Atụmatụ mmepụta ihe n'oge uto na ụmụ nwoke na nwanyị: nghọta nke obe na-agbanwe (2011)

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PMCID: PMC3134153
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Mbubre ọhụụ nke onye bipụtara edemede a dị na Neurosci Biobehav Rev
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nkịtị

Oge uto bụ oge mgbanwe n'etiti nwata na ntozu nke na-agụnye nnukwu mgbanwe n'ime usoro ụbụrụ nke yiri ụfọdụ, ma ọ bụghị ihe niile, mgbanwe omume. Ngosi elu na mmeghachi omume mmetụta uche na nhazi ụgwọ ọrụ na-eso ụdị ntụgharị U n'usoro mmalite na mgbaghara, yana ọnụ ọgụgụ kasị elu na-eme n'oge uto. Otú ọ dị, nhazi ọgụgụ isi na-agbaso usoro mmepe nke linear karịa. Nyocha a ga-elekwasị anya na mgbanwe n'ime akụkụ ndị bụ isi ma gosipụta mmekọrịta dị n'etiti mgbanwe ụbụrụ na omume, yana ihe akaebe sitere na imaging resonance magnetik arụ ọrụ (fMRI) n'ime ụmụ mmadụ na ọmụmụ ihe omimi nke ndị na-anabata ihe na ihe mgbaàmà na anụmanụ. A ga-eji mgbanwe ndị nọ n'afọ iri na ụma mee ihe iji ghọta ka omume na-adịghị ahụkebe na-esi ebili n'oge uto. Anyị na-ebute ọmụmụ ihe gbasara ụlọ ọgwụ na nke izizi iji nye usoro akwara maka ịkọwa oge uto na ọrụ ọ na-eme na ngbanwe ruo n'ịbụ okenye.

Keywords: Oge uto, ihe isi awọ, ịkwachaa, esemokwu mmekọahụ, ihe ọcha

Okwu Mmalite

Oge uto bụ oge pụrụ iche na mmepe ụbụrụ mammalian. A kọwawo nghọta n'oge uto n'ọtụtụ nyocha na ọkwa omume (McCutcheon na Marinelli, 2009; Okwu, 2000; Steinberg, 2010; Laviola et al., 1999; Laviola et al., 2003) na usoro usoro (Ernst na Fudge, 2009), ma ọ bụ naanị na-atụle n'ókè dị oke na ọkwa nke mgbanwe neuronal (Andersen, 2003; McCutcheon na Marinelli, 2009; O'Donnell, 2010; Okwu, 2000). Anyị ga-enyocha neuroanatomy, njikọta ọrụ, mkpụrụ ndụ ihe nketa, na mgbanwe akara na-eme n'oge uto. N'ikpeazụ, usoro dị n'ime usoro akwara ozi ga-emepụta ka mgbanwe ndị ntorobịa na akara ndị a si emetụta omume.

1. Isi

1.1 Ịkọwa oge uto

Enwere ike ịkọwa oge uto dị ka oge dị n'agbata afọ 10-19 n'ime mmadụ (WHO, 2010s), n'etiti afọ abụọ na anọ na primates (Schwandt et al., 2007), na n'etiti 35-60 ụbọchị na-eto eto na òké (Andersen et al., 2000; McCutcheon na Marinelli, 2009). Okwu (2000) malite mkparịta ụka ya banyere oge a site n'oge uto nke akọwara dị ka oge mgbanwe omume. A na-ahụkarị mgbanwe omume omume dị otú ahụ n'ofe ụdị anụ ahụ dị iche iche site n'inwewanye mmetụta maka ndị ọgbọ na ihe gbasara ọha mmadụ (Blakemore, akwụkwọ akụkọ a; Forbes na Dahl, 2005; Steinberg, 2010; Panksepp, 1981), ize ndụ (Laviola et al., 2003), na njikwa ọgụgụ isi na-eto eto (Casey et al., 2008). Nkọwa nke oge uto nwekwara ike ịdabere n'ụzọ ezi uche dị na ya na mgbanwe gonadal dịka ọ dị mkpa na ntozu oke mmekọahụ (Egwu na Foster, 2004). Arụmụka ndị e depụtara n'ebe a adịghị agwụ agwụ ma e kwesịghị iji ya mee ihe n'ụzọ doro anya, kama dị ka ebe ntụnyere aka.

Ọkwa mmepe ọhụrụ, ntolite na-etolite, na-apụta n'etiti afọ 18-29 n'ime mmadụ.Arnett, 2000). Akọwapụtara n'ọdịnala, ntozu ntozu n'ime mmadụ na-akọwa nleba anya na ọ bụ ezie na ọtụtụ mgbanwe neurobiological metụtara oge uto agwụla, akụkụ ahụ 'etozubeghị' dịka ọ na-egosi site na igbu oge n'inweta ọrụ ma ọ bụ alụmdi na nwunye. N'akụkọ ihe mere eme, G. Stanley Hall (1904) kọwara oge 'ọhụrụ' ntozu okè nke kọwara ntolite site n'echiche akụ na ụba nke ọha na eze nke mere ka e nwekwuo nnabata nke ọkwa dị iche. N'ihi ya, anyị achọpụtala mgbanwe neurobiological pụrụ iche na nke dị mkpa nke na-akọwa oge uto. Ọ bụ ezie na nyocha a lekwasịrị anya n'ụzọ bụ isi na akara nrịanrịa neurobiological nke oge uto, ọ dị mkpa ịghọta na n'ụdị òké na oge dị adị nke nwere ike weghara okenye na-etolite (obere ozi dị na primates na-abụghị mmadụ). Dị ka a tụlere n'okpuru, oke na-egosi mgbanwe dị ịrịba ama n'etiti 40-60 ụbọchị, ma oge n'etiti 60-100 ụbọchị na-ejikọta ya na nwayọọ nwayọọ na-agbanwe agbanwe nke na-eji nwayọọ nwayọọ na-edozi. Nke a nwere ike ịbụ oge "onye toro eto" ọhụrụ nke kwesịrị nlebara anya nyocha, ọ bụghị ihe ngosi mgbasa ozi iji kọwaa mgbanwe omenala ọhụrụ na mba ndị mepere emepe? Ihe dị mkpa nke ịkọwapụta ọkwa bụ ịbịarute nkwenye nke ọnọdụ ntozu okè nke organism nke a kọwara iji kwado ụdị obe na ntụnyere mmekọahụ.

