka mokhoa oa ho qobella litlhaloso tsa boits'oaro ba bohlankana. Joalokaha ho boletsoe pejana, nts'etsopele ea "cortex ea pele" ho lumeloa hore e bapala karolo ea bohlokoa ho ntlafatseng bokhoni ba tsebo e phahameng joalo ka ho etsa liqeto le taolo ea kelello (Casey, Tottenham, le Fossella 2002b; Casey, Mokoetlisi et al., 1997). Ho sebelisitsoe lipapatso tse ngata, hammoho le fMRI, ho lekola motheo oa methapo ea bokhoni bona, ho kenyelletsa mesebetsi ea Flanker, Stroop le go / nogo (Casey, Castellanos et al., 1997; Casey, Giedd, le Thomas, 2000a; Durston et al., 2003). Ka kopanelo, lithuto tsena li bonts'a hore bana ba hira libaka tse ikhethileng empa hangata li le ngata, li senya libaka tsa pele ha ba etsa mesebetsi ena ho feta batho ba baholo. Mokhoa oa ts'ebetso kahare ho likarolo tsa boko o bohareng ba tšebetso ea mesebetsi (ke hore, o tsamaellanang le ts'ebetso ea kelello) o shebane haholo le ho hlophisoa hantle ke lilemo, ha libaka tse sa tsamaellaneng le ts'ebetso ea mosebetsi li fokotseha ts'ebetsong le botsofaling. Sebopeho sena se bonoe likarolong tsohle tse peli tsa sefapano (Brown et al., 2005) le lithuto tse telele (Durston et al., 2006) le ho fetela lipapatso tse fapaneng. Leha lithuto tsa neuroimaging li ke ke tsa bonts'a ka mokhoa o hlakileng mokhoa oa liphetoho tse joalo tsa ntlafatso (mohlala, dendritic arborization, synun, pruning) liphuputso li bonts'a ntlafatso kahare, le ho nchafatsa, ho nahanisisa ho, le ho tsoa, ​​likarolo tsa ts'ebetso ea ts'ebetso ea mmele le ts'ebetso mme li fana ka tlhahiso ea hore liphetoho tsena li etsahale ka nako e telele. nako ea nako (Brown et al., 2005; Bunge, Dudukovic, Thomason, Vaidya, le Gabrieli, 2002; Casey, Mokoetlisi et al., 1997; Casey et al., 2002a; Crone, Donohue, Honomichl, Wendelken, & Bunge, 2006; Luna et al., 2001; Mose et al., 2002; Schlaggar et al., 2002; Tamm et al., 2002; Thomas et al., 2004; Turkeltaub, Gareau, Lipalesa, Zeffiro, le Edene, 2003).

Mokhoa ona o ka re tsebisa joang hore na bacha ba hlile ba haelloa ke taolo e lekaneng (ba sa susumetse) kapa ba kotsi likhethong le liketsong tsa bona? Taolo ea tšusumetso joalo ka ha e lekantsoe ke mesebetsi ea taolo ea kelello joalo ka mosebetsi oa ho ea / nogo li bonts'a mohlala oa nts'etsopele ea bongoaneng le bongoaneng joalo ka ha ho hlalositsoe kaholimo. Leha ho le joalo, lithuto tsa morao-rao tsa morao-rao li se li qalile ho hlahloba ts'ebetso e amanang le moputso e tobileng ho ho bea bacha kotsing (Bjork et al., 2004; Ernst et al., 2005; May et al., 2004). Lithuto tsena li shebile haholo ka tikoloho ea li-accumbens, karolo ea basal ganglia e amehang ho bolela moputso, ho e-na le tšobotsi ea nts'etsopele ea sebaka sena ka kopanelo le libaka tsa taolo tse holimo (preortalal cortex). Leha ho bonts'itsoe tlaleho ea haufinyane ea mosebetsi o tlang pele ho bacha lilemong tsa bocha e amanang le batho ba baholo nakong ea ts'ebetso ea ho etsa liqeto mabapi le boitšoaro bo behang kotsing.Eshel, Nelson, Blair, Pine, & Ernst, 2007).

