Litlhaselo tse tsoelang pele nakong ea bocha ho banna le basali: kutloisiso ea mefuta-futa ea liphetoho tsa boko bo ka sehloohong (2011)

E hatisitsoe ka mokhoa o qetelle o hlophisitsoeng e le:
PMCID: PMC3134153
NIHMSID: NIHMS296109
Moetso oa khatiso oa ho qetela oa mohatisi o fumaneha ho Neurosci Biobehav Rev
Sheba lihlooho tse ling ho PMC hore Qotsa sehlooho se hatisitsoeng.

inahaneloang

Nako ea bocha ke nako ea phetoho lipakeng tsa bongoana le ho ba motho e moholo e akaretsang liphetoho tse kholo tsamaisong ea boko tse tšoanang le tse ling, empa eseng kaofela, liphetoho tsa boits'oaro. Liphetoho tse bang teng kelellong le tšebetsong ea moputso li latela sebopeho se phetleng sa U ho latela ho qala le ho tšoarela, ka tlhoro e hlahang nakong ea bocha. Leha ho le joalo, ts'ebetso ea kelello e latela kemiso e kholo ea kholo. Tlhahlobo ena e tla shebisisa liphetoho tse sebopeho sa likarolo tsa bohlokoa 'me e tla totobatsa likamano lipakeng tsa liphetoho tsa boko le boits'oaro,' me bopaki bo nka nako ho tloha lipapaling tsa tšebetso ea matla a khoheli (fMRI) bathong ho ea ho lithuto tsa limolek'hule le lintlha tse tšoaeang liphoofolo. Liphetoho tsa lilemong tsa bocha tsa "neuronal substrates" li tla sebelisoa ho utloisisa hore na boits'oaro bo tloaelehileng le ba sebopeho ba hlaha joang lilemong tsa bocha. Re latela lithuto tsa tliliniki le tsa pele ho fana ka moralo oa neural oa ho hlalosa bocha le karolo ea ona phetohong ea ho ba motho e moholo.

Keywords: Bocha, ntho e putsoa, ​​ho faola, phapang ea thobalano, taba e tšoeu

Selelekela

Nako ea bocha ke nako e ikhethang kholisong ea 'mele oa mammary. Ho utloisisa lilemo tsa bocha ho hlalositsoe litlhahlobo tse ngata maemong a boitšoaro (McCutcheon le Marinelli, 2009; Spear, 2000; Steinberg, 2010; Laviola et al., 1999; Laviola et al., 2003) le boemo ba sistimi (Ernst le Fudge, 2009), empa ho buisanoa ka eona feela ho isa tekanyong e lekantsoeng ea liphetoho tsa neuronal (Andersen, 2003; McCutcheon le Marinelli, 2009; O'Donnell, 2010; Spear, 2000). Re tla shebisisa neuroanatomy, kgokahano e sebetsang, liphatsa tsa lefutso, le ho supa liphetoho tse etsahalang nakong ea bocha. Kamora nako, moralo o kahare ho mekhoa ea neural e tla akaretsa hore na liphetoho tsa bocha ho matšoao ana li susumetsa boitšoaro joang.

1. Kakaretso

1.1 Ho hlalosa bocha

Nako ea bocha e ka hlalosoa e le nako e pakeng tsa lilemo tsa 10-19 ho batho (WHO, 2010s), lipakeng tsa lilemo tse peli le tse 'ne ho li-primates (Schwandt et al., 2007), le pakeng tsa matsatsi a 35-60 a lilemo tsa litoeba (Andersen et al., 2000; McCutcheon le Marinelli, 2009). Spear (2000) e qala moqoqo oa hae oa nako ena ka nako ea bocha e hlalosoang e le nako ea phetoho ea boits'oaro. Liphetoho tse joalo tsa boits'oaro li lula li bonoa khafetsa ho mefuta e fapaneng ea liphoofolo tsa mammolimo ka ho eketseha ha maikutlo ho lithaka le litloaelo tsa sechaba (Blakemore, koranta ena; Forbes le Dahl, 2005; Steinberg, 2010; Panksepp, 1981), ho ipeha kotsing (Laviola et al., 2003) le taolo e hlakileng ea taolo (Casey et al., 2008). Litlhaloso tsa bohlankana li ka boela tsa theoa ka lebaka la liphetoho tsa gonadal kaha li amana le kholo ea thobalano (Sisk le Foster, 2004). Lipheho tse boletsoeng mona ha se mokhoa o fetelletseng ebile ha oa lokela ho sebelisoa ka mokhoa o makatsang, ho fapana le moo e le ntlha ea tšupiso.

Mohato o mocha oa nts'etsopele, ho ba motho e moholo ea hlahang, o hlaha lipakeng tsa lilemo tsa 18-29 ho motho (Arnett, 2000). Boholo bo hlalositsoeng ka setso, bo hlahang bathong bo hlalosa maikutlo a hore le ha liphetoho tse ngata tsa methapo ea kutlo tse amanang le bocha li felile, 'mele ha o e-so hōle moeeng joalokaha ho pakoa ke ho lieha ho fumana mosebetsi kapa lenyalo. Ho latela nalane, G. Stanley Hall (1904) o hlalositse nako e ncha ea ho hola e hlalosang bocha ho latela maemo a moruo le moruo a ileng a lebisa tlhokomelong e eketsehileng ea mohato o ikhethang. Ka lebaka leo, re supile liphetoho tse ikhethileng le tsa bohlokoa tsa methapo ea kutlo tse khethollang bocha. Le ha tlhahlobo ena e shebane haholo le li-indices tsa methapo ea bocha, ho bohlokoa ho hlokomela hore mefuteng ea litoeba ho na le nako e ka nkang batho ba baholo ba hlahang (tlhaiso-leseling e fokolang e fumaneha ho litloholo tseo eseng batho). Joalokaha ho hlalositsoe ka tlase, likhoto li bonts'a liphetoho tse tšoaeang lipakeng tsa matsatsi a 40-60, empa nako e lipakeng tsa matsatsi a 60-100 e amahanngoa le phetoho e liehang, e tsitsitseng eo butle-butle e tsitsang. Na ena ekaba nako e ncha ea "motho e moholo ea hlahang" e lokelang ho tsotelloa ke lipatlisiso, ho fapana le ketsahalo ea bophatlalatsi ho hlalosa phetoho e ncha ea setso lichabeng tse tsoetseng pele? Bohlokoa ba mekhahlelo e hlalositsoeng ke ho fihlela tumellano ea boemo ba kholo ba mmele bo hlalosoang ho nolofalletsa mefuta ea mefuta le lipapiso tsa thobalano.

1.2 Hobaneng nako e fetoha joalo?

Ho tloha ponong ea ho iphetola ha lintho, boitšoaro bo bōpiloe ke khetho ea tlhaho ho itokisetsa motho hore a atlehe lefatšeng la bophelo bo botle le 'meleng e le motho e moholo, ho kenyelletsa ho fumana molekane le katleho le katleho. Mokhoa ona o fihla sehlohlolong lilemong tsa bocha. Ka boitsoaro, liphoofolo tse anyesang tse tlohang litlamong ho isa bathong kaofela li ba le nako e kholo ea phetisetso eo ho tsamaea ho kena bonneng kapa hanyane le ts'usumetso e fokotsehileng ea motsoali e tsamaeang le ts'usumetso e eketsehang ea lithaka, tlholisano ea thobalano le mathata a macha a ho etsa liqeto (a lekantsoe ke Spear, 2000). Neuroplasticity e lumella karabelo e nepahetseng maemong a hlahang mme sena se bonahala ho nts'etsopele ea moputso le litsamaiso tse amanang le tsona (Galvan, 2010). Leha ho le joalo, lits'ebetso tse ling tsa nts'etsopele li bonts'a keketseho e tsitsitseng ea taolo ea kelello nakong ea bocha e tsamaisang ho etsa liqeto (Geier le Luna, 2009; Somerville le Casey, 2010). Ka kopanelo, yin ena le tsoelo-pele e tloaelehileng ea tsoelo-pele, moo boholo ba bacha ba loanang le phetoho ea ho ikhetholla ho lithaka le ho batsoali mme ba hlaha e le batho ba baholo ba ikemetseng, ba itaolang ha lits'ebetso tsena li fihlella tekano. Ha liphetoho tsena li hola ka tloaelo, ho etsoa liphetoho tse fapaneng le tse ikhethileng tikolohong le sechabeng. Leha ho le joalo, liphoso tsa ts'ebetso ena li baka boitšoaro bo bobe. Ho hlaha ha psychopathology ho ka hlahisoa ka tsela e 'ngoe e fapaneng le ho kheloha tloaelo e tloaelehileng ea ho feto-fetoha ha maemo le maemo, e hlahisang mathata a bophelo bo bolelele ka ts'ebetso e amanang le moputso le maikutlo. Ntle le bokhopo bo tsamaisoang ke liphatsa tsa lefutso, liphoso tsa ho hlahisa ka bongata le ho faola li-neuron kapa li-receptor, ntlafatso e fosahetseng ea ts'ebetso ea fiber kapa ho se rohakoe ha nyefolo ea bophelo ba pele kaofela e kanna ea kenya letsoho. Tlhahlobo ena e tla shebisisa lits'ebetso tsena tsa nts'etsopele bokong ba mammary, ka ho totobatsa kakaretso ka mokhoa o tloaelehileng ho fapana le aypypical (mohlala, Andersen le Teicher, 2008; 2009; Marco et al., 2011).

1.3 mofuta oa phetoho

Mokhoa oa neural system o fana ka leseli ho rarahaneng ea tlhaho ea kholo ea bongoana. Joalokaha ho hlalositsoe ke Paus (Paus et al., 2008), lits'oants'o tsa likarolo tse fapaneng tsa ts'ebetso ea ts'ebetso ea boko li bontša ka ho hlaka kamoo ho fapana ha tikoloho le tšebetso li kenyang letsoho bokong ba bongoana. Thutong ena, re hlahloba se tsebahalang ka liphetoho mokhoeng oa tsoelo-pele ka ho shebisisa mekhoa ea bongoana joalo ka ha ho hlalositsoe mefuta ea liphoofolo tse anyesang le lipakeng tsa bong. Moralo oa rona o ipapisitse le mofuta oa mofuta oa boraro, o hlalositsoeng ke Ernst le basebetsi mmoho (Ernst le Fudge, 2009; Ernst le Korelitz, 2009). Mefuta ea boraro ea methapo e fetoha ka mekhoa e meraro kapa li-node tse tharo, e leng sistimi e sebetsang, tsamaiso ea moputso, le thibelo ea kelello / karabo. Li-node tsena tse tharo tse fapaneng li sebetsa 'moho ho hlahisa boits'oaro bo tšoaeang bofetoheli ba bocha. Node e 'ngoe le e' ngoe e na le "trajectory" ea eona ea nts'etsopele, e bōpang tsamaiso ea bocha maemong a ho fetoha. Liphetho tsa ho qetela tsa boits'oaro li kanna tsa itšetleha ka moelelo o matla oa sethala se fanoeng kapa li ka bakoa ke node e fokolang e sa atleheng ho etsa mesebetsi ea taolo. Mohlala oa boraro bo felletseng ka tsela ea ona e nolofalitsoeng o fana ka ho hlalosa ho ikamahanya le maemo ha bohlankana ka bokhutšoanyane ba maikutlo, liphetoho mokhoeng oa meputso, le phetoho e tšoailoeng taolong ea cortical le nts'etsopele ea kelello. Mona, re tla sebelisa sebopeho sena ho hlalosa liphetoho tse hlakileng tlhahisong ea bacha lilemong tsa mefuta eohle le thobalano ka ho shebana le likarolo tsa boko le maoto.

2. Ho etsoa hoa mokokotlo: Liphetoho tsa Neuroanatomical

Boemong ba neuronal, ts'ebetso ea ho holisa bongoana ke e 'ngoe ea ntlafatso ea synaptic. Li-Neurons qalong li raloa ka mokhoa o kantle ho kenyelletso ea ho boloka sebaka ka cortex (Rakic ​​et al., 1986). Li-Neurons tse ileng tsa tsoaloa pele li hlokomela likarolo tse tebileng tsa cortex, ha ho bolokiloe likarolo tse ling tse sa bonahaleng tsa cortex li etsahala hamorao kholong. Ho lebisoa ho Neuronal ho tataisoa ke lisele ka bobeli tsa glia (Rakic ​​et al., 1986; Vernadakis, 1975) le li-gradients tsa lik'hemik'hale tse ikemiselitsoeng ke polelo ea neurotransmitter (Landis le Keefe, 1983; Purves le Lichtman, 1980). Polelo ea Neurotransmitter e ka ba ea ka ho sa feleng, ea baka ho kenella sebakeng se fuoeng, kapa ectopic, le li-transmit li hlahisoa hanyane ka hanyane bakeng sa nts'etsopele ea tataiso. Manonyeletso a thehoa ha li-neuron li fihla libakeng tseo li hlaseloang ho tsona. Ho rarahana ha likarolo tsa bokhachane le pelehi tsa ts'ebetso ena li hlahlojoa ka botebo bo bong hape (Levitt, 2003; Tau le Peterson, 2010), 'me e ke ke ea buisanoa ka lintlha tse joalo mona. Ha nako ea bocha e ntse e atamela, li-synapses lia hlahisoa ebe li lahleha ka morao ho moo, li bitsoa ho faola. Ho faola ke ts'ebetso e sa ts'oaneng le apoptosis le tahlehelo ea lisele, kaha ho roka ke ntlafatso ea makala a dendritic le khokahano ea synaptic le apoptosis e etselitsoe lefu la sele. Ho faola hoa li-synapses ho hlahella haholo bokong ba bongoaneng ho mefuta eohle mme ho ka hlalosoa ka tlhahlobo ea kamora lefuAndersen le Teicher, 2004; Huttenlocher, 1979; Lewis, 1997) kapa e theotsoe ho tsoa ho MRI, moo liphetoho tsa tikoloho tse putsoa le tse tšoeu li hlahang lilemong tsa bocha le hanyane ka hanyane ha batho ba ntse ba atamela lilemong tsa bona tsa boraro tsa bophelo (Giedd et al., 1999a; Huttenlocher, 1979; Sowell et al., 2004). Le ha ho reoa ha synaptic ka karolo ka 'ngoe ha ho lumeloe ho ama boleng ba tlhahlobo ea molumo (Rakic ​​et al., 1986; buiswang ke Giedd et al., 2008), liphetoho molemong oa taba e putsoa le e tšoeu li bonts'a phetoho ea likarolo tsa synaptic holima kholo.