1.2 Gịnị mere e ji nwee oge mgbanwe dị otú ahụ?

Site n'echiche evolushọn, e mebere omume site na nhọrọ eke iji kwado onye ọ bụla ka ọ nwee ihe ịga nke ọma n'ime ọha mmadụ na nke anụ ahụ dịka onye toro eto, gụnyere ịchọta onye ọlụlụ na ịmụ nwa. Usoro a na-ejedebe n'oge uto. N'omume, ụmụ anụmanụ na-esi na òké gaa na mmadụ niile na-enweta oge mgbanwe mgbanwe ebe ịnyagharị site na ntolite na mbelata mmetụta nne na nna jikọtara ya na mmetụta ndị ọgbọ na-abawanye, asọmpi mmekọahụ, na ihe ịma aka ime mkpebi ọhụrụ (nke nyochara ya. Okwu, 2000). Neuroplasticity na-enye ohere maka nzaghachi kwesịrị ekwesị na gburugburu ebe a na-apụta na nke a pụtara ìhè na mmepe nke ụgwọ ọrụ na usoro metụtara mmetụta (Galvan, 2010). Agbanyeghị, usoro mmepe ndị ọzọ na-egosipụta mmụba na-aga n'ihu na njikwa ọgụgụ isi n'oge uto nke na-akwado ime mkpebi (Geier na Luna, 2009; Somerville na Casey, 2010). Ọnụ, Yin na Yang a na-adabere na mmepe mmepe, ebe ihe ka ọtụtụ n'ime ndị na-eto eto na-agbasi mbọ ike na mgbanwe nke ndị ọgbọ na ndị nne na nna na-apụta dị ka ndị nwere onwe ha, ndị na-achịkwa onwe ha ka usoro ndị a na-enweta nguzozi. Mgbe mgbanwe ndị a na-etolite nke ọma, a na-eme mgbanwe n'otu n'otu na ike gburugburu ebe obibi na ọha mmadụ pụrụ iche. Agbanyeghị, njehie na usoro a na-ebute omume adịghị mma. Mwepụta nke psychopathology nwere ike ịpụta n'ụzọ ụfọdụ site na mgbagha site na nhụsianya nkịtị nke maturation, na-ebute okwu ogologo ndụ na nhazi ụgwọ ọrụ na mmetụta mmetụta. Ewezuga ihe na-adịghị mma nke mkpụrụ ndụ ihe nketa na-akpata, njehie na mmepụta nke oke na ịkwachaa neurons ma ọ bụ ndị na-anabata ya, nchacha nke ọma nke eriri eriri ma ọ bụ ikpughe mkparị nke nwata nwere ike inye aka. Nlebanya a ga-elekwasị anya na usoro mmepe ndị a na ụbụrụ mammalian, na-emesi ike n'ozuzu ya na ụdị kama ọ bụghị ihe atụ (dịka ọmụmaatụ, Andersen na Teicher, 2008; 2009; Marco et al., 2011).

1.3 Ụdị mgbanwe ahụ

Usoro usoro akwara ozi na-enye nghọta maka mgbagwoju anya nke ọdịdị mmepe ntorobịa. Dị ka Paus kwuru (Paus et al., 2008), Usoro nke akụkụ dị iche iche nke ụbụrụ na-arụ ọrụ na-egosi n'ụzọ doro anya otú mpaghara mpaghara na ọrụ dị iche iche si eme ka ụbụrụ dị n'oge uto nwee ọtụtụ akụkụ. N'ime nyocha a, anyị na-enyocha ihe a maara banyere mgbanwe na ọnọdụ mmepe na-elekwasị anya na usoro ntorobịa dị ka akọwara n'ofe ụdị anụ mammali na n'etiti nwoke na nwanyị. Usoro anyị gbadoro ụkwụ n'otu akụkụ n'ụdị triadic, nke Ernst na ndị ọrụ ibe kọwapụtara (Ernst na Fudge, 2009; Ernst na Korelitz, 2009). Ihe atụ triadic na-agbanye mgbanwe omume n'ime usoro atọ bụ isi, ma ọ bụ ọnụ, ya bụ usoro mmetụta, usoro ụgwọ ọrụ, na mgbochi / nzaghachi nzaghachi. Ọnụ atọ ndị a dị iche iche na-arụkọ ọrụ ọnụ iji mepụta àgwà ndị na-egosipụta ntozu oke ntorobịa. Ọnụ ụzọ nke ọ bụla nwere usoro mmepe nke ya, nke na-emepụta usoro ntorobịa na ọnọdụ mgbanwe. Nsonaazụ omume ikpeazụ nwere ike ịdabere na ọnụ ọnụ nke ọkwa enyere ma ọ bụ nwere ike pụta site na ọnụ adịghị ike nke na-adịghị arụ ọrụ nhazi. Ụdị triadic n'ụdị ya dị mfe na-enye ịkọwa mmekọ mmekọ nke nwata na ọtụtụ mkpali mmetụta uche, mgbanwe na mmetụta ụgwọ ọrụ, yana mgbanwe akara na njikwa cortical na mmepe ọgụgụ isi. N'ebe a, anyị ga-eji usoro a kọwaa mgbanwe zuru ezu na mmepe ntorobịa n'ofe ụdị na mmekọahụ na-elekwasị anya na mpaghara ụbụrụ cortical na limbic.