Ka kakaretso, lithuto tse 'maloa li sekasekile hore na nts'etsopele ea meputso ea meputso libakeng tse maemong a tlasa maemo a tlase joang (mohlala, li-accumbens) e fetoha hammoho le nts'etsopele ea libaka tsa pele. Ho feta moo, hore na liphetoho tsena tsa neano li tsamaellana joang le ho batla moputso, ho susumetsa le boitšoaro bo nkang kotsi ho ntse ho sa tsejoe. Moetso oa rona oa "neurobiological" o akantsha hore ho kopanngwa ha karabelo e phahamisitsoeng ho meputso le ho se tsitse ha libaka tsa taolo ea boits'oaro li ka rata bacha ba batlang kapele, ho e-na le ho fumana melemo ea nako e telele, mohlomong ba hlalosetsa keketseho ea bona ea ho etsa liqeto tse kotsi le boits'oaro bo sa susumetseng. Ho latedisa subcortical (mohlala, bokapele) le nts'etsopele (mohlala, pele-pele) ea ho etsa liqeto ho fihlela bongoaneng, e fana ka litšitiso tse ling tsa hore na liphetoho tse tlalehiloeng bongoaneng li ikhethile nakong ena ea kholo kapa li bontša ho hola ho etsahalang ka mokhoa o tsitsitseng nakong ea kholo. mofuta o itseng oa methapo ho tloha bongoaneng ho ea ho motho e moholo.

Bopaki bo matlafatsang bo tsoang phuputsong ea morao-rao ea fMRI bo thusa ho ts'ehetsa mohlala oa rona oa neurobiological mme bo nka mokhoa oa phetoho ho utloisisa bocha ka ho hlahloba liphetoho pele le ho latela lilemo tsa bocha. Thutong ena (Galvan et al., 2006), re ile ra hlahloba likarabo tsa boits'oaro le maikutlo a ho fana ka moputso molemong oa nts'etsopele, ra tsepamisa maikutlo libakeng tsa boko tse kenyellelitsoeng ho ithuteng ho amanang le moputso le boits'oaro ho phoofolo (Hikosaka le Watanabe, 2000; Pecina, Cagniard, Berridge, Aldridge, le Zhuang, 2003; Schultz, 2006) le lithuto tsa ho nahana ka batho ba baholo (mohlala, Knutson, Adams, Fong, & Hommer, 2001; O, Doherty, Kringelbach, Rolls, Hornak, Andrews, 2001; Zald et al., 2004) le lithutong tsa bokhoba ba tahi (Hyman le Malenka, 2001; Volkow & Li, 2004). E ipapisitse le mehlala ea litoebaLaviola et al., 1999; Spear, 2000) le mosebetsi oa ho nahana ka pele (Ernst et al., 2005), re kholoa hore setho ho bana le batho ba baholo, bacha ba ka bonts'a ts'ebetso e fetelletseng ea li-accumbens, ka konsarete le ho hira batho ba maemong a holimo ba laoloang ka pele. Mosebetsi oa morao-rao o bonts'ang ts'ebelisano e liehang ea ts'ebetso lipakeng tsa libaka tsena tsa pele le tsa maoto tse hanyane ka hanyane lilemong tsa bocha tse amanang le batho ba baholo, e fana ka mokhoa oa ho hloka taolo ea libaka tse tlase (Hare et al., 2007a).