2.1. Litšobotsi tsa Tlhahiso e Ntle le ea Petso

2.1.1. Synaptogenesis le Pruning

Ts'ebetso ea synaptogenesis le ho faola e bolokiloe haholo ho mefuta ea liphoofolo tse anyesang. Lithuto tsa motho tsa pele ho litopo tsa motho ka Huttenlocher (Huttenlocher, 1979; Huttenlocher le de Courten, 1987) le Benes (Benes et al., 1987) e bile ba pele ba bonts'ang liphetoho tse kholo litabeng tsa bohlooho le bosoeu nakong ea bongoana. Ka ho khetheha, ho prunun ka hare ho mela ea 3 ea cortex ea motho ea morao-rao ho bohlokoa haholo 'me 40% ea li-synapses e lahlehile pakeng tsa 7 le 15 ea lilemo. Mohlala, letšoao la synaptic la synaptophysin ho batho le nyoloha butle pakeng tsa lilemo tsa ho tsoaloa le 5 ea lilemo, le fihla sethaleng ka lilemo tsa 10, 'me le oela maemong a batho ba baholo ka lilemo tsa 16 tsa dorsolateral prefrontal cortex (PFC) (Glantz et al., 2007). Tlhahlobo e qaqileng ea synaptogeneis ho rhesus monkey motor cortex e senola mokhoa o ts'oanang molemong oa hore tlhahiso ea synaptic e tsoella ka morao ho nako le ho fihlella maemo a arohanang a habeli ho feta ho a batho ba baholo. Tekanyo ea li-synaptogeneis e ea fokola ha litšoene li fihlela kholo ea thobalano (lilemo tsa 3), ebe li fokotseha ka sekhahla maemong a batho ba baholo (Zecevic et al., 1989). Ka mokhoa o ts'oanang, litekanyetso tsa methapo ea kutlo ea "rataptic" li nyoloha pakeng tsa matsatsi a 25 le 40, 'me li lula li tsitsitse kamora moo (Andersen le Teicher, 2004). Leha ho le joalo, ha se liphetoho tsohle tse amanang le lilemo tse lekanang le bophahamo ba molumo li bakoang ke ho roka ha synaptic (mohlala, dendritic retraction). Mekhoa e hlakileng ea ho bala lisele ho likhoto e senola tahlehelo e amanang le lilemo tsa li-neuron kahare ea "cortex" ea mantlha likarolong tsohle (ntle le IV) likhethong ka mor'a bocha (Yates le Juraska, 2008). Phapang ea tahlehelo ea sele, joalo ka letsoalo la synaptic, le eona e bonoa. Ha cortex ea pono e bonts'a tahlehelo ea 18-20% liseleng, tahlehelo ea sele e nyane ea 5% e bonoa ka har'a ventromedial, empa eseng dorsal lateral, PFC in rats (Markham et al., 2007). Le ha tlhahiso e kholo le ho fahla e ntse e fapana lipakeng tsa libaka le lipakeng (lipakeng tsa libaka tse fapaneng), ts'ebetso e hlokomeleha mefuta eohle e fapaneng ka mehla.

Ho faola ho etsahala hangata libakeng tsa asymmetric synapses tse lutseng masapong a dendritic, joalo ka ha ho bontšitsoe ho motor cortex (Zecevic et al., 1989), molek'hule oa "gippus" ea hippocampal dentate le PFC ea dorsolateral (Eckenhoff le Rakic, 1991; Molisa, 1990). Li-synapses tsa asymmetric li hlasimolohile haholo ka tlhaho, athe li-symmetric synapses li sitisa haholo. Matšoao a methapo ea methapo ea kutlo ea "GABA" ("inhibitory transmitter") e lula e tsitsitse ho fihlela lilemo tsohle.Brenhouse et al., 2008; Vincent et al., 1995), e ts'oanang le palo e tsitsitseng ea baahi ba li-symmetric synapses tse shafts tsa dendritic (Zecevic et al., 1989). Mokhoa oa motheo oa ho faola limela ha o utloisisehe ka botlalo. Leha ho le joalo, litlhahlobo tsa morao-rao li bontšitse ka mokhoa o itseng taolo ea liphatsa tsa lefutso ea li-synapses tsa thabo. Phokotso ea bongoana ho NRG1, mofuta o amehang ho letseng la neuregulin, o ka bapala karolo ea boitšisinyo / khokahano ea inhibitory le khetho ea synaptic (Harris et al., 2009). Li-complexin, e leng liprotheine tsa presynaptic tse tataisang tokollo ea li-neurotransmitter mme li amanang le SNARE tata, le tsona li fetoha le lilemo. Complexin 2 (CX2), letšoao la li-synapses tse thabisang, e bonts'a mokhoa oa nts'etsopele oa nts'etsopele le plata ka lilemo tsa 10 ho motho. Ho fapana le moo, bothata ba "complexin 1 (CX1)" bo amanang le "inhibitory synapses" bo hola butle-butle ka ho ba motho e moholo lilemong ho batho ba boemong ba dorsolateral PFC (Salimi et al., 2008).

Le ha methapo ea kutlo ea glutamatergic e fetoha nakong ea bocha, GABA e bonts'a liphetoho tse tebileng tse amanang le lilemo tse buang ka tsona. Liphetoho tsena tsa GABA li sebetsa ka tsela ea tlhaho, athe liphetoho tsa glutamatergic li hlophisitsoe. Pele, GABA e na le liketso tse nyarosang mathoasong a pelehi ea pelehi. GABA e fumana tšusumetso ea eona ea thibelo ka nts'etsopele ea chloride Channel e fetohang bekeng ea bobeli ea bophelo ka hara rat; GABA e boloka ketso ena ea thibelo ka ho ba motho e moholo (Ben-Ari, 2002). Phetoho ena e thabisang-inhibitory e hlahisoa ke li-oscillation tse kholo maemong a calcium nakong ea kholo, e thusang nts'etsopele ea synaptic (Ben-Ari, 2002). Neonatal blockade ea mochine o ikarabellang bakeng sa ts'ebetso e phahameng ea chloride ea pele (mohlala, Na (+) - K (+) - 2Cl (-) cotransporter [NKCC1] e hlahisa liphetoho tse sa feleng maemong a potoloho ea cortical ha a le moholo.Wang le Kriegstein, 2010). Kahoo, liphetoho tse kholo mosebetsing oa li-neuronal nakong ena ea phetoho li ka hlakola boemo ba kemolo bo sa feleng.

Li-neurons tsa GABA li bapala karolo ea bohlokoa ho hokahanyang tšebetso ea cortical ka tsela e rarahaneng ea tšebeliso ea lijo le litlatsetso tse laolang phallo ea tlhaiso-leseling lipakeng tsa palo ea methapo ea methapo ea methapo (pyramidal neurons)Constantinidis et al., 2002; Di Cristo et al., 2007). Ketso tsena tse thibelang kelello tsa GABA tse holileng tsebong hammoho le nts'etsopele ea ts'ebetso e rarahaneng ea temohisiso (Luna et al., 2010) le ho eketseha haholo nakong ea bohlankana ho batho (Lewis et al., 2004),Cruz et al., 2003; Erickson et al., 1998) kapa ka likhoto (Tseng le O'Donnell, 2007). GABA e fumaneha haholo-holo liphoofolong tse tharo tse fapaneng tse bontšang calcium liprotheine tse tlamang parvalbumin, calbindin le calretinin. Immunohistochemistry ea liprotheine tsena tse fapaneng e ka sebelisoa ho lekola nts'etsopele ea GABA. Mohlala, li-neurons tsa parvalbumin-immunoreactive le GABA membrane transporter (GAT1) maemong a sa rateheng a ho phahama ha batho butle-butle, li nyoloha kapele bophelong 'me li lula li phahamisitsoe ho fihlela likhoeli tsa 15 tsa lilemo, ebe li fahla nakong ea bocha ho ba maemong a batho ba baholo (Anderson et al., 1995; Conde et al., 1996; Cruz et al., 2003). Ntle le moo, liprotheine tse hlalosang lipallo tsa GABA ho li-cortical pyramidal neurons (mohlala, likarolo tse ngotsoeng ka lebitso la "gephryin" le "karolo" ea axon) nakong ea bocha (Cruz et al., 2009). GABA e hokahanya tlhahisoleseling ea sele ea pyramidal ka ho feto-fetola lebelo la litlatsetso tse fapaneng libakeng tsa cortical (glutamatergic e ngata). Ts'ebetso ena e bonahala hantle ka ho fetisisa ha tlhaiso-leseling e phahameng e kenyelletsang mabaka a sa utloahaleng nakong ea phetoho lipakeng tsa bocha le ho ba motho e moholo. Ha e kopantsoe, bokaholimo ba bokhachane bo bopehile haholo ka ts'ebetso e khahlisang le GABA e tlatsetsang ts'ebetsong ena pele ho nako ea bophelo pele e fetoha thibelo nakong ea bocha.

2.1.2 Heterosynchrony le prun

Heterosynchrony nts'etsopele ea bokong e bua ka liphapang tsa tikoloho nakong eo ea ho rola nakong ea thupelo. Ho hlahisa le ho lema haholo ho sa tsoa bonoa haufinyane ka lithuto tsa ho nahana ka mokhoa oa sebopeho (Giedd et al., 1999a; Giedd et al., 1996b; Giedd et al., 1996c; Sowell et al., 2002; Sowell et al., 2001; Sowell et al., 2004; Tau le Peterson, 2010). Liphetoho tsa bophahamo ba litaba tsa bokhutšoaane joalo ka ha li sibolotsoe le MRI li bontša mokhoa oa tlhahiso e fetang le ho itlhohlolla kamora moo le maemo. Liphetoho tsena li bontša liphetoho tse feto-fetohang ka ho fetisisa tsa kutlo, kaha tsena ke ntlha e sa amaneng le neuron. Mokhoa oa MRI o lumella tlhahlobo e telele ea likaroloana tse ngata tsa boko kahare ho thuto e le 'ngoe, e ke keng ea khonahala ka mekhoa e meng. Lithuto tse joalo tsa nako e telele li fane ka limmapa tse hlakileng tsa hore na heterosynchrony e shebahala joang ka filimi e feteletsoeng ke nako (http://www.loni.ucla.edu/~thompson/DEVEL/dynamic.html). Ka har'a cortex, mokhoa ona o mosesaane oa ho faola o hlaha ka morao o ka pele, ka meaho ea pele e ntseng e ntlafala ea sensorimotor cortex e qala pele, ebe mekhatlo ea mekhatlo e ikopanye le mapolanka a pele-pele a ntseng a tsoela pele (Paus et al., 2008). Liphuputso tse entsoeng ka mor'a lefu la motho li bontša hore Ka hare likarolo tse fapaneng tsa libaka tsa pono, somatosensory, motor, le libaka tsa pele, leha ho le joalo, li etsahala ka nako e tšoanang (Rakic ​​et al., 1986).

Ka tloaelo, libaka tse subcortical li hola pejana ho feta libaka tsa cortical (Tau le Peterson, 2010). Amygdala e ka ba e 'ngoe ea libaka tsa pejana ho nts'etsapele le ho nts'etsapele ka mokhoa o hlephileng oa ho kopanela liphate. Banana, amygdala e bonts'a phetoho e fokolang ea bophahamo ba litaba tsa bohlooho nakong ea bocha, kaha e fihla bophahamo ba eona bo phahameng ka 4 ea lilemo li; ho bashanyana, bophahamo ba li-amygdala butle-butle bo eketseha ho fihlela lilemo tsa 18 ka 53%. Likarolo tse ling, ho kenyelletsa le caudate, putamen, le cerebellum li bonts'a sebopeho sa "U" se nang le molumo oa bohlooho se fumanehang ka nako ea bocha se nang le lipalo tse fokotsehang ka 15% (e ​​hlahlojoe (Durston et al., 2001)). Likaroloana tsa sebopeho se fanoeng li boetse li senotse liphetoho tse amanang le lilemo tse hlahelletseng (Gogtay et al., 2006). Lithuto tsa pele tsa hippocampus tse nang le MRI li bontšitse keketseho e itekanetseng ea molumo (12%) ho fihlela lilemo tsohle. Ho hlahlojoa bocha ha data ena lilemong tse leshome hamorao ho senola liphetoho tse makatsang kahare ho karohano. Mohlala, likarolo tsa ka morao tsa hippocampus li bonahala li hlahisa le ho nka lintho tse putsoa haholo ho feta likarolo tse ka ntle (Gogtay et al., 2006; Insausti et al., 2010).