2. Ime nke trajectory: Neuroanatomical mgbanwe

N'ọkwa neuronal, usoro nke mmepe ụbụrụ nke ndị nọ n'afọ iri na ụma bụ otu n'ime nhazi nke synapti. Neurons na-edobere na mbụ n'ụdị ime ime nke innervation na cortex (Rakic ​​et al., 1986). Neurons ndị a mụrụ na mbụ na-abanye n'ígwé dị omimi nke cortex, ebe ntinye nke elu nke cortex dị elu na-eme ma emechaa na mmepe. Mkpụrụ ndụ glia abụọ na-eduzi ebumnuche Neuronal.Rakic ​​et al., 1986; Vernadakis, 1975) na gradients kemịkalụ nke a na-ekpebi site na nkwupụta neurotransmitter (Landis na Keefe, 1983; Purves na Lichtman, 1980). Okwu Neurotransmitter nwere ike ịdị na-adịgide adịgide, na-ebute ntinye n'ime mpaghara enyere, ma ọ bụ ectopic, na ndị na-ebugharị na-egosipụta nwa oge maka ebumnuche mmepe nke nduzi. A na-etolite synapses ka neurons na-abata na mpaghara ha lekwasịrị anya. A na-enyocha mgbagwoju anya nke usoro ọmụmụ na mmalite ọmụmụ nke usoro a n'ime omimi dị ukwuu n'ebe ọzọ (Levitt, 2003; Tau na Peterson, 2010), na a gaghị atụle ya n'ụzọ zuru ezu ebe a. Ka ntorobịa na-eru nso, synapses na-emepụta nke ukwuu ma emesịa na-efunahụ ya, nke a na-akpọ kwachaa. Ịkwachaa bụ usoro na-abụghị otu ihe dị ka apoptosis na ọnwụ cell, ebe ọ bụ na ịkwachaa bụ nhazi nke alaka dendritic na njikọ synapti na apoptosis na-eme atụmatụ ọnwụ cell. Ikwachaa nke synapses bụ ihe a ma ama na ụbụrụ ndị nọ n'afọ iri na ụma n'ofe ụdị dị iche iche ma enwere ike ịgụta ya na nyocha nke ọnwụ (post-mortem analysis).Andersen na Teicher, 2004; Huttenlocher, 1979; Lewis, 1997) ma ọ bụ sitere na MRI, ebe mgbanwe mpaghara na isi awọ na ihe ọcha na-egosi n'oge uto na ngwa ngwa ka ụmụ mmadụ na-abịaru nso afọ iri nke atọ nke ndụ ha.Giedd et al., 1999a; Huttenlocher, 1979; Sowell et al., 2004). Ọ bụ ezie na ekwenyeghị na ịkwachaa synaptik kwa se ọ na-emetụta oke nyocha nke olu.Rakic ​​et al., 1986; tụlere site Giedd et al., 2008), mgbanwe na mpịakọta isi awọ na ọcha nwere ike gosipụta mgbanwe nke akụkụ synapti n'elu mmepe.

2.1. Àgwà ndị e ji emebiga ihe ókè na ịkwachaa

2.1.1. Synaptogenesis na kwachaa

A na-echekwa usoro nke synaptogenesis na ịkwachaa nke ukwuu n'ofe ụdị anụ mammali. Ọmụmụ ihe ọmụmụ mmadụ nke izizi nwụrụ anwụ site n'aka Huttenlocher (Huttenlocher, 1979; Huttenlocher na de Courten, 1987) na Benes (Benes et al., 1987) bụ ndị mbụ gosipụtara mgbanwe dị egwu n'ime ihe isi awọ na ọcha n'oge oge ntorobịa. Kpọmkwem, ịkwachaa n'ime oyi akwa 3 nke cortex ihu mmadụ dị ezigbo mkpa na ihe dịka 40% nke synapses na-efunahụ n'etiti afọ 7 na 15. Dịka ọmụmaatụ, akara synapti nke synaptophysin n'ime mmadụ na-ebili nwayọọ nwayọọ n'etiti ọmụmụ na afọ ise, rute n'ala dị larịị na afọ 5, ma daa n'ogo okenye site na afọ 10 na dorsolateral prefrontal cortex (PFC) (Glantz et al., 2007). Nyocha zuru ezu nke synaptogenesis na rhesus monkey moto cortex na-ekpughe ụkpụrụ yiri nke ahụ na mmepụta synapti na-aga n'ihu ma na-enweta ọkwa synapse nke dị okpukpu abụọ karịa nke ndị okenye. Ọnụ ọgụgụ nke synaptogenesis na-ebelata ka enwe na-eru ntozu okè mmekọahụ (afọ 3), wee gbadaa ngwa ngwa ruo ọkwa okenye (afọ XNUMX).Zecevic et al., 1989). N'otu aka ahụ, ụkpụrụ njupụta synapti nke oke na-ebili n'etiti ụbọchị 25 na 40, ma kwụsie ike ma emesịa (Andersen na Teicher, 2004). Agbanyeghị, ọ bụghị mgbanwe niile metụtara afọ na olu bụ n'ihi ịkwachaa synapti (dịka ọmụmaatụ, mgbagha dendritic). Ụzọ agụta ọnụ ọgụgụ sel n'ụzọ ziri ezi na oke na-ekpughe mfu nke neurons nke afọ na cortex a na-ahụ anya na ọkwa niile (ma e wezụga IV) na oke mgbe ọ dị afọ iri na ụma (Yates na Juraska, 2008). A na-ahụkwa ọdịiche mpaghara na mfu cell, dị ka njupụta synapti. Ọ bụ ezie na cortex a na-ahụ anya na-egosi mfu 18-20% na sel, a na-ahụ obere mfu 5% na ventromedial, mana ọ bụghị n'akụkụ azụ, PFC na oke (Markham et al., 2007). Ọ bụ ezie na oke mmepụta na ịkwachaa dịgasị iche n'etiti mpaghara na n'ime mpaghara (n'etiti ọkwa dị iche iche), a na-ahụ usoro ahụ n'ofe ụdị dị iche iche na-emekarị.

Ikwachaa na-emekarị na synapses asymmetric dị na spines dendritic, dịka egosiri na cortex moto (Zecevic et al., 1989), oyi akwa molekul nke hippocampal denate gyrus na dorsolateral PFC (Eckenhoff na Rakic, 1991; Onye na-azụ atụrụ, 1990). Synapses asymmetric bụ isi ihe na-akpali akpali na okike, ebe synapses symmetric na-egbochikarị. Njupụta nke neurons GABA (onye na-ebunye ihe mgbochi mbụ) ka kwụsiri ike karịa afọ (Brenhouse et al., 2008; Vincent et al., 1995), nke na-eme ka ọnụ ọgụgụ ndị mmadụ kwụsiri ike nke synapses symmetric na ogwe dendritic (Zecevic et al., 1989). Aghọtachaghị usoro nke ịkwachaa. Otú ọ dị, nyocha ndị na-adịbeghị anya achọpụtawo akụkụ ụfọdụ usoro usoro mkpụrụ ndụ ihe nketa nke ịkwachaa synapses excitatory. Mbelata nke ndị nọ n'afọ iri na ụma NRG1, mkpụrụ ndụ ihe nketa na-etinye aka na nrịbama neuregulin, nwere ike ịrụ ọrụ na nguzozi mkpali / inhibitory na nhọrọ synaptic (Harris et al., 2009). Complexins, nke bụ protein presynaptic nke na-achịkwa ntọhapụ neurotransmitter ma jikọta ya na mgbagwoju SNARE, na-agbanwekwa na afọ. Complexin 2 (CX2), ihe nrịbama nke synapses excitatory, na-egosiputa usoro mmepe nke curvilinear na ala dị larịị site na afọ 10 n'ime mmadụ. N'ụzọ dị iche, complexin 1 (CX1) njupụta, nke jikọtara ya na synapses inhibitory, ji nwayọọ nwayọọ na-ebili site na nwata na PFC dorsolateral mmadụ.Salimi et al., 2008).