Seo re se fumaneng se ne se lumellana le mehlala ea litoeba (Laviola, Macri, Morley-Fletcher, le Adriani, 2003) le lithuto tse fetileng tsa monahano (Ernst et al., 2005) Ho fana ka tlhahiso ea ts'ebetso e ntlafalitsoeng ea ho bokella melemo ea bongoana. Ho joalo, ho amana le bana le batho ba baholo, bacha ba bontšitse karabelo e fetelletseng ea bokhoni ba ho lebella moputso. Le ha ho le joalo, bana le bacha ba bontšitse karabelo e fokolang tsebong libakeng tsa taolo ea pele ho feta batho ba baholo. Liphumano tsena li fana ka maikutlo a fapaneng a ntlafatso ea libaka bakeng sa libaka tsena tse ka eketsang katoloso ea tšebetso ea bokapele, e amanang le bana kapa batho ba baholo, e ka amanang le boits'oaro bo eketsehang le bo kotsi bo hlokometsoeng nakong ena ea ntlafatso (bona Feie. 4).

Feie. 4    

Localization ea ts'ebetso ka ho lebella sephetho sa moputso bokong ba li-nucleus accumbens (A) le orbital frontal cortex (B). Boholo ba tšebetso libakeng tsena bo hlophisitsoe e le mosebetsi oa lilemo, bakeng sa thuto ka 'ngoe e bonts'a kholo e tsoetseng pele ...

Ho bokelloa ka mokhoa o fapaneng oa libaka tsa pele le tsa subcortical ho tlalehiloe ho pholletsa le lithuto tse 'maloa tsa nts'etsopele ea fMRI (Casey et al., 2002b; Monk et al., 2003; Thomas et al., 2004). Ka tloaelo liphumano tsena li hlalositsoe ho latela libaka tse tlileng pele ho pele ho feta tikoloho ho fapana le ho se lekane pakeng tsa nts'etsopele ea sebaka sa pele le subcortical. Ho fanoe ka bopaki ba libaka tsa pele ho tataisa liketso tse nepahetseng maemong a fapaneng (Miller & Cohen, 2001) ts'ebetso ea pele ho nako e ka sitisa khakanyo e nepahetseng ea sephetho sa kamoso le ho lekola likhetho tse kotsi, 'me ka hona, e ka ba le tšusumetso ho boleng ba moputso ho feta ho bokelloa. Mokhoa ona o lumellana le lipatlisiso tse fetileng tse bonts'ang maemo a tlase a amanang le ts'ebetso ea maikutlo ha liqeto li khethoa ke melemo ea nako e telele (McClure, Laibson, Loewenstein, & Cohen, 2004). Ho feta moo, ts'ebetso ea bokhabane e bontšitsoe le fMRI ho lumellana hantle le boits'oaro bo latelang ba ho ipeha kotsing (Kuhnen le Knutson, 2005). Nakong ea bocha, ho amana le bongoana kapa ho ba motho e moholo, cortex ea bongoana e tlang pele ho nako e kanna ea se ke ea fana ka taolo e lekaneng ea holimo ea libaka tse sebetsanang le moputso (mohlala, li-accumbens), tse hlahisang tšusumetso e nyane ea litsamaiso tsa pele (li-orbitofrontal cortex) tse amanang le maqhubu a moputso. boleng.

Hobaneng ha boko bo hlophisitsoe hore bo hōle ka tsela ee?

Nako ea bocha ke nako ea phetoho lipakeng tsa bongoana le ho ba motho e moholo e atisang ho ba teng nakong ea boroetsana. Ho kena bohlankaneng ho tšoaea qaleho ea nako ea thobalano (Graber & Brooks-Gunn, 1998) hape e ka hlalosoa ke matšoao a tlhaho. Nako ea bocha e ka hlalosoa e le phetoho e tsoelang pele ea ho ba motho e moholo ka thuto ea nakoana ea ho etsa joalo (Spear, 2000). Ha re bua ka mokhoa o iphetolang, nako ea bocha ke nako eo ka eona tsebo ea boipuso e fumanoang ho eketsa katleho holima tšireletso ea lelapa, leha ho le joalo eketsa menyetla ea maemo a kotsi (mohlala, ho lemala, khatello ea maikutlo, ho tšoenyeha, tšebeliso ea lithethefatsi le bokhoba ba tahi (Kelley, Schochet, & Landry, 2004). Boitšoaro bo batlang boipuso bo atile mefuta eohle ea mefuta, joalo ka keketseho ea likamano tse tataisoang ke lithaka le ho matlafatsa boits'oaro bo batlang bo sa tloaelehang le bo behang motho kotsing. Lintlha tsa kelello le maikutlo li ama tekano ea bocha bakeng sa boits'oaro bo kotsi. Leha ho le joalo, boits'oaro bo kotsi ke sehlahisoa sa ho se lekane pakeng tsa tlhaho le lipono tsa maikutlo ka kopanelo le "bokhoni ba ho itaola" (Steinberg, 2004). Lintlha tsa rona tsa neurobiological li bontša hore sena se etsahala ka nts'etsopele ea lits'ebetso tsena tse peli (maoto le taolo).