Ho fapana hoa libaka tse kang tsena ho supa linako tse fapaneng tsa ho ba kotsing ea ho rohakoa ho ka ba teng tse sa ananeloang ka botlalo ka lebaka la ho fetoloa hoa sebaka sa boko se fanoeng (Andersen, 2003; 2005; Andersen le Teicher, 2008). Boithuto mabapi le litlamorao tsa ho pepeseha litsietsing nakong ea bongoana li bonts'a phokotso ea kakaretso ea 12-15% ho molumo oa litaba tsa grey ho batho (mohlala, Bremner et al 1997), haholo-holo, lihlahlobo tsena li shebile haholo lintlha tsena tsa kamora nako tse fetang tse kholo. liphetoho tsa kholo. Heterosynchrony nts'etsopele ka har'a maemo a mangata a tekolo (mohlala, sebaka, subregion, le dikarolo) e hloka ho tsotelloa ha ho ithuta nts'etsopele e tloaelehileng kapa nts'etsopele e fetotsoeng kamora ho rohaka.

Le ha MRI e bile bohlokoa bakeng sa ho lekola liphetoho litabeng tsa bohlooho ho fihlela bokong bohle, mokhoa ona o fana ka kutloisiso e fokolang ea liphetoho tse matla tse etsahalang kahare ho lits'ebetso tse fapaneng tsa methapo ea kutlo. Litekanyo tsa litaba tsa Grey li bonts'a likhakanyo tse fosahetseng tsa letsoalo la synaptic tse sa bontsheng liphetoho tse sebetsang tse hlahang nakong ea kholo, joalo ka tse boletsoeng ka holimo. Leha ho le joalo, tlhahlobo ea polelo ea liphatsa tsa botona tsa botšehali nakong ea bongoana liseleng tsa motho ka mor'a lefu (ke hore, mokhoa o hlaselang o ke keng oa khoneha le MRI) o ka fana ka lintlha tse ling mabapi le mofuta oa liphetoho tse etsahalang nakong ena. Mefuta e amanang le ts'ebetso ea nts'etsopele ea neuronal, ho kenyelletsa tataiso ea axon, morphogeneis le synaptogeneis, e fokotseha lilemong tsa bocha ho likhoto (Harris et al., 2009). Mehlala e ikhethileng e kenyelletsa li-netrins, semaphorin, neuropilin, neurexin le neurolignin. Liphetoho tse amanang le lilemo ho neurexin li tsamaisana le "axon" e khethollang ho chona le ho fokotseha hoa bohlokoa ha polelo ea gene e hlokometsoeng lipakeng tsa matsatsi a 45 le 90 ho rat (Cressman et al., 2010). Tlhatlhobo ea sehlopheng sa polelo ea liphatsa tsa lefutso le microarray e ka fana ka leseli mabapi le mefuta e mecha e amehang ho hlahiseng bongoana le ho poma. Tlhatlhobisong e joalo, liphatsa tsa lefutso li hlophisitsoe ka lihlopha tse tharo tsa mantlha tse sebetsang: Xtolkeletal cluster (25 e khethollotsoe), sehlopha se amanang le Ras / GTP (12 e hlalositsoe), le lipid metabolism le li-process tse amanang le steroid (13). Sehlopha sa cytoskeletal se ntlafatsa boemo ba ho hlophisoa bo ka morao ho bocha bo bang teng lilemong tsa bocha, sehlopha sa Ras / GTP se boetse se fana ka tlhahiso ea liphetoho, ha sehlopha sa boraro se ka etsahala se bonts'ang liphetoho tse amanang le bophelo ba ka. Kamora nako, lipalo tsa bocha tsa "protein" adhesion molecule (NCAMs) ea bocha li bontša hore liphatsa tsena tsa lefutso li hlahisoa ka tsela e tšoanang le liphumano tse fumanehang (Cox et al., 2009).

Ha se liphetoho tsohle tse hlahisoang ke mofuta oa gene tse amanang le liprotheine tsa sebopeho. Mohlala, liphatsa tsa lefutso tse amanang le phetoho ea li-glucocorticoid receptors nakong ea bocha (Perlman et al., 2007; Pryce, 2008). Ho batho le li-primates tse seng tsa batho, li-glucocorticoid receptors lia eketseha ebile li le tlhoro nakong ea bocha. Leha ho le joalo, li-isoforms tse fumanehang ho glucocorticoid receptors (GR) li bontša li-trajectories tse fapaneng, ka GR isoforms GRalpha-A le 67-kDa GRalpha e nyoloha ka har'a bana ba banyenyane hape hape nakong ea bocha; ka lehlakoreng le leng, litlhoko tse fapaneng tsa GRalpha-D li hola pele ho nts'etsopele 'me li fokotseha kamora moo (Sinclair et al., 2010). Liprotheine tsena tsa GR li hlahisoa haholo li-neuron tsa "pyramidal" empa li bonts'a polelo ea nakoana ho taba e tšoeu ea linaleli ka neonatally.

Tlhahlobisong e ikhethileng ea liphatsa tsa lefutso tsa 2,979 e ka hlalosang heterochrony (ke hore, mefuta ena e hlahisoa ka mokhoa o fapaneng lipakeng tsa libaka, khetlong lena, dorsolateral PFC le ntlha ea caudate bathong), 58% ea li-genes li ikarabella bakeng sa phetoho butle butle lipakeng tsa cortical le subcortical libaka (Somel et al., 2009). Ho ile ha hlahlojoa liphatsa tsa lefutso bakeng sa phapang ea mefuta lipakeng tsa batho le lichimpanzi mabapi le lefu la heterochrony le kamora ho hlaha. Lichimpanzi li arolelana kholo ea batho le batho, empa li na le mokhoa o khuts'oane oa bophelo, o fanang ka mokhoa o mong oa ho utloisisa heterochrony. Papisong ena, polelo e tšoanang ea liphatsa tsa lefutso e arohana lipakeng tsa mefuta e sa le qalong ea kholo ea thobalano (Somel et al., 2009), le liphetoho tse amanang le nts'etsopele ea litaba tse putsoa.

2.1.3. Ho itšetleha ka thobalano

Lithuto tsa morphology tsa MRI ho batho li bonts'a hore ba batona ba na le bophahamo ba khoele ea 9% ho feta ba batšehali, ba nang le phapang e fapaneng ea thobalano e bonahang meahong e tlatselletsang ea (Giedd et al., 1996a). Mookotaba oa caudate o moholo ho basali, empa phapang e eketsehileng e bonoa litekanyetsong tsa kholo. Boholo ba li-amygdala bo eketseha ka potlako ho ba batona ho feta ba batšehali, ka lehlakoreng le leng le bonoa ka boholo ba hippocampal. Caudate e tona e thella ka boholo, athe boholo ba caudate ba mosali ha bo fetohe haholo ho theosa le lilemo (Giedd et al., 1996a). Caviness et al (Caviness et al., 1996) e entse tlhahlobo ea MRI ea volumetric e bonts'ang hore subcortical forebrain nuceli (neostriatum) ho basali ke bophahamo ba batho ba baholo pakeng tsa lilemo tsa 7-11. Ho fapana le hoo, lits'oants'o tse tšoanang ho banna ba lilemo tse lekanang li kholo ho bophahamo ba tsona ba batho ba baholo, mme ka moelelo o tlameha ho khutla pele e ba motho e moholo. Ka ho ba moholo ka hara rat, banna ba baholo ba na le 18% e kholo ea ventral medial PFial (mPFC) ho feta basali, seo se bakoa ke li-neuron tse peli (13% tse amanang le ba batona) le lisele tsa glia (18%) (Markham et al., 2007). Liphetoho tse tšoanang li se li hlalositsoe ho "cortex" ea mantlha ea rat, moo banna ba nang le "20%" molumo oa taba e sootho ka lebaka la karolo ea 19% ho feta ea basali (Nunez et al., 2002; Reid le Juraska, 1992).

Tsela eo liphapang tsena li susumetsang tšebetso e sebetsang ka eona ke mohopolo feela. Ho faola limela ka boeona ho lumeloa ho tsamaisa tšebetso ea (Changeaux et al., 1976; Purves le Lichtman, 1980). Hang ha marang-rang a li-neuronal a thehiloe bokong bo ntseng bo hola, tlhokeho ea marang-rang ha e sebetse ebile li-synapses lia pongoa. Joalokaha ho boletsoe kaholimo, phokotso ea letsoalo la synaptic le nomoro ea sele ho lumeloa hore e eketsa ts'ebetso ea ts'ebetso. Liphetoho tsena tsa sebopeho li boetse li bapisoa le phokotso ea ts'ebeliso ea tsoekere (sesupo sa tšebetso ea boko; e tšohloang ka tlase ho Karolo ea 4.1), e phahameng haholo bongoaneng le bocheng pele e faoloa. Litlamorao tsa ts'ebetso ena li bonahala haholo ha li sa tsamaee hantle. Caudate e tona e faola limela tse amanang le kotsi e kholo ea mathata a amanang le tloaelo le makoloi, ho kenyeletsoa Tourette's Syndrome le khaello ea tlhokomelo ea khatello ea maikutlo (Teicher le Andersen 1995). Libaka tse amanang le tloaelo li kanna tsa ntlafatsoa ka nako ea ho fetoha ha mmele; libaka tse ling tse amehang mekhatlo e mecha le memori e ntseng e nchafatsoa khafetsa e ka se sebetse ka mokhoa o tšoanang (Teicher et al. 1995). Li-neuron tse fokolang sebakeng leha e le sefe, ho kenyeletsa le mPFC, li ka eketsa ts'ebetso ho sebetsa ka lebelo.

Liphapang tsa botona le botšehali li ka hlophisoa esale pele ka bophelo ke lihormone tsa gonadal tse bopehang bokong ba bongoana (bo sa tsoa hlahlojoa ho Viveros et al, 2010). Nakong ea bokhachane, phetoho ea li-androgens ho estrogen ka neural aromatase li kenya letsoho litlamorao tsa gonadal steroids mosebetsing oa boko, ho kenyelletsa phapang ea thobalano ka "masculinizing" boko ba basali (MacLusky et al., 1994). Polelo ea pele ea libaka tse hloahloa tsa androgen tse kopanyang li-androgen le li-enzymes tsa metabolic li fumanoa nakong ea nts'etsopele ea pele ho hypothalamus, amygdala, dorsolateral and orbital PFC le somatosensory cortex (ka maemo a sa ts'oanang le a batho: Clark et al 1989; rat: Reid le Juraska, 1992)). Ts'oaetso ea testosterone bokong e etsa hore ho be thata le ho feta ho tseba hore na ke lihormone tsa thobalano tse ikarabellang bakeng sa phapang ea thobalano. Liteko tse sebelisang androgen e sa aromatizable, 5cy-dihydrotestosterone (DHT), li thusa ho pepesa litlamorao tsena tsa steroidal, empa ts'ebeliso e joalo e lekanyelitsoe ho ithuteng mefuta e tlase kapa tlhekefetso ea chromosomal.

Litekong tsa tlhaho tse kenyelletsang ho hlohlona ha XPY ea chromosomal XXY (mohlala, Klinefelters), batho bana ba fokotse taba ea bohloeki ka har'a insula, gyri ea nakoana, amygdala, hippocampus, le libaka tsa cingulate (Giedd et al., 1996a). Tšobotsi ea morao-rao ho batho e tlaleha hore boholo ba litaba tsa grey e ne e amana hampe le litekanyetso tsa estradiol ho banana (r = −0.32) mme e ntle le maemo a testosterone ho bashanyana (r = 0.32) (Peper et al., 2009). Phapang ea tikoloho bakeng sa litlamorao tsa li-hormone, leha ho le joalo, e teng, joalo ka likamano tse matla lipakeng tsa boemo bo tlase ba "gyrus" le "estrogen" e tlase ho banana (r = -0.72). Ho kenyelletsa moo, manoplication a androgens kapele bophelong a hae a na le litlamorao tse sebetsang mabapi le ts'ebetso ea cortical. Mohlala, khethollo ea ntho, mosebetsi o amanang le PFC, e ntlafala ho batona ba tloaelehileng ba batona le basali ba pepesehileng ba androgen ba amanang le basali ba tloaelehileng (Clark le Goldman-Rakic, 1989). Ka lehlakoreng le leng, keketseho ea "pubertal" ea lihormone tsa thobalano e kenella pele ho ho teba, e ka ananeloang ke litlamorao tsa tlhophiso mosebetsing oa suboportical dopamine (Morris et al., 2010).