Ọ bụ ezie na synapses glutamatergic na-agbanwe n'oge uto, GABA na-egosipụtakwa mgbanwe dị ukwuu metụtara afọ nke na-ekwu. Mgbanwe GABA ndị a na-arụ ọrụ na okike, ebe mgbanwe glutamatergic bụ nhazi. Na mbụ, GABA nwere omume mkpali n'isi mmalite mmalite ọmụmụ. GABA na-enweta mmetụta mgbochi ya site na mmepe ọwa chloride nke na-agbanwe n'ime izu nke abụọ nke ndụ na oke; GABA na-akwado ihe mgbochi a site n'ịbụ okenye (Ben-Ari, 2002). A na-emepụta mgbanwe a na-akpali akpali-inhibitory site na nnukwu oscillations na ọkwa calcium n'oge mmepe, nke na-eme ka mmepe synaptic dịkwuo mma (Ben-Ari, 2002). Mgbochi nwa ọhụrụ nke usoro na-ahụ maka ọrụ chloride n'oge dị elu (dịka ọmụmaatụ, Na (+) -K (+) -2Cl (-) cotransporter [NKCC1]) na-emepụta mgbanwe na-adịgide adịgide na cortical circuitry n'oge okenye.Wang na Kriegstein, 2010). Ya mere, mgbanwe dị ịrịba ama na ọrụ neuronal n'oge mgbanwe a nwere ike ịkpụgharị sekit na-eto eto na-adịgide adịgide.

GABA neurons na-arụ ọrụ dị mkpa n'imekọrịta ọrụ cortical site na mkparịta ụka dị mgbagwoju anya nke ntinye aka na usoro nzaghachi nke na-achịkwa mgbasa ozi spatiotemporal nke dị n'etiti ọnụ ọgụgụ nke pyramidal neurons.Constantinidis et al., 2002; Di Cristo et al., 2007). Omume mgbochi ndị a nke GABA tozuru oke na mmepe nke nhazi nghọta dị mgbagwoju anya (Luna et al., 2010) na mmụba nke ukwuu n'oge uto na ụmụ mmadụ (Lewis et al., 2004), ndị na-abụghị mmadụ (Cruz et al., 2003; Erickson et al., 1998) ma ọ bụ na oke (Tseng na O'Donnell, 2007). A na-achọta GABA n'ụzọ bụ isi n'ime mmadụ atọ dị iche iche na-egosipụta protein protein parvalbumin, calbindin, na calretinin. Enwere ike iji immunohistochemistry nke protein ndị a dị iche iche iji soro mmepe GABA. Dịka ọmụmaatụ, neurons parvalbumin-immunoreactive na onye na-ebugharị akpụkpọ anụ GABA (GAT1) na ịrị elu nke na-abụghị nke mmadụ, na-arị elu n'oge ndụ wee nọgide na-ebuli elu ruo ọnwa 15, wee kwachaa n'oge uto na ọkwa okenye (Anderson et al., 1995; Conde et al., 1996; Cruz et al., 2003). Na mgbakwunye, protein ndị na-akọwapụta ntinye GABA na neurons cortical pyramidal (dịka ọmụmaatụ, akụkụ gephryin nke akpọrọ axon nke mbụ) prune n'oge uto.Cruz et al., 2009). GABA na-emekọrịta ozi cell pyramidal site n'ịgbanwe ọsọ nke ntinye dị iche iche na mpaghara cortical (ọtụtụ glutamatergic). Usoro a na-egosipụta nke ọma site n'ichepụta nghọta dị elu nke na-agụnye echiche nkịtị n'oge mgbanwe n'etiti oge uto na okenye. N'ịchịkọta ọnụ, ụbụrụ na-etobeghị bụ nke a na-akpụzi nke ọma site na nhazi mkpali na GABA na-enye aka na usoro a na mmalite nke ndụ tupu ọ ghọọ ihe mgbochi n'oge uto.

2.1.2 Heterosynchrony na kwachaa

Heterosynchrony na mmepe ụbụrụ na-ezo aka na ọdịiche mpaghara na oge ịkwachaa n'oge mmepe. A na-ahụta oke mmepụta na ịkwachaa n'oge na-adịbeghị anya site na ọmụmụ ihe nhazi nhazi (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 na Peterson, 2010). Mgbanwe ụda isi awọ dị ka achọpụtara ya na MRI na-atụ aro usoro mmepụta ihe na-esote na ịkwachaa ya na maturation. Mgbanwe ndị a na-egosipụtakarị mgbanwe synaptik, n'ihi na ndị a bụ isi ihe na-enweghị myelinated nke neuron. Usoro MRI na-enye ohere maka nyocha ogologo oge nke ọtụtụ mpaghara ụbụrụ n'ime otu isiokwu, nke na-agaghị ekwe omume na ụzọ ndị ọzọ. Ihe ọmụmụ ogologo oge dị otú ahụ enyela maapụ doro anya nke ihe heterosynchrony dị ka ihe nkiri na-ewe oge (http://www.loni.ucla.edu/~thompson/DEVEL/dynamic.html). N'ime cortex, usoro nke ịkwachaa a na-eme n'azụ azụ n'ihu, ya na ihe ndị mbụ na-emepe emepe nke sensorimotor cortex kwachaa na mbụ, mgbe ahụ, njikọ cortices na-ebute ụzọ okporo ụzọ na-emepe emepe n'oge.Paus et al., 2008). Ọmụmụ ihe gbasara ọnwụ na-egosi na ịkwachaa n'ime ọkwa dị iche iche nke visual, somatosensory, moto, na mpaghara prefrontal, agbanyeghị, na-eme n'otu oge (Rakic ​​et al., 1986).

A, mpaghara subcortical na-etolite tupu mpaghara cortical (Tau na Peterson, 2010). Amygdala nwere ike ịbụ otu n'ime mpaghara mbụ na-etolite ma na-etolite n'ụdị dimorphic mmekọahụ. N'ime ụmụ agbọghọ, amygdala na-egosi obere mgbanwe na ụda isi awọ n'oge uto, ebe ọ na-erute oke ya site na afọ 4; N'ime ụmụ nwoke, amygdala olu na-eji nke nta nke nta na-abawanye ruo afọ 18 site na 53%. Mpaghara ndị ọzọ, gụnyere caudate, putamen, na cerebellum na-egosi udi ntụgharị-U na olu isi awọ nke na-arị elu n'oge uto yana mpịakọta na-agbada site na 15% (nyochaa (nyochaa)Durston et al., 2001)). Nkebi nke ihe owuwu enyere ekpugherekwa mgbanwe ndị metụtara afọ ndị pụtara ìhè (Gogtay et al., 2006). Nnyocha mmalite nke hippocampus na MRI gosipụtara mmụba dị ntakịrị na olu (12%) n'ofe afọ. Nyochagharị data a ka afọ iri gachara na-ekpughe mgbanwe dị egwu n'ime mpaghara mpaghara. Dịka ọmụmaatụ, akụkụ azụ nke hippocampus na-apụta na ọ na-amịpụta ma kwachaa ihe isi awọ karịa akụkụ nke mbụ (Gogtay et al., 2006; Insausti et al., 2010).