Ho nahanoa ho ka fana ka maikutlo a hore mokhoa ona oa ntšetso-pele ke karolo ea ho iphetola ha lintho. O hloka ho etsa boits'oaro bo kotsing e kholo ea ho siea ba lelapa le motsana oa hau ho fumana molekane le ho nka likotsi ka nako e ts'oanang ha li-hormone li khanna bacha ho batla balekane ba thobalano. Sechabeng sa kajeno ha boroetsana bo ka nyoloha ka nako e sa lekanyetsoang, le bana ba lulang le batsoali mme ba na le ts'epo ea chelete mme ba khetha balekane hamorao bophelong, khopolo ena e ka nkoa e sa nepahala.

Ho na le bopaki ka mefuta eohle ea ho batla bocha le ho ipeha kotsing nakong ea lilemo tsa bocha. Ho batla lithaka tsa lilemong tse ts'oanang le ho loana le batsoali, tseo kaofela li thusang ho etsa hore mocha a hole le sebaka sa lapeng bakeng sa ho ikatisa ho bonoa mefuta e meng e kenyelletsang litoeba, li-primates tse seng tsa motho le linonyana tse ling.Spear, 2000). E amanang le batho ba baholo, likhoto tsa periadolescent li bonts'a mekhoa e mengata ea bo-ralitaba e batlang e le mokhoa oa ho ithuta ka mokhoa oa mahala oa papali oa palesa (Laviola et al., 1999). Bopaki ba neurochemical bo bontša hore tekano bokong ba bocha pakeng tsa litsamaiso tsa dopamine ea cortical le subcortical dopamine, e qala ho fetohela maemong a dopamine a maholoanyane a dopamine nakong ea bocha.Spear, 2000). Tlhabollo e tšoanang ea nako e telele ea dopaminergic ka ho ba motho e moholo e bontšitsoe ho cortex e seng ea botho ea pele ho motho hape (Rosenberg le Lewis, 1995). Kahoo ho bonahala kotsi e ka holimo-limo e nka sebaka sa mefuta eohle ebile e na le sepheo sa bohlokoa.