Boithuto ba Rodent bo fana ka maikutlo a hore neonatal estrogen e hatella tlhahiso e ngata ea li-neuronal libakeng tsa basali tsa basali (ho kenyeletsa le libaka tsa prelimbic le infralimbic) (Juraska le Markham, 2004; Markham et al., 2007), e fapaneng le litlaleho tsa pele tsa bokhoni ba estrogen ba ho hlohlelletsa arborization e pharalletseng libakeng tse ling tsa boko joalo ka hippocampus ho batho ba baholo (Hajszan et al., 2009; Toran-Allerand, 1996). Prepubertal ovariectomy e fokotsa methapo ea methapo ho basali, e ka 'nang ea hlalosa lintlha tse tlase tsa bohlooho ho basali ((Nunez et al., 2002). Maemo a ntseng a eketseha a testosterone nakong ea bokhachane e thusa ho itlhoekisa ha dendrite ka har'a amygdala ea monna ea lilemong tsa bocha.Zehr et al., 2006). Ka bobeli, lithuto tsena li fana ka maikutlo a hore li-hormone tsa gonadal li bapala karolo e rarahaneng ho pepesa boko ba bongoana.

2.2. Ho hlahisa haholo le ho poma lits'ebetso tsa li-receptor

Tlhahiso e ngata ea 2.2.1 ea li-receptor tsa monoamine

Ho hlahisa ka bongata le ho poma ha mekhoa ea li-receptor ho thata haholo ha ho bapisoa le liphetoho tsa synaptic, 'me ho hlaha maqhubu a mabeli a liphetoho tse amanang le lilemo tsa letsoalo. Tsamaiso e 'maloa ea li-neurotransmitter, ho kenyelletsa dopamine (Gelbard et al., 1990; Kalsbeek et al., 1988; Lankford et al., 1988; Todd, 1992), norepinephrine (Feeney le Westerberg, 1990; Kline le al., 1994) le serotonin (Kuppermann le Kasamatsu, 1984; Lauder le Krebs, 1978; Whitaker-Azmitia le Azmitia, 1986) Ho ba le likarolo tse sa lekanyetsoang tsa bokhelohi bokong. Polelo ea ectopic ea li-subtypes tse fapaneng tsa "receptor" nakong ea ho hola kamora nako ea bokhachane e amahanngoa le ho hola ho kopaneng ha synaptic, kholo ea axonal le sebopeho sa synapse. Mohlala, polelo ea ectopic ea li-receptor tsa serotonin 5-HT7 kahare ho hippocampus e etsahala hanyane nakong ea libeke tse peli tsa pele tsa bophelo litlelong (Louiset et al., 2006; Vizuete et al., 1997). Ka mokhoa o ts'oanang, serotonin transporter (5-HTT) e fumaneha ho li-neurons tse se nang serotonergic ka mokhoa o kopaneng le neuroepithelia ea cortical le striatal neuroepithelia le sensory thalamic pathways postnatally ho P0-P10 (Zhou et al., 2000). Polelo ea nakoana ea 5-HTT le vesicular monoamine transporter (VMAT) e ile ea boela ea bonoa methapong ea methapo ea kutlo, hippocampus, cortex ea "cerebral" septum le amygdala (Lebrand et al., 1998). Barekisi bana le / kapa li-receptors ho lumeloa hore li tataisa ho bonoa ha methapo ea methapo. Litlamorao tsa li-neurophransmitters tsa trophic li itšetlehile ka khatello ea kelello (Mazer et al., 1997), ho fana ka maikutlo a hore maemo a motheo a bohlokoa ka botlalo molemong oa ts'ebetso. Polelo e tšoanang ea ectopic receptor le eona e bonoa litabeng tse tšoeu. Mohlala, "renadrenergic receptor α2" e bonoa ka har'a lintho tse tšoeu tse bosoeu ho Rate (Happye al al., 2004). Leha ho le joalo, ha se polelo eohle ea sebui se sebetsang.

Leqhubu la bobeli la polelo le utloang haholo ha le le lilemong tsa bocha, nakong eo li-receptors le mekhoa e bonts'ang li bonts'a khokahano ea mofuta oa U-e tsoetseng pele e hlahisang maemo a polelo a mamellang ho ba motho e moholo. Ho fapana le "ectopic", poleloana ea nakoana eo motho a bang le eona ha e sa le motho e moholo. Tlhahlobo ea liphetoho tsa phetoho ea bocha e fumanoa ho Lethathamo 1, ka ho totobatsa li-receptors kahare ho likarolo tse maoto le maoto. Nako ea nako ea tlhahiso e ngata le ho rema e itšetleha ka tikoloho (Andersen et al., 2000), 'me e bonoa letoto la matšoao a mangata. Lisebelisoa tse fapaneng tsa li-receptor li kenyelletsa: dopamine, serotonin, norepinephrine, glutamate, GABA, neurotensin, endocannabinoid, le cholinergic (Andersen et al., 2000; Eggan et al., 2010; Lidow et al., 1991). Ha a le monkey rhesus, Lidow et al (Lidow et al., 1991) li bontšitse hore methapo ea li-receptor e hlaha ho konsarete le synaptogeneis.

Haeba re tsepamisa maikutlo ho li-microcircuits ho hlahloba phetisetso e amanang le lilemo tsa li-receptor, liphetho tsa morao-rao li bontša liphetoho tse rarahaneng le ho feta lilemong tsa bocha. Phetiso ea li-receptor ka boeona e fetoha pakeng tsa li-phenotypes tse fapaneng tsa neuronal. Mohlala, li-receptor tsa D1 dopamine ha li bonahale li fetola boemo ba tsona ba polelo haholo pakeng tsa lilemo tsa kamora ho khoesoa ho ba motho e moholo ho li-neurons tsa GABAergic (Brenhouse et al., 2008; Vincent et al., 1995). Ho fapana le hoo, ho eketsoa le ho pongoa ha li-receptor tsa D1 ho etsahala haholo ho li-neurons tse hlahisang glutamatergic (Brenhouse et al., 2008). Ka ho khetheha, ke 2% feela ea likhakanyo tsena tsa glutamatergic tse sebetsang tsa D1 ho litoeba tsa bana, li nyolohelang ho 44% ho P40, 'me li oela tlase ho 6% ka kholo ea P100. Hore na li-receptors tse ling li bonts'a polelo e khethollang ho li-subtypes tse ling tsa neuronal nakong ea bocha li hloka ho hlahlojoa. Lethathamo 1 e fana ka tlhahisoleseling mabapi le liphetoho tse ling tsa lihlopha tsa li-receptor, empa ho tsebahala ka mefuta e khethehileng ea neuronal hangata ha ho tsejoe. Ka lehlakoreng le leng, li-receptor tsa D2 li thibela tšebetso ea li-interneurons tse potlakileng tsa Gaba kamora ho kena bohlankaneng (O'Donnell, 2010; Tseng le O'Donnell, 2007). Li-neuron tsena ke tsa bohlokoa bakeng sa ho kopanya ka nepo lipalo tse ngata ka nako ea sebele. Ka hona, kabo ea li-receptor kahare ho li-microcircuits le matla a tsona a sebetsang a fetoha haholo nakong ea bocha.

Ho itšetleha ka thobalano ea 2.2.2

Bopaki ba pele ba phapang ea thobalano ho polelo ea li-receptor bo tsoa thutong ea PET ea motho moo DA le 5HT recensor density e theohelang haholo ho banna ho feta basali ho tloha lilemong tsa 19-30 (Wong et al., 1984). Re boetse re bonts'itse phapang ea thobalano ho striatum nakong ea bonyenyane lilemong tsa bocha, le basali ba bonts'a tlhahiso e ngata hape e sa sebetseng haholo (Andersen et al., 1997). Mohlala, letsoalo la li-receptor tsa D2 le eketsehile 144 ± 26% ho banna ho bapisoa le 31 ± 7% ho basali pakeng tsa 25 le matsatsi a 40 a lilemo ho rat. Ka mokhoa o tšoanang, prunor ea preptor e ne e le kholo haholo ho banna ho feta basali mme e etsahetse lipakeng tsa matsatsi a 40 le 120 (motho e moholo). D1 striatal density e fokotsehile 34 ± 4% ho banna, empa ke feela 7 ± 8% ho basali. Bakeng sa li-bokella tsa li-nucleus, li-curls tsa "D1" tsa banna le tsa basali li ne li tšoana kamora matsatsi a 40, 'me e' ngoe le e 'ngoe e bonts'a ho hlapanya hanyane ka matsatsi a 80. Leha ho le joalo, liphapang tsa botona le botšehali tsa D1 receptor density li ile tsa phehella P120, moo li-receptors tsa D1 li neng li le kholo ho feta li-57.8 ± 21.2% ho ba batona ho feta basali. Ka kakaretso, ho ne ho se na phapang ea bong ho D2 density ho li-bokellase tsa nucleus. Phapang ea thobalano ea thobalano, leha ho le joalo, e ne e sa khonehe hore e sebelisoe ke manonyeletso a gonadal nakong ea bohlankana.Andersen et al., 2002). Gonadectomy hang-hang pele D1 le tlhahiso ea tlhahiso ea D2 e sa fetole khatello ea methapo ka kakaretso nakong ea bocha; le gonadectomy pejana bophelong. Liphetho tsena li fana ka maikutlo a hore ho pepeseha ha testopubertal ha ho khothalletse tlhahiso e ngata ea dopamine, hape estrogen ha e hatelle tlhahiso e ngata ho feta tekano. Mefokolo ea bohlahlobo e kanna ea ba le leseli la ho hlokomela liphetoho tse amanang le thobalano. Le ha Autoradiography e tšoaneleha hantle bakeng sa liphetoho tsa letsoalo la receptor tsa tikoloho ka kakaretso, mokhoa ona o hloleha ho senola hore na ke li-neuron life tse hlalosang li-receptor tsena. Kahoo, menyetla e lula e le hore liphetoho tse amanang le thobalano, le tlhaselo ea tsona ea li-hormone, li hlaha liphoofolong tse fapaneng tsa li-neuron tse seng li lokela ho ba teng.

Tlhahlobo ena e ke ke ea shebana le litlamorao tse sebetsang tsa liphetoho tsena tsa li-receptor, joalo ka tse hlahlobang karabelo ho li-agonists tse ikhethileng kapa bahanyetsi. Leha ho le joalo, ho bohlokoa ho hlokomela hore liphapang tsa thobalano ka mekhoa e tšoaeang maikutlo li susumetsoa ke li-hormone tsa gonadal, hape li ba le liphetoho tsa kholo nakong ea bocha.Andersen et al., 2002; Kuhn et al., 2001).

3. Khokahano

3.1. Kahare e ikhethang ea li-system tsa neurotransmitter

Karolong ena, re tšohla hore na li-system tsa neurotransmitter tse ikhethang li kenella joang bokong bo itseng ba boko. Ho tseba lintho tsa tlhaho ho qala ka tlhaho, empa ka mafolofolo ho tsoela pele nakong ea bocha le ho ba motho e moholo. Leha ho le joalo, liphuputso tse ngata ha li felle monahanong oa bocha 'me li nka hore bolulo bo tsoella ka mokhoa o hlakileng. Boithuto ba ho hokela setopo sa motho ka mora setopo bo batla bo sa khone ho tsamaisoa, ka ha litsi tsa methapo ea methapo ea kelello li hleka lisele tsa mmele libakeng tse nyane tse thibelang ho ts'oaroa hoa pampitšana. Tharollo ea MRI ha e lumelle ho ts'oaroa ha pampitšana manonyeletso a itseng a neuronal ho buisana le e mong le e mong (ntle le ka ho sebelisa tractography, e hlahlobang myelin le axon caliber ka nako e le ngoe). Tekanyo ea transporter hangata e sebelisoa e le sesupo sa mekhoa ea bolulo (mohlala,Moll et al., 2000)). Le ha ho le joalo, mesaletsa ea transporter e ka fapana ka boits'oaro ba ho iphumanela mme ka hona e kanna ea se ke ea tšoaneleha bakeng sa merero e joalo.

Ho ipapisitsoe le lithuto tse 'maloa tsa liphoofolo tse sebelisang mekhoa e metle ea ho ts'oara hore e be lilemong tsa bocha, tse ling li bonts'a tsoelo-pele ea tlhaiso-leseling nakong ea kholo.Brenhouse et al., 2008; Brummelte le Teuchert-Noodt, 2006; Cunningham et al., 2002; Erickson et al., 2000), athe ba bang (Cressman et al., 2010; Rios le Villalobos, 2004) bonts'a mokhoa o bopehileng oa sebopeho sa U. Re hlokometse tsoelopele ea mohala oa ho boloka polokelo ea mothapo oa "V glutamate" oa "" "" "" "" "" "" LIT "" "tsa" "methapo ea" methapo ea methapo "bokong ba li-nucleus accumbens pakeng tsa 25, 44 le matsatsi a 100 ho rat (Brenhouse et al., 2008). Phuputsong e entsoeng ke Cunningham le basebetsi mmoho (Cunningham et al., 2002), mokhoa oa ho boloka matlo ka mohala o boetse o fumanoe likamanong tsa glutamatergic lipakeng tsa amygdala le PFC, tse tsoelang pele ho tloha ho tsoalo ho fihlela lilemong tsa bocha / ho ba motho e monyane lilemong (matsatsi a 60) ho rat. Phapang ea lilemo khokahanong ea li-synaptic le eona ea sebetsa. Mohlala, li-neuron tsa glutamate li thehile axo-dendendritic (36.5%), axo-spinous (7.7%), le axo-somatic synapses (5.8%) ho li-neurons tsa GABAergic, empa 17.3%, 30.8% le 1.9% ho li-neuron tse seng tsa GABAergic. Ho theha mabitso ana hangata ho ne ho latela mokhoa o ntlafalitsoeng ho theosa le lilemo.