Ọdịiche mpaghara dịka ndị a na-egosi oge dị iche iche nke adịghị ike maka mkparị nwere ike ịdị adị nke a na-enwebeghị ekele zuru oke n'ihi nleba anya nke mpaghara ụbụrụ nyere (Andersen, 2003; 2005; Andersen na Teicher, 2008). Nnyocha banyere mmetụta nke ikpughe na nhụjuanya n'oge nwata na-egosi mbelata 12-15% n'ozuzu oke okwu isi awọ hippocampal n'ime mmadụ (dịka, Bremner et al 1997), na nke ọma, nyocha ndị a lekwasịrị anya na akụkụ ndị a dị n'azụ nke na-enweta ihe kachasị mma. mgbanwe mmepe. Heterosynchrony na mmepe n'ime ọtụtụ ọkwa nyocha (dịka, mpaghara, mpaghara, na ọkwa) kwesịrị iburu n'uche mgbe ị na-amụ mmepe nkịtị ma ọ bụ mmepe mgbanwe na-eso mkparị.

Ọ bụ ezie na MRI bara uru maka nyochaa mgbanwe na ihe isi awọ n'ofe ụbụrụ dum, usoro a na-enye obere nghọta nke mgbanwe mgbanwe na-eme n'ime usoro neurotransmitter dị iche iche. Ntụnye ihe isi awọ na-egosipụta atụmatụ nhụsianya nke njupụta synapti nke na-egosighi mgbanwe ọrụ na-apụta ìhè n'oge mmepe, dị ka ndị a tụlere n'elu. Otú ọ dị, nyocha nke okwu mkpụrụ ndụ ihe nketa n'oge uto na anụ ahụ post-mortem nke mmadụ (ya bụ, ụzọ mgbochi na-agaghị ekwe omume na MRI) nwere ike inye nkọwa ndị ọzọ gbasara ọdịdị mgbanwe na-eme n'oge a. Mkpụrụ ndụ ihe nketa metụtara usoro mmepe neuronal, gụnyere ntụzịaka axon, morphogenesis na synaptogenesis, na-ebelata n'oge uto na oke.Harris et al., 2009). Ihe atụ akọwapụtara gụnyere netrins, semaphorins, neuropilin, neurexin na neurolignin. Mgbanwe ndị metụtara afọ na neurexin kwekọrọ na nkwụghachi azụ axon nke na-akọwapụta ịkwachaa na mbelata dị ukwuu na mkpụrụ ndụ ihe nketa hụrụ n'etiti ụbọchị 45 na 90 na oke.Cressman et al., 2010). Nyocha ụyọkọ nke mkpụrụ ndụ ihe nketa na microarray nwere ike ime ka a ghọtakwuo mkpụrụ ndụ ihe nketa ọhụrụ ndị na-etinye aka n'ịmebiga ihe ókè na ịkwachaa. N'ime nyocha dị otú ahụ, mkpụrụ ndụ ihe nketa gbakọtara n'ime ụyọkọ ọrụ atọ bụ isi: ụyọkọ cytoskeletal (25 chọpụtara), ụyọkọ metụtara Ras/GTP (12 chọpụtara), na metabolism lipid na usoro usoro steroid (13 chọpụtara). Ụyọkọ cytoskeletal na-emezigharị ọkwa nhazigharị ahụ nke na-eme n'oge uto, ụyọkọ Ras/GTP na-atụkwa anya mgbanwe ọrụ, ebe ụyọkọ nke atọ na-egosipụtakarị myelination na mgbanwe ndị metụtara oge uto. N'ikpeazụ, ọnụ ọgụgụ kasị elu nke ndị nọ n'afọ iri na ụma na protein neural cell adhesion molecule (NCAMs) na-egosi na a na-egosipụta mkpụrụ ndụ ihe nketa ndị a na-arụ ọrụ n'otu n'otu na nchọpụta òké.Cox et al., 2009).

Ọ bụghị mgbanwe niile n'okwu mkpụrụ ndụ ihe nketa na-emetụta protein nhazi. Dịka ọmụmaatụ, mkpụrụ ndụ ihe nketa metụtara ndị na-anabata glucocorticoid na-agbanwe n'oge uto (Perlman et al., 2007; Prince, 2008). N'ime mmadụ na primates na-abụghị mmadụ, ndị na-anabata glucocorticoid na-abawanye na elu n'oge uto. Otú ọ dị, isoforms na ndị na-anabata glucocorticoid (GR) na-egosi ụzọ dị iche iche, na GR isoforms GRalpha-A na 67-kDa GRalpha na-eto eto na-eto eto na ọzọ n'oge uto; N'ụzọ dị iche, ụdị GRalpha-D na-arị elu n'isi mmalite ma na-ebelata ma emesịa (Sinclair et al., 2010). A na-egosipụta protein ndị GR ndị a nke ọma na neurons pyramidal, mana na-egosipụta okwu na-adịru nwa oge na astrocytes ọcha n'oge ọhụrụ.

Na nyocha pụrụ iche nke mkpụrụ ndụ ihe nketa 2,979 nke nwere ike ịkọwa heterochrony (ya bụ, a na-egosipụta mkpụrụ ndụ ihe nketa ndị a n'ụzọ dị iche iche n'etiti mpaghara, na nke a, PFC dorsolateral na caudate nucleus na ụmụ mmadụ), 58% nke mkpụrụ ndụ ihe nketa na-eme ka uto dị nwayọọ n'etiti mpaghara cortical na subcortical (Somel et al., 2009). A na-enyochakwa mkpụrụ ndụ ihe nketa maka ọdịiche dị n'etiti mmadụ na chimpanzees gbasara mmepe heterochrony na mgbe ọmụmụ nwa gasịrị. Chimpanzees na-eso ụmụ mmadụ na-ekekọrịta nnukwu ihe ọmụmụ, mana ha nwere ogologo ndụ dị mkpụmkpụ, nke na-enye ụzọ ọzọ iji ghọta heterochrony. N'iji atụnyere nke a, ụdị mkpụrụ ndụ ihe nketa yiri nke ahụ na-esi n'etiti ụdị dị iche iche na mmalite nke ntozu oke mmekọahụ (Somel et al., 2009), na mgbanwe ndị metụtara mmepe isi awọ.