Melemo ea tlhaho ea tlhaho, nts'etsopele le kotsi

Phapang e fapaneng ea taolo ea tšusumetso le ho nka likotsi e lemohile ho psychology ka nako e telele (Benthin, Slovic, & Severson, 1993). Mohlomong e 'ngoe ea mehlala ea khale ea liphapang tse tlalehiloeng ka bokhoni bona lithutong tsa kelello, tsa kelello le tsa tsoelopele ke ho lieha ho khotsofatsa (Mischel, Shoda, & Rodriguez, 1989). Ho lieha ho khotsofala ho hlahlojoa hangata ho 3- ho bana ba 4 ba lilemo li. Mosha o botsoa hore na ba ka rata moputso o nyane (cookie e le 'ngoe) kapa moputso o moholo (li-cookie tse peli). Joale ngoana o bolelloa hore ea lekang o tla tsoa ka phapusing e le hore a itokisetse liketsahalo tse tlang mme o hlalosetsa ngoana hore haeba a lula setulong sa hae mme a sa jele cookie, o tla fumana moputso o moholo. Haeba ngoana a sa emele kapa a sa khone ho emela, o lokela ho letsa tšepe ho bitsa sebapali ebe o amohela moputso o nyane. Ha ho hlakile hore ngoana o utloisisa mosebetsi, o lutse tafoleng ka meputso e 'meli le mohala. Litšitiso ka phapusing lia fokotsoa, ​​ho se na lintho tsa ho bapala, libuka kapa litšoantšo. Teko e khutla kamora metsotso ea 15 kapa kamora hore ngoana a hlake tšepe, a je meputso, kapa a bontše matšoao a khatello. Mischel e bonts'itse hore bana ka tloaelo ba phela ka tsela e 'ngoe ea tse peli: (1) ekaba ba letsa tšepe hang-hang hore ba be le cookie, ho bolelang hore ba fumana e le' ngoe feela; kapa (2) ba emela le ho ntlafatsa menyetla ea bona, 'me ba amohela li-cookie ka bobeli. Tlhatlhobo ena e fana ka maikutlo a hore batho ba bang ba betere ho feta ba bang matleng a bona a ho laola litšusumetso ha ba shebane le tšusumetso e matla haholo mme khethollo ena e ka fumanoa bongoaneng.Mischel et al., 1989) 'me li bonahala li lula li le lilemong tsa bocha le ho ba motho e moholo.Eigsti et al., 2006).

Ke eng e ka hlalosang karohano ea motho ka mong ho etseng liqeto le boits'oaro bo nepahetseng? Ba bang ba theoriste ba supile hore dopaminergic mesolimbic circry, e kentsoeng ts'ebetsong ea moputso, e ka beha boitšoaro bo kotsi. Phapang pakeng tsa motho ka mong ho potoloho ena, joalo ka mefuta e fapaneng ea liphatsa tsa lefutso tse amanang le dopamine, e bakang dopamine e nyane haholo kapa e ngata haholo libakeng tse tlasa subportical, e kanna ea amana le matla a ho etsa boits'oaro bo kotsi (O'Doherty, 2004). Li-nucleus accumbens li bontšitsoe hore li eketsa ts'ebetso hanghang pele ho etsoa likhetho tse kotsi ho paradigms ea kotsi ea chelete (Kuhnen le Knutson, 2005; Matthews le al., 2004; Montague le Berns, 2002) 'me joalo ka ha ho hlalositsoe pejana, bacha ba bonts'a mosebetsi o mongata oa ho bokella chelete e ngata ho hlahisa litholoana tse khotsofatsang tse amanang le bana kapa batho ba baholo.Ernst et al., 2005; Galvan et al., 2006). Ka kopanelo, lintlha tsena li fana ka maikutlo a hore bacha ba ka ba monyetleng oa ho khetha likotsi tse le sehlopha e le sehlopha (Serapa sa jarete le Steinberg, 2005), empa bacha ba bang ba tla potlakela ho feta ba bang ho etsa lintho tse ka behang bophelo ba bona kotsing, ka ho ba beha kotsing e kholo ea litlamorao tse mpe. Ka hona ho bohlokoa ho nahana ka phapano ea motho ka mong ha u hlahloba likamano tse rarahaneng tsa boits'oaro ba boko bo amanang le ho nka kotsi le ts'ebetso ea meputso maemong a ntlafatso.