Ho fapana le hoo, mekhoa e meng ea bolulo e fana ka lithuto tse se nang mohala thekisong ea tsona. Mohlala, PFC ea medial (libaka tse ka pele tsa prelimbic le infralimbic) ho amygdala ea basolateral e lula e tsitsitse pakeng tsa matsatsi a 25 le 45 ea lilemo ho rat, empa e fokotseha ka 50% lipakeng tsa 45 le 90 (Cressman et al., 2010). Liphumano tse tšoanang li bonoa litoeba. Barekisi ba tsoang ho dorsalmedial thalamus ho ea pele ea cortex e eketseha ho fihlela matsatsi a 13, ba lateloa ke ho fokotseha ha 67% bekeng ea boraro ea bophelo, ha ba eketseha butle-butle ho fihlela bocha le ho tsitsa (Rios le Villalobos, 2004). Karolo ea pele ea tlhahiso ea matlo e se e hokahane le mokhatlo o sebetsang oa li-neuron tsa III, o fana ka maikutlo a hore glutamate input drive synaptogeneis. Li-neuron tsa Dopamine li latela mokhoa o ts'oanang oa ho beha sebakeng sa prort cortex (libaka tsa 4, 9, 46): li-axon tsa dopaminergic tse karolong ea III li ile tsa eketseha makhetlo a mararo pele ho likhoeli tsa 5-7, ntle le liphetoho tse ananelang likarolo tsa 1 le V (Erickson et al., 1998). Li-varicosities tse lebisitsoeng li ile tsa tsoela pele ho eketseha, tsa fihla tlhōrōng (tse thataro-kholo ho feta litšoeneng tse nyane) ho liphoofolo 2-3 ea lilemo li (bongoaneng) pele li theohela maemong a batho ba baholo ba tsofetseng (Rosenberg le Lewis, 1995; Woo et al., 1997). Gerbils o bontša mokhoa o tšoanang. Ho kenella kahare ho amygdala ho eketsa libeke tse tharo tsa ho qala ka bophelo ka likokoana-hloko, pele ho theoha ho honyenyane ha letsoalo nakong ea bocha e tsofalang hore e be motho ea holileng ea ho ba motho e moholo.Brummelte le Teuchert-Noodt, 2006). Kahoo, ho ka etsahala (mme ha e koaheloa ka nepo tlhahlobisong ena) hore lits'ebetso tse ling tsa li-neurotransmitter li bonts'a liphetoho tse ts'oanang mekhoeng ea polokelo.

Mothating ona, ha ho hlake hore na ke hobaneng ha mekhoa e fapaneng ea bolulo (mohlala, mola o bapileng le o entsoeng ka popego ea U) e hlaha ka likarolo tse fapaneng tsa cortical (Setšoantšo sa 2). Monyetla oa pele o ka fumanoa ke mehlala ea lilemo, moo ho ka 'nang ha e-ba le likhohlano tse tebileng tse neng li sa hlahe ka nepo. Monyetla oa bobeli o ka sebopeho sa tikoloho e sebetsitsoeng. Re hlahisitse khatiso ena pejana maemong a li-dopamine receptors (Teicher et al., 1995) le tse ling bakeng sa boluloErickson et al., 1998). Ka ho khetheha, libaka tse fapaneng tse kenyellelitsoeng mesebetsing tse hlokang ho ntlafatsoa khafetsa li ka rua molemo ka ho eketseha ha mela ho etsahalang pejana bophelong (pele ho nako ea bocha). Ka lehlakoreng le leng, libaka tse amehang ho ithuteng mosebetsi o mong oa bophelo bohle, joalo ka tloaelo, li rua molemo ka ho o hlophisa o amanang le ho faoloa. Monyetla oa boraro ke hore bolulo bo bonts'a methati e ikhethileng ea lilemo mokhatlong oa laminar, 'me karolo ea III ka sethaleng e bonts'a sebopeho se senotsoeng sa U, le likarolo tse tebileng le tse sa bonahaleng tse bonts'ang sebopeho se tsoelang pele. Ha ho kopantsoe hammoho, khokahano e ikhethileng ea litumellano tse kenang kahare le tse tsoang ho batona li thusa ka ho hlaka ho betla methapo ea methapo nakong ea bocha (Benes, 2009).

Setšoantšo sa 2 

a) Litšoantšo tsa "cortical lamination" tse emeng ka nqane ho sefapano se emisitsoeng ke Santiago Ramon y Cajal ka mor'a Nissl (ka ho le letšehali, bohareng) ho motho e moholo le Golgi stain (ka ho le letona) ka lesea le nang le khoeli la 1½. B) lipaterone tsa liphetoho tse lumellanang tse etsahalang nakong ea phetoho ...

3.2.1 Myelination

Tsoelopeleng eohle, boholo ba phaello e felletseng ea bophahamo ba bokong bo tsoa lipapatsong tse tšoailoeng tsa lipampitšana tsa fiber (Benes et al., 1994). Myelination e eketsa lebelo la phapanyetsano ea tlhahisoleseling, mme bonyane e ikarabella bakeng sa ho hlaha ha "repertoire" ea boitšoaro bo botle ba bo-mmeMasimo, 2005). Myelination bokong ba motho e fapana ka thobalano le sebaka (Benes et al., 1994; Giedd et al., 1999b). Myelination e tsoela pele hanyane ka hanyane le hanyane ka hanyane ka bong ba batona, ho latela lithuto tsa kamora lefuBenes et al., 1987) le lithuto tsa MRI tse hlahlobang liphetoho tse joalo ka ho khetholla taba e tšoeu le e putsoa (Paus et al., 1999) kapa ka tšebeliso ea menahano ea lits'oants'o tsa bomolimo (DTI) (Paus et al., 1999). Boholo ba tse tsejoang ka liphetoho tsa ntlafatso ea myelination li thehiloe ho lithuto tsa Corpus callosum, pampitšana e kholo ka ho fetisisa ea myelin bokong (mohlala, (Keshavan et al., 2002; Teicher et al., 2004)). Ho fapana le liphetoho tsa taba e bohlooho, mokhoa oa rostral-caudal oa taba e tšoeu o ntse o tsoela pele ho eketsa boholo ba li-callus ho ba motho e moholo (Giedd et al., 1996a). Liphetoho tse amanang le lilemo li hlaha karolong e ka morao ea (Paus et al., 1999). Lipampitšana tse ling tse tšoeu, e leng capule e ka hare le setšoantšo se tlohang ka ho le letšehali, li tsoela pele ho ipabola ka ho fetoha ha mmele. Delel myelination of frontocortical connances e etsahalang nakong ea lilemo tsa bobeli le tsa boraro bathong e kanna ea amahanngoa le taolo e ntlafalitsoeng ea boitšoaro le taolo ea tšusumetso e hlahang kamora bocha (Luna et al., 2010; Paus, 2005).

DTI e hloaea likhakanyo tsa motsamao oa metsi, ka mehato ea matla a ho fapana (MD) le anisotropy (FA). Ka har'a voxel e fanoeng, mehato ea FA e fapana ho tloha 0 (hantle-ntle isotropic infusion) ho 1 (infishene ea anisotropic e phethahetseng), 'me e khethiloe ke bophara ba fiber le segokanyipalo, tumellano le boemo ba myelination (Basser le Pierpaoli, 1996). FA e hlahloba tekanyo ea ho tsamaisoa ha metsi. Ho tsamaea ha metsi ka nqa e le 'ngoe, joalo ka se etsahalang ka pampitšana, ho na le boleng bo phahameng ba FA. Tšobotsi e pharalletseng ea kamoo MD le FA li fetohang ho theosa le lilemo (5-30 lilemo) libakeng tse fapaneng tsa boko li ka fumanoa litlalehong tsa Lebel et al (Lebel et al., 2008) le Qiu et al (Qiu et al., 2008). Likarolong tse tšoailoeng pampiring ea Liebel et al, tahlehelo e tebileng ka ho fetisisa ea MD e etsahala ka har'a leseli la caudate nakong ea bocha athe splenium ea corpus callosum e fihlella tahlehelo ea eona e felletseng (~ 8%) pele ho lilemo tsa 15. Leha ho le joalo, litekanyo tsa FA li bonts'a ho feta ho ts'oaroa feela, 'me li kenyelletsa likhakanyo tsa phapang ea mofuta oa lipampitšana tsa fiber (ka mohlala, ho lekanya ha li-axon ka bomong le ho lekana ha tsona ka "botsena"; Paus, 2010). Ka hona, liphetoho tse hakantsoeng tsa myelination tse ipapisitseng le mehato ea FA li hloka ho nahanela bophara ba myelin le axon. Sekhahla sa `g '(bophara ba axon: bophara ba axon + botenya ba myelin sheath) se entsoe hore se ikarabelle ka bophara ba axon le bophara ba fiber. Kaha bobeli ba axon le botenya ba myelin bo ama lebelo la ho khanna empa li sa nyolohe ho isa tekanyong e ts'oanang kamora ho kena bohlankaneng, karolelano ea `g 'e kanna ea bonahatsa liphetoho tsa nts'etsopele litabeng tse tšoeu le conductivity (Paus le Toro, 2009). Ho lekola sekhahla sa myelination le kamano ea eona ho bophara ba axonal ho hloka microscopy ea elektrone. Ka tekanyo, mehato ea khatello ea maikutlo e sa khetholleng e bonts'a hore palo ea lisele tsa glial e fetoha ka tsela e itšetlehileng ka tikoloho. Palo ea sele ea Glia e tsitsitse ho PFC e tsitsitseng pakeng tsa bocha le ho ba motho e moholo, empa e eketsa hoo e batlang e le 40% e nang le phapang ho PFC ea dorsal (Markham et al., 2007). Ka hona, liphetoho ho DTI li bonts'a liphetoho tsa bophara ba glia le axonal bophara.

Mokhoa o mong oa ho khetholla liphetoho mqoqong ke ho hlahloba mofuta oa gene. Tumellanong le litekanyo tse ntlafalitsoeng haholo tsa anatomical, liphatsa tsa tlhaho tse amanang le myelination le tsona li eketsa polelo nakong ea bocha bathong (Harris et al., 2009). Mohlala, liphatsa tsa lefutso tse kenyeletsang MBP (protheine ea mantlha ea myelin), MOG (myelin oligodendrocyte glycoprotein), le MAG (myelin e amanang le glycoprotein) e eketsa polelo ea bona ka ho hola. Le ha MBP le MOG li amana le liphetoho tsa sebopeho sa myelin, MAG e kentse letsoho ho kopanya axonal caliber (ketso) le degree ea myelination (Yin et al., 1998). Ha ho kopantsoe hammoho, masapo a taba e tšoeu a eketseha ka tsela e tsoetseng pele, ea kahare e fapaneng le sebopeho sa molora sa U sa mofuta oa bohlooho se tšoaeang bohlankana.

3.2.2 Bothata ba ho ts'epahalla thobalano

Liphapang tsa botona le botšehali li hlaha ho myelination mme li bonoa nakong ea ho qala bongoana. Lithuto tse ngata li bonts'a keketseho e kholo mokhoabong oa likarolo tse ngata tsa boko nakong eohle ea bocha ho ba motho e moholo ho banna, empa eseng basali.Blanton et al., 2004; Leussis le Andersen, 2008; Paus, 2010). Ho fapana le moo, ho bonahala eka myelination e hlaha pejana ho basali. Mohlala, liphapang tsa botona le botšehali ho tsa pelehi ea hippocampal myelination li hlaha kamora lilemo tsa 5, ka karolelano ea 37% tekanyo e kholo ea myelination ho basali ho feta ba batona (Benes et al., 1994). Phapang e tšoanang ea botona le botšehali e bonoa ho mefuta eohle (mohlala, batho, likhoto (Kodama, 2008). Ha e se e le motho e moholo, myelination in the corpus callosum e kholo ho banna, leha basali ba na le lisele tse fokolang tsa glia tse kenyang letsoho (Nunez le Juraska, 1998; Kim et al 1997). Ka mokhoa o ts'oanang, "NFC" ea rat e na le lisele tsa "glia" tse nyenyefalitsoeng ka tlase ho basali ho feta banna ka ho ba motho e moholo.Markham et al., 2007).