2.1.3. Ndabere mmekọahụ

MRI morphology ọmụmụ n'ime ụmụ mmadụ na-egosi na ụmụ nwoke nwere 9% ibu ụbụrụ ụbụrụ karịa ụmụ nwanyị, yana ọdịiche mmekọahụ ndị ọzọ na-ahụ anya na akụkụ nke subcortical.Giedd et al., 1996a). Nucleus caudate buru ibu na ụmụ nwanyị, mana a na-ahụ ọdịiche ndị ọzọ na ọnụego mmụba nke nha. Ogo amygdala na-abawanye ngwa ngwa n'ime ụmụ nwoke karịa ụmụ nwanyị, nke a na-ahụ maka nha hippocampal. Nwoke caudate na-agbada na nha, ebe nwanyị caudate nke nwanyị anaghị agbanwe nke ọma n'ofe afọ (Giedd et al., 1996a). Caviness et al (Caviness et al., 1996) mere nyocha nke MRI volumetric nke gosipụtara na subcortical forebrain nuceli (neostriatum) na ụmụ nwanyị nọ n'ogo okenye n'etiti afọ 7-11. N'ụzọ dị iche, otu ihe ahụ n'ime ụmụ nwoke nke otu afọ dị ukwuu karịa ụda ha toro eto, na site na ntinye ga-alaghachi azụ tupu ha etoo. Site na ntozu oke na oke, ụmụ nwoke toro eto nwere 18% ventral medial PFC (mPFC) karịa ụmụ nwanyị, nke a na-esite na neurons ole na ole (13% metụtara ụmụ nwoke) na sel glia (18%) (Markham et al., 2007). A kọwawo mgbanwe ndị yiri ya na cortex visual isi nke oke, ebe ụmụ nwoke nwere ~ 20% karịa ụda isi awọ n'ihi akụkụ nke 19% karịa neurons karịa ụmụ nwanyị (Nunez et al., 2002; Reid na Juraska, 1992).

Kedu ka ndịiche nhazi ihe ndị a si emetụta ọrụ bụ ntule. Ekwenyere na ịkwachaa n'onwe ya na-eme ka nhazi dị mma (Changeaux et al., 1976; Purves na Lichtman, 1980). Ozugbo netwọk neuronal guzobere n'ụbụrụ na-eto eto, redundancy n'ime netwọk adịghị arụ ọrụ ma na-akwacha synapses. Dị ka a tụlere n'elu, a na-ekwenye na mbelata nke njupụta synaptik na ọnụ ọgụgụ cell na-eme ka arụmọrụ nke nhazi dịkwuo elu. Mgbanwe nhazi ndị a na-agbakwunyere site na mbelata nke itinye glucose n'ọrụ (ihe na-egosi ọrụ ụbụrụ; a tụlere n'okpuru ebe a na Nkebi 4.1), nke dị elu na nwata na oge uto tupu ịkwachaa. Mmetụta nke usoro a na-apụta ìhè karịsịa mgbe ọ na-adịghị mma. Nwoke caudate na-akwachaa nke jikọtara ya na nnukwu ihe ize ndụ maka omume na nsogbu metụtara moto, gụnyere Tourette's Syndrome na nlebara anya enweghị ike hyperactivity.Teicher na Andersen 1995). Mpaghara jikọtara ya na omume nwere ike bụrụ nke na-eto eto; mpaghara ndị ọzọ metụtara mkpakọrịta ọhụrụ na ebe nchekwa a na-emelite mgbe niile nwere ike ọgaghị akwachaa otu oke (Teicher et al. 1995). Neuron ole na ole na mpaghara ọ bụla, gụnyere mPFC, nwere ike ịbawanye arụmọrụ na ọsọ nhazi.

Enwere ike ịhazi esemokwu mmekọahụ na mmalite nke ndụ site na homonụ gonadal nke na-akpụ ụbụrụ akabeghị aka (nyochaa nso nso a na Viveros et al, 2010). N'ime oge nwa ọhụrụ, ntụgharị nke androgens na estrogen site na neural aromatase na-enye aka na mmetụta nke gonadal steroid na ọrụ ụbụrụ, gụnyere ọdịiche mmekọahụ site na "ịma nwoke" ụbụrụ nwanyị (MacLusky et al., 1994). A na-ahụ mmalite mmalite nke saịtị na-ejikọta androgen na-ejikọta ya na enzymes metabolic n'oge mmepe mmalite na hypothalamus, amygdala, dorsolateral na orbital PFC na somatosensory cortex (na primate na-abụghị mmadụ: Clark et al 1989; oke: Reid na Juraska, 1992)). Aromatization nke testosterone na ụbụrụ na-eme ka ọ dịkwuo mgbagwoju anya iji chọpụta nke hormone mmekọahụ bụ maka esemokwu mmekọahụ. Nnwale ndị na-eji androgen na-abụghị aromatizable, 5a-dihydrotestosterone (DHT), na-enyere aka ịkọwa mmetụta ndị a na-akpata steroid, ma ojiji ndị dị otú ahụ na-ejedebe na ọmụmụ nke ụdị ala ma ọ bụ chromosomal abnormalities.

N'ime nnwale eke nke gụnyere chromosomal abnormality XXY (dịka ọmụmaatụ, Klinefelters), ndị a ebelatala isi awọ n'ime insula, gyri temporal, amygdala, hippocampus, na mpaghara cingulate.Giedd et al., 1996a). Nkọwapụta n'oge na-adịbeghị anya na ụmụ mmadụ na-akọ na mkpokọta ihe isi awọ jikọtara ya na ọkwa estradiol na ụmụ agbọghọ (r= -0.32) yana nke ọma na ọkwa testosterone n'ime ụmụ nwoke (r= 0.32).Peper et al., 2009). Esemokwu mpaghara maka mmetụta hormonal, Otú ọ dị, dị, dị ka mmekọrịta siri ike n'etiti gyrus ihu dị ala na ọkwa estrogen na ụmụ agbọghọ (r=-0.72). Na mgbakwunye, njikwa androgens n'oge ndụ ha nwere nsonaazụ na-arụ ọrụ na ọrụ cortical. Dịka ọmụmaatụ, ịkpa oke ihe, ọrụ metụtara PFC, ka mma n'ime ụmụ nwoke na-eto eto na ụmụ nwanyị na-ekpughere androgen n'ihe gbasara ụmụ nwanyị nkịtị (Clark na Goldman-Rakic, 1989). N'ụzọ dị iche, mmụba n'oge uto na ọkwa homonụ mmekọahụ na-ebelata mgbochi pre-pulse, nke enwere ike ịgbazite ya site na nsonaazụ nhazi na ọrụ dopamine subcortical.Morris et al., 2010).