Ho hlahloba ho se tšoane ka boits'oaro bo nkang kotsi, Galvan et al. (2007) Haufinyane tjena o ile a hlahloba kamano lipakeng tsa ketsahalo e amanang le moputso o amanang le neural ka tebello ea moputso o moholo oa chelete o nang le litšobotsi tsa botho ba ho ipeha kotsing le ho kenella kapele bohlankaneng. Melemo ea maiketsetso ea ts'ebetso ea matla a boselamose le litekanyetso tse sa tsejoeng tsa ho itlaleha tsa boitšoaro bo kotsi, pono le kotsi li fumanoe ho batho ba pakeng tsa 7 le 29 lilemo. Ho ne ho na le kamano e ntle lipakeng tsa ketsahalo ea accumbens le monyetla oa ho etsa boitšoaro bo kotsi holima kholo. Ketsahalo ena e ne e fapana joaloka litekanyetso tsa batho ka bomong tsa litlamorao tse ntle kapa tse mpe tsa boitšoaro bo joalo. Batho bao ba boneng boits'oaro bo kotsi e le bo lebisang liphellong tse mpe, ba ile ba kenya chelete eo hore e fuoe. Mokhatlo ona o ne o tsamaisoa haholo ke bana, mme ba baholo ba hakanya litholoana tsa boitšoaro bo joalo kamoo ho ka khonehang. Likeletso tsa impulsivity li ne li sa amane le ts'ebetso ea accumbens, empa ho fapana le lilemo. Liphumano tsena li fana ka maikutlo a hore nakong ea bocha, batho ba bang ba ka ba le tšekamelo ea ho etsa boits'oaro bo kotsi ka lebaka la liphetoho tsa nts'etsopele ea concert le ho feto-fetoha ha maemo ha motho ea ikemiselitseng ho nka boits'oaro bo kotsi, ho fapana le liphetoho tse bonolo tsa ho potlaka (bona. Feie. 5).

Feie. 5    

Bacha ba bonts'a ts'ebetso e ntlafalitsoeng ea li-accumbens tse amanang le bana le batho ba baholo (A). Ts'ebetso ea Accumbens e amahanngoa hantle le litekanyetso tsa ho itšupa menyetla ea ho etsa boitšoaro bo kotsi (B) mme e amana hantle le boitekolo ba ...

Boitšoaro ba bongoaneng bo 'nile ba tšoauoa khafetsa e le bo sa susumetseng ebile bo le kotsi (Steinberg, 2004, 2007), leha ho le joalo tlhahlobo ena ea lingoliloeng tse inahaneloang e fana ka likarolo tse fapaneng tsa "neurobiological substrates" le li-trajectories tse fapaneng tsa nts'etsopele bakeng sa boits'oaro bona. Ka ho khetheha, ho susumetsoa ho amana le nts'etsopele ea bokhachane ba pelehi mme butle-butle e fokotseha ho tloha bongoaneng ho ea ho motho e moholo (Casey, Galvan et al., 2005). Kamano e mpe lipakeng tsa litekanyetso tsa impulsivity le lilemo ho thuto ke Galvan et al. (2007) e ts'ehetsa taba ena hape. Ho fapana le hoo, ho nka likotsi ho amana le keketseho ea ketsahalo ea liqoqiso (Kuhnen le Knutson, 2005; Matthews le al., 2004; Montague le Berns, 2002), seo se eketsoa ho feta lilemong tsa bocha, se amanang le bana le batho ba baholo (Ernst et al., 2005; Galvan et al., 2006). Kahoo likhetho le boits'oaro ba bongoaneng li ke ke tsa hlalosoa ka ho qobelloa kapa ho nts'etsopele hoa "cortex ea pele" feela, joalo ka ha bana ba tla boleloa esale pele hore ke bona ba ka behang kotsing e kholo. Se fumaneng se fana ka motheo oa neural bakeng sa hore na hobaneng bacha ba bang ba kotsing e kholo ho feta ba bang, empa e tsoela pele e fana ka motheo oa hore na boitšoaro ba bongoaneng bo fapane joang le ba bana le batho ba baholo ba ipehang kotsing.