Ha tlhahlobo ea DTI e arotsoe likarolo tsa FA le MD, lipapatso tse fapaneng li ba teng lipakeng tsa mehato, thobalano, le sebaka ka seng (Asato et al., 2010). Lipampitšana tsa fiber tsa arcuate fasciculus (e hokahanyang sebaka sa Wernicke le sebaka sa Broca) le libaka tse tlase tsa fronto-occipital fasciculus (e hokahanyang sensorimotor le libaka tse ka pele) li bonts'a keketseho ea FA ho banana, empa e fokotse FA ho bashemane ba pakeng tsa lilemo tse 6-20. ; ha ho phapang ea thobalano e neng e bonoa bakeng sa MD (Ashtari et al., 2007; Schmithorst et al., 2008). Liphetoho tsena li amana le IQ le ts'ebetso e phahameng ea mantsoe ho basali ba batona ho feta banna (Ashtari et al., 2007; Schmithorst et al., 2005). Ka lehlakoreng le leng, lipampitšana tse ling li sitoa ho bontša keketseho e lebelletsoeng e amanang le lilemo tsa FA, athe MD e fokotsehile (Eluvathingal et al., 2007). Mehato e bonts'ang keketseho ea FA ha ho se na liphetoho liphihlelong tsa radial (index ea demyelination) e ka supa phetoho ho tloha ho tlokotsi e fokotsoang ho ea mokhatlong o moholo oa axonal fiber (kapa likhoele tse thata) nakong ea bohlankana.Ashtari et al., 2007). Ts'ebetso e sebetsang haholoanyane e ka ba litholoana tse boletsoeng esale pele tsa liphetoho tse joalo.

Methati ea testosterone e amana le liphetoho tsa `g 'ho banna ba batho (Perrin et al., 2008). Sekhahla sa `g" se eketseha ho banna, empa ha se fetohe ho basali (Paus le Toro, 2009). Liphetoho tsa "calic" ea Xxonal nakong ea nts'etsopele li ka 'na tsa hlalosa keketseho ea DTI ho banna, athe liphetoho tsa basali ho DTI li ka bonahatsa myelination (Perrin et al., 2009) Boithuto ba mantlha bo bonts'a hore "femos" ea "femosum" e ameha ka lihormone tsa lefutso, 'me li-ovariectomy ka matsatsi a 20 a le lilemo ka rateng li fokotseha palo ea li-axel tsa myelinated ha li bapisoa le li-control (Yates le Juraska, 2008); palo eohle ea li-axons thutong ena ha ea ka ea ameha, ho fana ka maikutlo a hore liphetoho tsena li entsoe ka lebaka la tahlehelo ea myelin eseng lisele. Tlhaloso e le 'ngoe ke hore phapang ea thobalano e teng nakong ea ho phela ha li-oligodendrocyte, moo lisele li shoang kapele ho basali ba batona ho feta ba batona (Cerghet et al., 2006). Monyetla o mong o kenyelletsa litlamorao tsa estrogenic tse fetolang li-hormone tse ling tsa gonadal (mohlala, progesterone), li-hormone tse amanang le khatello ea maikutlo, kapa le lintho tse holang tseo ka tsona li hlahisang myelination (e tšohliloeng ho Yates le Juraska, 2008). Patlisiso e eketsehileng e tla tlatsa likheo tse sieo tsa mak'henike kamoo estrogen e fetolang myelination.

Re se re qalella ho utloisisa hore na synaptogeneis le prunun li sebelisana joang le lits'ebetso tsa myelinating le tšebetso ea boko ho etsa boitšoaro ba bocha (Paus et al., 2008). Myelin o bapala karolo ea bohlokoa ntlafatsong, empa ho bohlokoa le ho feta, ho hokahanya lebelo la lipakanyo tse fapaneng ho tloha libakeng tse fapaneng ho ea sebakeng se itseng. Ho supa Synchronous ho bohlokoa haholo hore tsoelo-pele e tloaelehileng e tsoele pele (Masimo, 2005), le liphetoho tse bakoang ke myelination li kenyellelitse maloetse a mangata a kelello.

4.0. Ts'ebetso ea liphetoho tse sebetsang

Tlhahlobo ena e koahetse liphetoho tse hlophisitsoeng tse etsahalang nakong ea bongoana ho ea ho liphetoho tsa batho ba baholo, empa liphetoho tse sebetsang li ka bonts'a mekhoa ea bona. Boko bo holang bo sebelisa sebopeho sa eona sa lisebelisoa le lisebelisoa (mohlala, metabolism ea glucose) ho buisana lipakeng tsa likarolo tse ka hare ho susumetsa boitšoaro. Kamoo likarolo tsa boko li sebetsang ka tsela e fapaneng ho arabela ts'usumetso e fanoeng li ka boela tsa re bolella hore li hokahane joang ka tšebetso. Karolong ena, "khokahano e sebetsang" joalo ka ha e lekantsoe ke MRI e bua ka likamano tse amanang pakeng tsa libaka tse peli.

4.1. Ts'ebeliso ea matla

Liphetoho tsa morphological tse hlalositsoeng kaholimo li etelletsoa pele ke liphetoho tse sebetsang kahare ho bokong. Lithuto tsa mantlha tse mabapi le liphetoho tse sebetsang li sebelisitse PET ho nahana ka tsoekere ho hlahisa tšebeliso ea matla ka moralo oa likarolo (Chugani, 1998; Feinberg, 1988). Ts'ebeliso ea glucose bathong e fihla maemong a batho ba baholo ka lilemo tse peli tsa bophelo (Chugani et al., 1987) empa ebe e nyoloha a le lilemo li 4-5 ebile e boloka sehlaba sena ho fihlela a le lilemo li 10 pele a hlaha ka ~ 50% ka 16-18 ea lilemoChugani, 1998). Liphatsa tsa lefutso tse amanang le metabolism ea glucose, mohlala, gene acyl coA dehydrogenase (ACADSB), li hlahisoa ka maemo a phahameng nakong ea bocha, leha bohlokoa ba bona ba ts'ebetso bo sa tsejoe ka nako ena (Harris et al., 2009).

Matšoao a mang a tšebetso ea boko a hlahlobang metabolism ea boko, joalo ka n-acetylaspartate (NAA; letšoao la li-neurons le lits'ebetso), phosphocreatine (PCr; matla a khoheli), le membrane phospholipid metabolism (e nang le baetsi ba sPME le sPDE) e hlahlobiloe ka matla a magne imagon resonance imagroscopic imaging (MRSI) ho fana ka index ea tlhahiso e sa hlaseleng. Liphetoho ho mats'oao ana li ile tsa bonts'oa ho methapo ea methapo ea boko ho banna le basali ba 6-9.5, 9.5-12, le 12-18 ea lilemo li le ho liithuti tsa 106 (Goldstein et al., 2009). Lipapiso pakeng tsa 6-9.5 ea lilemo li 12 le 18 ea lilemo li bontša ha ho na phapang ho NAA, e bonts'ang liphetoho tse sa bonahaleng tsa "neuronal". Tlhahlobo ena e fapane ka ho toba le tahlehelo ea neuronal e bonts'itsoeng hantle ka tekanyo e otlolohileng lithaoleng tsa kamora lefu (mohlala, Huttenlocher, 1979). Leha ho le joalo, NAA e fana ka acetate bakeng sa olidgodendrocyte tse ikarabellang bakeng sa tlhahiso ea myelin. Kahoo, ha ho phetoho e kholo ho NAA ho ntlafatseng ea bongoana e ka bonts'ang ho leka-lekana pakeng tsa tahlehelo ea methapo le ho eketseha ha myelination. PCr e fokotsehile sehlopheng sa batho ba banyenyane, empa maemo a phahameng a pherese le sPME / sPDE ratios, a bonts'a phallo ea phospholipid, a ne a phahame. PCr le peresente ea lintho tse putsoa li ne li amana hantle le lilemo, empa NAA, sPME, sPDE, le sPME / sPDE li ne li se joalo. Leha liphetoho tse ka bang teng li kanna tsa fetoloa ka ho kopanya banna le basali, lintlha tsena li fana ka maikutlo a hore MRSI ha e bonts'e liphetoho tse amanang le lilemo tsa metabolic.

Khokahano e sebetsang ea 4.2 joalo ka ha e hlalositsoe le MRI

Khokahano e sebetsang ke mokhoa o mong o sebelisoang ho bonts'a likamano tsa nakoana lipakeng tsa libaka tsa ts'ebetso nakong ea ho phomola mmuso kapa nakong ea mosebetsi oa fMRI (Fair et al., 2008; Supekar et al., 2009; Thomason et al., 2009; Zuo et al., 2010). Limmapa tsa khokahano e sebetsang li boetse li bitsoa li-connomes (Biswal et al., 2010), ka lits'ebetso ho fMRI tse emelang ts'ebeliso ea morao-rao ea tšimo ena (Lichtman le Sanes, 2008). Mokhoa ona o fana ka leseli mabapi le kholo ea boko ba bocha, leha e haelloa ke maikutlo a mang a hore `` khokahano e sebetsang '' e bonoa libakeng tse se nang likamano tsa 'nete tsa anatomikiMahe a linotsi et al., 2009; Koch et al., 2002). FMRI ea naha ea phomolo e ipapisitse le maikutlo a hore liphetoho tse kholo tsa maqhubu a maholo (<0.1 Hz) lia etsahala (Biswal et al., 2010). Mekhoa ea ho utloisisa khokahano e sebetsang e kenyelletsa peo e thehiloeng ho peo (moo qalo e khetholloang ka letsoho ho khetholla qalo), tlhahlobo ea likarolo tse ikemetseng (ICA), le tlhahlobo ea domain-frequency. Nts'etsopele e sebetsang ea lits'ebetso tse fapaneng tsa boko e kenyelletsa kopanyo ea likhokahano tse fokotsehang tsa nako e khuts'oane (ke hore, khethollo) le ho eketsa likhokahano tse telele (ke hore, kopanyo) (Fair et al., 2007; Stevens et al., 2009). Ka mantsoe a mang, nts'etsopele e tsoa sebakeng ka seng ho ea marang-rang a arolelanoang haholo ha libaka tse fapaneng li hokahana haholo (Fair et al., 2009). Khokahano ena ha e hokahane, empa libaka tse ikemetseng lia hokahana ebe li hokahana (Supekar et al., 2010).

Lipatlisiso tse sebetsang tsa ho hokahanya fMRI ea ho phomola li bonts'a hore "marang-rang a sa feleng" a teng bokong ha a sa sebetse ka botlalo tlhahisoleseling. Khokahano ea marang-rang e na le "posterior cingrate cortex", mPFC, "lobial temporal lobes" le "angular gyrus". Mehaho ena e bonts'a li-oscillations tse kopanetsoeng, tse tlase ha 0.1 Hz) ha motho a le sebakeng se khutsitseng, se phomotsang. Ha kelello e ntse e hokahana hanyane ka hanyane pakeng tsa bongoana le bocha (Fair et al., 2008), khokahano e eketsehang ka har'a marang-rang ea kamehla e etsahala nakong ea phetoho ena (lipakeng tsa lilemo tsa 9-12; Broyd et al., 2009). Khokahano ea marang-rang e kentsoe ts'ebetso ho bapala karolo ea bokhoni ba ho qapa, athe phokotso kahare ho marang-rang ea kamehla e amanang le schizophrenia le autism.

Likhokahano tse ling tse sebetsang, leha ho le joalo, ka sebele li teng bokong. Phuputsong e bapisang bacha ba lilemong tsa bocha (ho bolela lilemo tsa 12.5 ± 0.51 [SD] lilemo) le batho ba baholo ba banyenyane (22.2 ± 1.67 [SD] lilemo) ka lihlopha tse kopaneng tsa thobalano, li-network tsa 13 tse sebetsang hantle li ile tsa tsebahala (Jolles et al., 2010). Ho li-network tsena, tse robeli li bonts'itse ts'ebetso e matla lipakeng tsa libaka tsa cortical nakong ea bohlankana, tse peli ha li supe phapang ts'ebetsong, 'me tse tharo li ne li amana le mesebetsi ea mantlha ea pono kapa sensorimotor (ke hore, sensorimotor, pono ea pono, le marang-rang a phallo ea marang-rang) mme li bonts'itse ts'ebetso e nyane nakong ea bocha. ho feta ho ba motho e moholo lilemong. Ho tsebahala ka marang-rang ana, hona joale ho tla nolofalletsa lipatlisiso tsa nako e tlang hore na ke hobaneng li bontša liphetoho tse amanang le lilemo.

5. Ntlafatso ea ts'ebetso ea lipotoloho

Nakong ea bocha, liphetoho tse makatsang tsa boitšoaro li tlangoa le liphetoho tse amanang le botsofali kahare ho bokong. Tlhahlobo e batsi ea boitšoaro ba bocha e fumanoa kae kapa kae (Spear, 2000), empa re fana ka setšoantšo se akaretsang sa hore na liphetoho tse ikhethang tšebetsong ea tšebetso nakong ea bocha li ka hlalosa a mang a boits'oaro bona. Ka har'a sehlopha sa 'mino oa kaho ea bokong, tikoloho ka' ngoe e na le kemiso ea eona ea ntlafatso (Tau le Peterson, 2010). Ka kakaretso, libaka tsa cortical li holile kamora nako ho feta libaka tse tlasa bohlasoa, joalo ka ha ho boletsoe ka holimo. Ho lieha ha nts'etsopele kapa nts'etsopele ea kelello kahare ho methapo ea methapo ea methapo ea methapo e kanna ea qala kemiso e ts'oanang le liketsahalo tsa nts'etsopele tse fetolang boemo ba likelello tsa likarolo tse ngata tsa boko (Ernst le Fudge, 2009; Haber le Rauch, 2010). Ho latela pono ena, lithuto tse telele li tla ba molemo ho khethollang tatellano ea liphetoho tsa boko ba libaka ha liketsahalo tse fapaneng li sa etsahala (Gogtay et al., 2006; Sowell et al., 2004). Mohlala, Shaw le basebetsi mmoho (Shaw le al., 2007) li bonts'itse hore bana ba nang le ADHD ba amanang le lithaka tsa bona, ba ts'oaroa ke kholo ea bona. Ka lehlakoreng le leng, ho qala ha lefu la kelello la bongoana ho amahanngoa le ho itšoaea setaelehileng ho feta ho bonoang ho bana ba tloaelehileng (Rapoport et al., 1999). Boithuto bo joalo ka tsena bo bohlokoa molemong oa ho lekola nako ea koluoa, empa hape ka nako e ts'oanang bo totobatsa lifensetere tsa nts'etsopele tse ka bang ts'oarellang le litšusumetso tsa kantle.