Ọmụmụ rodent na-atụ aro na estrogen ọhụrụ na-egbochi mmepụta neuronal na nwanyị ventromedial PFC (gụnyere mpaghara prelimbic na infralimbic)Juraska na Markham, 2004; Markham et al., 2007), nke dị iche na akụkọ ndị gara aga banyere ikike estrogen nwere ike kpalite nnukwu arborization na mpaghara ụbụrụ ndị ọzọ dị ka hippocampus na ndị okenye (Hajszan et al., 2009; Toran-Allerand, 1996). Prepubertal ovariectomy na-ebelata njupụta neuronal n'ime ụmụ nwanyị, nke nwere ike ịkọwa oke isi awọ n'ime ụmụ nwanyị.Nunez et al., 2002). Mmụba nke testosterone n'oge uto na-enyere aka kwachaa dendrites n'ime amygdala nwoke na-eto eto (Zehr et al., 2006). N'ịkọkọ, ọmụmụ ihe ndị a na-atụ aro ka homonụ gonadal na-ekere òkè dị mgbagwoju anya n'ịkpụ ụbụrụ ụbụrụ na-eto eto.

2.2. Mmepụta na ịkwachaa sistemu nnabata

2.2.1 Mmepụta nke ndị na-anabata monoamine

Mmepụta na ịkwachaa nke sistemu nnabata dị mgbagwoju anya ma e jiri ya tụnyere mgbanwe synapti, na ebili mmiri abụọ nke mgbanwe afọ na njupụta na-eme. Ọtụtụ sistemụ neurotransmitter, gụnyere dopamine (Gelbard et al., 1990; Kalsbeek et al., 1988; Lankford et al., 1988; Todd, 1992), norepinephrine (Feeney na Westerberg, 1990; Kline et al., 1994na serotonin (Kuppermann na Kasamatsu, 1984; Lauder na Krebs, 1978; Whitaker-Azmitia na Azmitia, 1986) nwere oke trophic ọrụ na ụbụrụ. A na-ejikọta okwu ectopic nke subtypes dị iche iche nke nnabata n'oge mmalite mmalite ọmụmụ ọmụmụ na mmụba nke omepụta synapti, uto axonal, na nhazi synapse. Dịka ọmụmaatụ, okwu ectopic nke ndị na-anabata serotonin 5-HT7 n'ime hippocampus na-eme obere oge n'ime izu abụọ mbụ nke ndụ na oke.Louiset et al., 2006; Vizuete et al., 1997). N'otu aka ahụ, a na-ahụ onye na-ebugharị serotonin (5-HTT) na neurons na-abụghị serotonergic embrayo na cortical na striatal neuroepithelia na sensory thalamic ụzọ postnatally na P0-P10 (Zhou et al., 2000). A na-ahụkwa ngosipụta nke 5-HTT na vesicular monoamine transporter (VMAT) na akwara cranial sensọ, na hippocampus, cortex cerebral, septum, na amygdala.Lebrand et al., 1998). Ekwenyere na ndị na-ebugharị na/ma ọ bụ ndị na-anabata ya na-eduzi innervation neuronal. Mmetụta nke trophic neurotransmitters na-adabere na ntinye uche (Mazer et al., 1997), na-atụ aro na ọkwa ntọala dị oke mkpa maka ọdịdị mmetụta. A na-ahụkwa okwu nnabata ectopic yiri ya na ihe ọcha. Dịka ọmụmaatụ, a na-ahụ α2 onye na-anabata noradrenergic n'ihe ọcha na-etobeghị na oke (oke).Happe et al., 2004). Agbanyeghị, ọ bụghị okwu nnabata niile na-arụ ọrụ trophic.

Ebili nke abụọ nke nnabata ihe karịrị nkwupụta na-eme n'oge uto, n'oge ndị nnabata na usoro akara ngosi na-egosi ngbanwe nke ọdịdị U-nke na-ebute ọkwa ngosipụta nke na-adịgide ruo n'oge. N'adịghị ka ectopic, okwu na-adịru nwa oge nke fọrọ nke nta ka ọ bụrụ okenye, ọnụ ọgụgụ ndị a na-anabata ya ji nwayọọ nwayọọ na-ebili, na-arị elu ma na-adalata n'oge ntozu okè. A na-ahụ ntụle mgbanwe nke nnabata ndị ntorobịa na Isiokwu 1, na-emesi ike na ndị nabatara n'ime mpaghara limbic na cortical. Ogologo oge mmepụta na ịkwachaa na-adabere na mpaghara (mpaghara).Andersen et al., 2000), a na-ahụkwa ya n'ọtụtụ nrịbama. Usoro nnabata dị iche iche gụnyere: dopamine, serotonin, norepinephrine, glutamate, GABA, neurotensin, endocannabinoid, na cholinergic (Andersen et al., 2000; Eggan et al., 2010; Lidow et al., 1991). Na enwe rhesus, Lidow et al (Lidow et al., 1991) egosila na njupụta nke ndị na-anabata ya na-etolite na concert na synaptogenesis.

Ọ bụrụ na anyị lekwasịrị anya n'ihu na microcircuits iji nyochaa nkesa nke ndị nnata metụtara afọ, nsonaazụ na-adịbeghị anya na-atụ aro ọbụna mgbanwe dị mgbagwoju anya n'oge uto. Nkesa nnata n'onwe ya na-agbanwe n'etiti phenotypes neuronal dị iche iche. Dịka ọmụmaatụ, ndị na-anabata dopamine D1 adịghị ka ọ na-agbanwe ọkwa ha nke ọma n'etiti afọ ịgbapụ ara ruo n'ịbụ okenye na neurons GABAergic (Brenhouse et al., 2008; Vincent et al., 1995). N'ụzọ dị iche, mmepụta na ịkwachaa nke ndị na-anabata D1 na-eme nke ọma na neurons mmepụta glutamatergic (Brenhouse et al., 2008). Kpọmkwem, naanị 2% nke amụma glutamatergic ndị a bụ D1 immunoreactive na oke ụmụ oke, na-ebili ruo 44% na P40, wee daa na 6% na ntozu oke na P100. Ọ dị mkpa ka enyocha ma ndị nnabata ndị ọzọ gosipụtara ngosipụta dị iche na subtypes neuronal ndị ọzọ n'oge uto. Isiokwu 1 na-enye ozi na klaasị nnabata ndị ọzọ na-agbanwe, mana amataghị ụdị neuronal akọwapụtara. N'ụzọ dị iche, ndị na-anabata D2 na-egbochi ọrụ nke GABA interneurons na-agba ọsọ ngwa ngwa mgbe ha tolitere (O'Donnell, 2010; Tseng na O'Donnell, 2007). Neuron ndị a dị mkpa maka ijikọta ọtụtụ ntinye nke ọma na ozugbo. Ya mere, nkesa nkesa n'ime microcircuits na ike ọrụ ha na-agbanwe nke ukwuu n'oge uto.