Ka kakaretso, lintlha tsena li fana ka maikutlo a hore le ha bacha e le sehlopha ba nkuoa e le ba nkang likotsi (Serapa sa jarete le Steinberg, 2005), bacha ba bang ba tla ba le tšekamelo e fetang ea ba bang ea ho etsa boits'oaro bo kotsi, ba ba behang kotsing e kholo ea litlamorao tse mpe. Liphumano tsena li totobatsa bohlokoa ba ho nahana ka phapang ea motho ka mong ha ho hlahlojoa likamano tse rarahaneng tsa boko le boits'oaro tse amanang le ho nka kotsi le ts'ebetso ea meputso ho palo ea batho ba hola. Ho feta moo, liphapang tsena tsa motho ka mong le tsa tsoelo-pele li ka thusa ho hlalosa ho ba kotsing ea ho itšireletsa ho batho ba bang ka ho beha kotsing ea ts'ebeliso ea lithethefatsi, 'me qetellong, ho lemalla.

Nahanisisa

Lithuto tsa ho nahana ka batho li bonts'a liphetoho tsa sebopeho le tšebetso libakeng tse ka pele.Giedd et al., 1996, 1999; Jernigan et al., 1991; Sowell et al., 1999; bakeng sa tlhahlobo, Casey, Galvan et al., 2005) tse bonahalang eka li tsamaisana le taolo ea taolo ea kelello le boitaolo (Casey, Mokoetlisi et al., 1997; Luna & Sweeney, 2004; Luna et al., 2001; Rubia et al., 2000; Steinberg, 2004;; bona hape Steinberg, 2008, makasine ona). Liphetoho tsena li bonahala li bonts'a phetoho ea ts'ebetsong ea libaka tsa pele ho tloha ho ferekanya ho isa ho tsepamisitsoeng pele ho feta ka nako (Brown et al., 2005; Bunge et al., 2002; Casey, Mokoetlisi et al., 1997; Durston et al., 2006; Mose et al., 2002) le ho hloa libaka tse phahameng tsa maemo a tlase lilemong tsa bocha.Casey et al., 2002a; Durston et al., 2006; Luna et al., 2001). Leha lithuto tsa neuroimaging li ke ke tsa bonts'a ka mokhoa o hlakileng mokhoa oa liphetoho tse joalo tsa nts'etsopele, liphetoho tsena molemong oa sebopeho le sebopeho li ka bonts'a ntlafatso kahare, le ntlafatso ea, likhakanyo tsa ho tsoa le ho tsoa libakeng tsena tsa boko nakong ea kholo ea kholo ea ho hlophisoa ha sistimi le nts'etsopele.

Ha ho kopantsoe hammoho, liphumano tse hlophisitsoeng mona li bonts'a hore boits'oaro bo eketsehileng ba ho beha bonyenyaneng bo amana le likarolo tse fapaneng tsa nts'etsopele ea mehopolo e tlase ea boithabiso le libaka tse laoloang ke maemo a phahameng ho fetisisa. Liphetoho tsena tsa nts'etsopele li ka eketsoa ke ho se tšoane hoa tšebetso ea lits'ebetso tsa meputso. Leha bocha bo khethollotsoe e le nako e tšoauoa ka boits'oaro ba ho batla moputso le boits'oaro bo behang kotsing (Serapa sa jarete le Steinberg, 2005; Spear, 2000) phapang e fapaneng le e 'ngoe ea likarabo tsa neural ho fana ka moputso, ho lebella bacha ba bang ho beha likotsi tse ngata ho feta tse ling, ho ba beha kotsing e kholo ea litlamorao tse mpe. Liphumano tsena li fana ka motheo oa bohlokoa ka ho kopanya liphumano tse fapaneng tse amanang le boits'ebetso bo beang bonyenyaneng lilemong tsa bocha le ho utloisisa liphapang tsa batho ka bomong le matšoao a nts'etsopele bakeng sa maano a ho etsa boitšoaro bo bobe.

lumela hore baa fokola

Mosebetsi ona o ile oa tšehetsoa ka lichelete ke lithuso tse tsoang ho National Institute of Drug Abuse R01 DA18879 le National Institute of Mental Health 1P50 MH62196.

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