Ho hlaha ha psychopathology nakong ea bongoana libakeng tse phahameng tsa moputso- le ts'ebetso e amanang le ts'ebetso ha se ntho e iketsahalletseng. Ha ho fanoa ka palo ea liphetoho tse makatsang tse etsahalang nakong ena, lits'ebetso tse tsamaeang li sa tsamaee hantle kapa tse ileng tsa fosahetse pejana bophelong 'me li sa fetoloe ke liphetoho tsena (Andersen, 2003; Andersen le Teicher, 2008; Weinberger, 1987; Laviola et al., 2003) e tla bonahala nakong ena. Bohlokoa ba ho hlasimolla le ho theola linako tse tebileng bo thehiloe ho utloisiseng litlamorao tse bohloko tsa litsamaiso tsa ntlafatso. Ntle le moo, mathata a mangata a na le motheo lits'ebetsong tsa methapo ea methapo ea methapo e sa sebetse. Ho pepeseha litsietsing ho sa le pele ho emela kotsi e kholo bakeng sa likotsi tse 'maloa. Mohlala, lithuto tsa lefu la ho kula li bontšitse hore ho pepesetsoa litsietsing ho baka tlokotsi e kholo pherekanong e kholo ea khatello ea maikutlo (Anda et al., 2006; Anda et al., 2002; Chapman et al., 2004), botho bo moeling, tšebeliso e mpe ea lithethefatsi (Andersen le Teicher, 2009), le ho ipolaea, ka khatello ea maikutlo e le mokhoa o tloaelehileng oa batho ba baholo oa tlhekefetso ea pele (Putnam, 2003; Zisook et al., 2007).

5.1. Ntlafatso ea ts'ebetso ea lipotoloho tse amehang

Nts'etsopele ea tšebetso ea li-circuits le lits'ebetso bokong e rarahane, e na le likarolo tse ngata tse tsamaeang ho e kopanya. E le mokhoa oa ho atamela lipotoloho tsa nts'etsopele, re fana ka litlhatlhobo tse latelang ka ha li amana le ho ama le ho putsa nakong ea bocha. Mekhoa ena ha e kenyeletse lithuto tse se nang palo le tsa bohlokoa tse hlahlobang phetoho ea boits'oaro le ea bongaka e etsahalang nakong ea bocha, empa e tsepamisitse maikutlo lithutong tse nang le likamano tsa neuroanatomical motso oa tsona.

Boholo ba boitšoaro ba motho le tšusumetso li hlaha ho litloaelano tse neng li kile tsa fumanoa lipakeng tsa tšusumetso e ntle kapa e nyarosang le maemo a etsahalang ho ona (Cardinal le al., 2002). Mekhatlo ena e matla, e ithutileng e khanna boitšoaro ba rona ba hona joale le ba nako e tlang (Cardinal le al., 2002) mme e etsahala ka mekhoa ea boemo ba Pavlovian (Rosenkranz et al., 2003). Tlhahisoleseling mabapi le tikoloho le maikutlo li sebetsoa ka har'a basolateral amygdala (BLA) (Grace le Rosenkranz, 2002), e bopang likamano tse matla pakeng tsa sepheo se rerang ho hlaha hoa takatso kapa sepheo se setle, hape e hlahisang "ama" ka har'a BLA (Cardinal le al., 2002; Laviolette et al., 2005; Schoenbaum, 2004; Sheba le al., 2003). Le ha ho le joalo, ho arabela mohopolong o fanoeng ho hloka ho hlaka le ho nepahala ho latela maikutlo, bohlokoa ba maikutlo, kapa tlhokomelo joalo ka ha ho amana le khetho (Paus et al., 1996). Ts'ebetso ena e hlaha ho PFC (Cardinal le al., 2003; Rebec le Letsatsi, 2005; Schoenbaum, 2004; Ventura et al., 2007). Li-receptor tsa Noradrenergic le dopaminergic ho PFC li buella taolo ea tlhokomelo, boits'oaro le maikutlo ka ho matlafatsa khokahano ea marang-rang pakeng tsa li-neurons tse nang le lipallo tse arolelanoang (Arnsten, 2009). Ka har'a mPFC, leseli la tlhahisoleseling le sebetsoa ho laola tlhokomelo e khethiloeng.

Kahoo, tlhaiso-leseling e tsoang BLA e fetisetsoa ho mPFC ke likhakanyo tsa glutamatergic (Bechara et al., 1999; Laviolette et al., 2005; McDonald le Pearson, 1989), moo e sebetsoang bakeng sa masose (Schultz, 1998) le liphoso tse lokelang ho bolela sephetho sa nako e tlang (Falkenstein et al., 2000; Theko, 1999). Ka lebaka leo, susumetso ea ho bolela sephetho se nyarosang e ka arabeloa ka mokhoa o nepahetseng (Pezze et al., 2003). Ts'ebetso ena e etsoa ke lipontšo tsa dopaminergic ho mPFC (Jackson le Moghaddam, 2004), e kenyeletsang tlhaiso-leseling e eketsehileng ea boits'oaro le bocha ka leseli la maikutlo (Cardinal le al., 2002; Milad le Quirk, 2002) ho susumetsa boits'oaro bo tataisoang ke sepheo. MPFC e romella tlhahisoleseling ena ho li-nucleus tse bokelloang ka kotloloho (Goto le Mohau, 2005; Voorn et al., 2004), kapa ka tsela e sa tobang ka amygdala. Kamora nako, ts'ebetso ea sephetho ka har'a mPFC, ka kotloloho kapa ka kotloloho e susumetsa boits'oaro bo susumetsoang ho li-nucleus accumbens.

Ho sebetsanoa le ts'ebetso e sa lekanyetsoang pakeng tsa amygdala le PFC ho khothalelitsoe ho etsa ts'ebetso ea lietsahala ea ho kula ho hlonephehang ho fihlela lilemong tsa bocha (Ernst et al., 2006). Kahare ho setšoantšo sa mofuta oa tlholo se hlahisitsoeng ke Ernst le basebetsi mmoho (Ernst et al., 2006), mokhoa oa ho qoba o amanang le amygdala drives boits'oaro bo sa sebetseng ke PFC e sa lemalang. Ho latela mofuta ona, li-nucleus li bokella matla a khokahano lipakeng tsa takatso ea lijo le maemo a leholimo (1)Horvitz, 2002). Khopolo ena ke e 'ngoe ea tse' maloa tse sa tloaelehang tse kenyelletsang se tsejoang ka neurobiology ea khatello ea maikutlo ka har'a moralo oa nts'etsopele. Le ha ho le joalo, mohopolo o bolela hore bana le bacha ba tla hola ka lebaka la khatello ea maikutlo le khokahano ea cortical e hlahang, ho seng joalo (Andersen le Teicher, 2004; 2008).

Re sa tsoa shebisisa liphetoho tsa nts'etsopele nakong ea bongoana tse ka eketsang tlokotsi ea khatello ea maikutlo (Andersen le Teicher, 2008). Ka bokhutšoanyane, bana ba na le ts'ebetso e fetang ea batho ba baholo ba amygdala ho arabela maikutlo a bona (Killgore le al., 2001), e eketsang le ho feta ho bana le bacha ba nang le bothata ba ho tšoenyeha ka litaba tsa sechaba (Beesdo et al., 2009). Le ha ho le joalo, li-nucleus accumbens li kenella haholo ts'ebetsong ea takatso ea takatso ea lijo le bohloeki bocheng ho fapana le ho ea ho amygala (Ernst et al., 2005). Ho amoheloa ha PFC ho ea ka maikutlo a ferekanyang maikutlo ha ho etsahale ho fihlela motho e moholo.Killgore le al., 2001). Pele ho moo, sena se lumellana le liteko tsa tlhahlobo ea pampitšana tse bontšang tsoelo-pele e tsoelang pele ea BLA ho ea sebakeng sa bolulo sa PFC nakong ea bocha (Cunningham et al., 2002), empa ho bohlokoa le ho feta ke hore, tlhōrō ea ho boloka PFC ho kenya litlatsetso tsa BLA nakong ea bocha (Cressman et al., 2010). Ka kopanelo, likhokahano tse eketsehileng tsa anatomical li ka fana ka motheo oa ho hlaha kapa ho lieha ha bokhachane ba matšoao a sithabetsang le ho thatafala maikutlong ho emisang naha ena ea maturational e le taolo ea taolo hodima ama e fetoha (kapa e hloleha ho nts'etsapele).

5.2. Ntlafatso ea ts'ebetso ea lipotoloho tsa meputso

Lipatlisiso tsa MRI tse sa tsoakoang le lithuto tsa electrophysiologic li bonts'a likarolo tse ikhethileng tsa likarolo tse kahare ho cortex ea pele ho ts'ebetsong ea moputso. MPFC (libaka tsa Broadman [BA] 10 / 12 / 32 hape e kenyelletsa anterior cingrate cortex; BA 24) e arabela sephethong sa moputso: e entsoe ts'ebetso haeba moputso o lebelletsoeng o amoheloa mme o sa sebetsoa ha o sa amoheloe (Knutson et al., 2003; Schulz et al., 2004). Khoutu ea orbital frontal cortex (OFC) e lebelletse sephetho le likhakanyo tsa boleng bo susumetsoang ho latela moputso o ka bang teng. OFC e bapala karolo ea bohlokoa ho khutlisetseng ho ithuta le ho liehisa ho matlafatsoa (Dalley et al., 2004) ka ho hokahana ha eona le libaka tsa sensory, limbic, frontal, and subcortical. OFC e arotsoe ka tsela e hlophisehileng le likarolo tsa bongaka tse arabelang ka mokhoa o ikhethileng ho fumana moputso, athe likarolo tsa morao-rao li hatella mekhoa e amanang le moputso e fetileng (Elliott et al., 2000; Elliott et al., 2003; London et al., 2000).

The accumbens (sebaka sa khubung ea naha) e arabela bohlasoa (Ernst et al., 2004), valence (ho rata lijo kapa ho hlapatsa) (Jensen et al., 2003) le tebello ea moputso (moputso o sa lebelloang o sebetsa ka matla ho feta moputso o boletsoeng esale pele)Berns et al., 2001; Elliott et al., 2000)), empa eseng motsoako oa koloi (Zink le al., 2004). Nakong ea bocha, liqoqibete li arabela tse kholo ho feta tsa OFC ho li putsa (Galvan et al., 2005). Ha li kopantsoe hammoho, lintlha tsena li fana ka maikutlo a hore mocha o bokella mokhanni phetoho ts'ebetsong ea moputso (Galvan, 2010).

Leha ho le joalo, bopaki ba hore na litsamaiso tsa cortical le subcortical li arabela joang moputsong oa moputso li bontša hore cortex e bapala karolo e kholo le ho feta phetolelong ea bocha ts'ebetsong ea moputso. Lithuto tsa liphoofolo li bontšitse hore moputso oa phetisetso ea moputso nakong ea bocha ka ho pongoa le ho shebana hape le likhokahano tsa cortical ha marang-rang a hola le ho ba motho e moholo ((Brenhouse et al., 2008; Crews et al., 2007). Thupelo ea bongaka ea fMRI ea bongaka e fana ka maikutlo a hore tšebetso ea "ventral striatum" le "MPFC" e hlohlellehe ho fumana moputso nakong ea bocha.Bjork et al., 2004). Pele ho phetoho ena, mesebetsi e amanang le BOLD e hlahisa pherekano e eketsehileng le ts'ebetso e tlase ea libaka tsa pele ho bana ho feta ho batho ba baholo.Durston et al., 2003). Leha ho le joalo, bana ba bonts'a ts'ebetso e kholo ho "ventral striatum"Ernst et al., 2005; Galvan et al., 2006). Joalokaha re tseba ka mokhoa o fokolang mabapi le kholo ea moputso bathong, re tla ts'ohla lipatlisiso tsa tlhaiso-leseling bakeng sa kutloisiso e kholo.

Ho fetoloa ha mPFC ho liehisoa ho latela likarolo tse ling tsa boko (Andersen et al., 2000; Huttenlocher, 1979) hape e fihla sehlohlolong sa khokahano ea "synaptic" haufi le ho ba motho e moholo.Benes et al., 2000). Ho eketseha ha leqhubu la dopamine neurons (Benes et al., 1996; Kalsbeek et al., 1988; Verney et al., 1982), letsoalo la receptor (Andersen et al., 2000; Leslie et al., 1991), le tšebetso ea tsamaiso ea man systemosa a bobeli (Andersen, 2002) fihla qetellong ea dopaminergic drive ho mPFC nakong ea bohlankana. Liphumano tsa morao-rao li boetse li bonts'a keketseho e amanang le lilemo tsa ts'ebetso ea D1 ea lisele tse sa potlakileng tsa spiking ho mPFC, e etsahalang kamora ho kena bohlankaneng (Tseng et al., 2006), le tlhōrō ea palo ea ho thunya ea li-neuron tsa VTA dopaminergic ka nako e tšoanang (McCutcheon le Marinelli, 2009). Phatlalatso e fetelletseng ea li-receptor tsa D1 litlamong tsa glutamatergic ho li-accumbens le tsona li eketseha nakong ea bocha e tsamaellanang le boitšoaro ba ho batla lithethefatsi (Badanich et al., 2006; Brenhouse et al., 2008). Baahi ba li-receptor bona ba bile le tšusumetso ho khutlisetsong ea lithethefatsi, ka hona ho tebisa maikutlo ha bona nakong ea bocha ho bohlokoa (Kalivas, 2005). Liphetoho tsena tšebetsong ea moputso oa cortical le tsona li ka susumetsa likarabo tsa subcortical ho psychostimulants.

Ka lehlakoreng le leng, litekanyetso tsa basal tsa dopamine tsa extracellular dopamine le likarabo tsa dopaminergic ho li-stimulants ha li fetohe ka mokhoa o ananelang pakeng tsa bacha le batho ba baholo ho li-accumbens (Frantz et al., 2007) kapa mPFC (Jezierski et al., 2007). Leha ho le joalo, karo-karolelano lipakeng tsa cortical: expressionbens expression ea mofuta oa pele oa pele c-fos karabelo ea lintho tse matlafatsang li eketseha pakeng tsa bocha le ho ba motho e moholo.Andersen et al., 2001). Ntle le moo, amphetamine e hlahisa subcortical> cortical activation patterns tsa c-fos ho bacha (Andersen et al., 2001), Empa cortical> subcortical activation ho bacha (Cao et al., 2007). Ha li kopantsoe, lintlha tsena li fana ka maikutlo a hore bacha ba fapane haholo le bacha, ba baholo ka ho fetesisa likarabo tsa bona ho tse matlafatsang. Ka mantsoe a mang, menyetla ea hore tšebeliso ea lithethefatsi e hlaha haholo nakong ea bohlankana e latela litlamorao tse tobileng kapa tse sa tobang tsa ts'ebetso ea cortical ka ts'ebetso ea subcortical.

5.3. Ntlafatso ea ts'ebetso ea kelello

Litekanyetso tsa liteko tse kang Stroop, Simon, Flanker, Go / No-Go, le Mesebetsi ea Stop-Signal li hloka khatello ea boits'oaro bo itlhommeng pele ho etsa e sa iketsahalletseng feela. Taolo e lebisang tlhokomelo, thibelo ea karabo, le likhang le tlhaiso ea liphoso ke lits'ebetso tse hlakileng tse sebetsanang le ts'ebeletso ea taolo ea kelello le ts'ebetso e atlehileng ea mosebetsi. Ts'ebetso ho mesebetsi ena kaofela e ntlafala hanyane ka hanyane ho hola, empa ha e atamele maemo a batho ba baholo ho fihlela bonyane bongoaneng kapa bonyenyaneng ba pele (Bunge et al., 2002; Casey et al., 1997; Davidson et al., 2006; Luna le Sweeney, 2004; Rubia et al., 2000). Joalo ka mohopolo oa ho sebetsa, matla a ho itaola a bana a ka sithabetsoa habonolo ke litlhoko tse eketsehileng tsa mesebetsi. Ho batho ba baholo, ho itaola ho itšetleha ka libaka tse pharaletseng tsa cortical tse kang sebaka sa tlatsetso sa makoloi, libaka tse ka pele tsa mahlo, cortex cortex, dorsolateral PFC, ventralPFC / lateral orbitofrontal cortex, hammoho le libaka tsa nakoana le tsa parietal tseo kaofela li nang le khokahano le striatum ho subcortex (Leung et al., 2000; Marsh et al., 2007).

Ho arabela ka mokhoa o atlehang ho tsusumetso ea tikoloho ho hloka tlhokomelo e ikhethileng le tataiso e susumetsang, hammoho le khatello ea liketso tse seng li sa hlokahale kapa tse sa lokelang. Tlhatlhobo ena e lekantsoe ka liteko ka karabo ea karabelo, e kenyeletsang lits'ebetso tse tharo tse hokahaneng, joalo ka ha ho hlahisitsoe ke Barkely (Barkley, 1997): 1) thibelo ea karabelo ea pele e nang le bokhoni, 2) ho emisa karabo e sa khaotseng kapa ho lieha ho araba, le 3) ho fokotsa ho kena-kenana kapa ho senyeha nakong ea ho lieha. The basal ganglia le PFC ka bobeli li na le moelelo lits'ebetsong tsena (Casey et al., 2008). Ka kakaretso, ha basal ganglia e laola tšitiso ea boitšoaro bo sa lokang (Mink, 1996), PFC e sebetsa ho thibela ho kena-kenana le tlhaiso-leseling e nepahetseng ka tlhaiso-leseling (Miller le Cohen, 2001).

Ho fapana le thibelo ea ts'ebetso, e hlokang tšusumetso ea litšusumetso tse matla mme e hokahanngoe haholo ka tšebelisano-mmoho ea PFC, striatum, le amygdala (e lekantsoe (Ernst le Fudge, 2009)), Inhibition ea karabelo e hira li-circuits tse tataisang ho rala le ho etsa nako (Deiber et al., 1999). Boikarabello ba mantlha ba marang-rang a boits'oaro bo botle bo fana ka ts'ebeliso e fapaneng ea ntlafatso ho feta ea ts'usumetso le lits'ebetso tsa tlhokomelo e ikhethileng.

5.4. Ntshetsopele ya thibelo ya karabo

Le ha bacha ba khona ho etsa mesebetsi e metle ea boiphihlelo, bokhoni ba ho etsa joalo bo lula bo ntlafala nakong ea bocha le ho ba motho e moholo, ntlafatso ena ea methapo e hola hore tsoelopele ea methapo ea kutlo e latela tsoelo-pele e tšoanang. Bana ba bonts'a matla a maholo a ts'ebetso ho feta batho ba baholo ba libakeng tsa pele tsa lobe (Bunge et al., 2002) ho kenyelletsa ligrisi tsa karolo ea pele tsa medial frontal le likarolo tsa moea tsa karolo e ikhethileng ea li-front front gyrus (Booth et al., 2003). Sena se tsamaisana le phapang e amanang le lilemo tsa ho nepahala le nako ea ho etsa lintho tse etsahalang bongoaneng. Ho khahlisang, phuputso e kopaneng ea DTI le fMRI e entsoeng ke Stevens le basebetsi mmoho (Stevens et al., 2009) e tlaleha kamano e ikhethileng lipakeng tsa liphetoho tse amanang le lilemo tsa khokahano e sebetsang pakeng tsa patellar ea naha e kopaneng, kotara e nepahetseng ea parietal le caudate e nepahetseng, ho eketseha ha myelination, le ntlafatso ea ts'ebetso ea Go / No Go. Phuputsong e 'ngoe ea DTI, karabo ea karabo ho ba nang le selemo sa 7-13 e ne e amahanngoa haholo le FA e phahameng le MD e tlase maemong ka bobeli a boemo bo tlase bo ka pele le bo nepahetseng ba pre-supplement motor cortex (Madsen et al., 2010). The linear developmental trajectory ea myelination e tšohliloeng ka holimo ka hona e lumellana le tlhaiso e hlakileng ea tatellano ea taolo ea kelello, e amanang le mokhoa o ts'oaroang oa U oa sebopeho sa tšusumetso le ts'ebetso ea moputso. Bana le bona ba bonts'a matla a maholo a ts'ebetso ho feta batho ba baholo ka har'a lesapo la caudate le letšehali nakong ea ho tsamaea / ho se ee (Booth et al., 2003) le ho emisa (Rubia et al., 1999) mesebetsi. Sehlopha sa basal ganglia se hlahisitsoe hore se kenye letsoho ho thibela boitšoaro bo sa lokang (Casey et al., 2001), 'me sehlopha sa basal ganglia se bonahala se hola hanyane ka hanyane ho tloha bongoaneng ka ho ba motho e moholo.

Neurobiology ea mantlha ea lipotoloho tsena e se e kile ea buisanoa ka holimo kapa e sa ntsane e ithutoa maemong a ntlafatso. Ha ho ntse ho na le leruo la data la neuroimaging le potileng mesebetsi ea karabo, ho bile le lipatlisiso tse fokolang tsa ts'ebetso ea methapo ka morao ho lits'ebetso tsena (bakeng sa tlhahlobo e felletseng, bona Eagle et al., 2008). E 'ngoe ea mathata a mantlha a amanang le ho khetha mokhoa ona pele ho libeke tse hlokahalang ho ruta liphoofolo ho etsa mesebetsi ena, e emelang ho ithuta ha tsona nakong ea kholo. Ka lebaka la bohlokoa ba taolo ea kelello le taolo nakong ea bongoana ho ba motho e moholo, tšimo ena e hloka tlhokomelo e fetang eo e e fumaneng.

6. Boiphihlelo bo hlahisa kholo ea bokong

Le ha liphatsa tsa lefutso li fana ka polane ea ho aha boko, bona litšoantšo tse betliloeng bokong hore li tsamaisane le litlhoko tsa tikoloho. Phello ea qetello ea synapse e fanoeng e thehiloe ho netefatso ea ts'ebetso. Boko ba bongoana ha bo angoe feela ke tšusumetso ea tikoloho, empa bocha ke nako eo boiphihlelo ba pele bo hlahang (Andersen, 2003; Andersen le Teicher, 2008). Likhokahano tse rarahaneng tsa neural li ba teng nakong ea bohlankana, 'me tsena li phuthoa ke mesebetsi e etsoang ka boiketsetso le boiphihlelo (Ben-Ari, 2002; Francis et al., 2002; Katz le Shatz, 1996; Zhang le Poo, 2001). Tlhahlobo ea rona ea pejana (Andersen, 2003) e buile ka tšusumetso e kholo eo matla a tikoloho a nang le eona holima kholo ea boko. Lipampiri tse ling tsa tlhahlobo-kholo li bua ka tšusumetso eo khatello ea maikutlo e bang le eona holima kholo ea bongo ea bongo (Andersen le Teicher, 2008; 2009). Ho pepesehela lithethefatsi tsa psychotropic nakong ea ntšetso-pele le hona ho tla fetola boemo ba khatello ea maikutlo, le litlamorao tse hlahang nakong ea bocha.Brenhouse et al., 2009; Ansorge et al., 2008).

Summary

Sebopeho le boholo ba liphetoho tsa bocha mahlaleng a neuroanatomy ea boko li lula li fetoha ha lisebelisoa tsa rona tsa ho sekaseka li ntse li hlaka hantle. Phapang e ka ananeloa feela ha libaka li ithutoa kahare ho karohano ea tšebetso (mohlala (Gogtay et al., 2006)), ka nako e felletseng ea tšobotsi, le ha liphihlelo tsa pele (Andersen le Teicher, 2008) le lintlha tse ling (mohlala, botona le botšehali, sethala sa Tanner) li nkuoa ka hloko. Li-timec tsa nako e sa felletseng lithutong tsa pejana li lebisitse liqetong tse sa nepahalang mabapi le nako ea mofuthu (e tšohliloeng ka McCutcheon le Marinelli, 2009) le hore na liphihlelo tsa pele li hlile li ama ntlafatso. Tlhahlobo ena e fana ka kakaretso ea kutloisiso ea rona ea hajoale ea liphetoho tsa bohlankana bokong nakong ea phetoho ea hae ho tloha bongoaneng ho ea ho motho e moholo. Ts'ebetso ena e makatsang e ts'oarella haholo ka lebaka la polasetiki e lumellang sisteme ea mammona ho ikamahanya le litlhoko tsa tikoloho ea eona.

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Setšoantšo sa 1 

Lenane la lits'ebetso tsa nts'etsopele ho batho le litoeba. Mekoallo ea pinki e emela tatellano ea linako ea basali, e tlang pele ho ea banna, e emeloang ka mehala ea boputsoa. Polelo ea nakoana ea li-receptors ("ectopic") e etsahala bongoaneng mme ...

Lintlha-khōlō Tsa

  • Re lekola bocha e le sethala sa nts'etsopele ea bohlokoa.
  • Liphetoho tsa Neuroanatomical li hlophisitsoe ka tšusumetso ea tikoloho le litlhoko tsa lona.
  • Li-trajector tsa nts'etsopele li sebelisana le liphetoho tse itšetlehileng ka thobalano.
  • Re tšohla kholo ea kutloisiso ka lisebelisoa tse ntseng li eketseha tse tebileng.

Mongolo o botlaaseng ba leqephe

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

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