2.2.2 Ndabere mmekọahụ

Ihe akaebe izizi maka ọdịiche nwoke na nwanyị na nkwupụta nnabata sitere na ọmụmụ PET mmadụ ebe DA na 5HT njupụta nnabata na-agbada karịa n'ime ụmụ nwoke karịa ụmụ nwanyị sitere na 19-30 afọ.Wong et al., 1984). Anyị egosipụtakwala ọdịiche dị n'ihe gbasara nwoke na nwanyị na striatum n'oge ntorobịa nke ntorobịa, yana ụmụ nwanyị na-egosipụta obere nnabata nnabata na obere kwachaa (Andersen et al., 1997). Dịka ọmụmaatụ, njupụta nke ndị na-anabata D2 mụbara 144 ± 26% n'ime ụmụ nwoke karịa 31 ± 7% n'ime ụmụ nwanyị n'etiti ụbọchị 25 na 40 na oke. N'otu aka ahụ, ịkwachaa ndị nnata dị ukwuu karịa ụmụ nwoke ma mee n'etiti ụbọchị 40 na 120 (okenye). D1 striatal density belatara 34 ± 4% n'ime ụmụ nwoke, mana naanị 7 ± 8% n'ime ụmụ nwanyị. Maka nucleus accumbens, nwoke na nwanyị D1 njupụta njupụta njupụta dị n'otu mgbe ụbọchị iri anọ gasịrị, nke ọ bụla na-egosipụta ntakịrị ntakịrị na ụbọchị 40. Agbanyeghị, ọdịiche mmekọahụ na njupụta nnabata D80 nọgidere na P1, ebe ndị na-anabata D120 bụ 1± 57.8% karịa ụmụ nwoke karịa ụmụ nwanyị. N'ozuzu, enweghị ọdịiche nwoke na nwanyị na njupụta D21.2 na nucleus accumens. Otú ọ dị, ọdịiche dị n'etiti nwoke na nwanyị na-enwe mmekọahụ, enweghị ike ime mgbanwe nke hormone gonadal n'oge oge ntorobịa (Andersen et al., 2002). Gonadectomy ozugbo tupu D1 na D2 nnabata nnabata emeghị ka njupụta zuru oke n'oge uto; ọbụghịkwa gonadectomy na mbụ na ndụ. Nsonaazụ ndị a na-egosi na mkpughe nke testosterone na-adịghị akpali mmepụta nke dopamine, ma ọ bụ estrogen anaghị egbochi mmepụta oke n'ozuzu ya. Mmachi nke nyocha ahụ nwere ike igbochi ịhụ mgbanwe ndị dabere na mmekọahụ. Ebe autoradiography dabara nke ọma maka ịkọwa mgbanwe njupụta nnabata nke mpaghara n'ozuzu ya, usoro a anaghị ekpughe ọnụ ọgụgụ nke neurons na-egosipụta ndị nnabata a. Ya mere, ihe ga-ekwe omume na-anọgide na mgbanwe ndị na-adabere na mmekọahụ, na nkwụsị nke hormonal ha, na-eme na ọnụ ọgụgụ dị iche iche nke neurons nke a na-amabeghị.

Nyocha a agaghị elekwasị anya na nsonaazụ arụ ọrụ nke mgbanwe ndị nnata a, dị ka ndị na-enyocha nzaghachi nke ndị agonists ma ọ bụ ndị na-emegide ya. Otú ọ dị, ọ dị mkpa iburu n'obi na esemokwu mmekọahụ na usoro mgbaàmà na-emetụta hormones gonadal, ma na-enwekwa mgbanwe mmepe n'oge uto.Andersen et al., 2002; Kuhn et al., 2001).

3. Njikọ

3.1. Innervation pụrụ iche nke sistemụ neurotransmitter

N'akụkụ a, anyị na-atụle ka usoro neurotransmitter kpọmkwem si abanye mpaghara ụbụrụ nyere. Innervation na-amalite tupu ịmụ nwa, mana ọ na-aga n'ihu ruo n'oge ntorobịa na ntozu okè. Otú ọ dị, ọtụtụ ọmụmụ ihe na-agafe n'ịkọwa oge uto ma na-eche na innervation na-aga n'ihu n'ụdị ahịrị. Ọmụmụ ihe gbasara njikọta ọnwụ mmadụ na-agaghị ekwe omume ime, ebe ọ bụ na ụlọ ọrụ akụrụngwa ụbụrụ na-ekesa anụ ahụ ụbụrụ n'ime obere mpaghara na-egbochi ịchụ traktị. Mkpebi nke MRI anaghị ekwe ka nchọta traktị ọnụ ọgụgụ neuronal kpọmkwem ịkparịta ụka n'etiti onwe (ọ bụghị site na traktị, nke na-enyocha ma myelin na axon caliber n'otu oge). A na-ejikarị njupụta nke ndị njem dị ka ihe ngosipụta nke ụkpụrụ innervation (dịka ọmụmaatụ, (Moll et al., 2000)). Agbanyeghị, njupụta nke ụgbọ njem nwere ike ịdịgasị iche na-adabereghị na innervation ma yabụ na ọ gaghị adabara nke ọma maka ebumnuche ndị ahụ.

Dabere na ọmụmụ anụmanụ ole na ole nke na-eji ụzọ nchọta ọkọlọtọ akọwa ntozu ntozu, ụfọdụ na-egosi ọganihu n'ahịrị nke innervation n'ofe ntozu okè (dịka, (Brenhouse et al., 2008; Brummelte na Teuchert-Noodt, 2006; Cunningham et al., 2002; Erickson et al., 2000), ebe ndị ọzọ (Cressman et al., 2010; Rios na Villalobos, 2004) gosi ụkpụrụ U-atụgharị. Anyị ahụla ọganihu ahịrị ahịrị nke innervation nke oyi akwa V glutamate neurons nke medial PFC n'ime isi ihe na-akpata nucleus n'etiti 25, 44, na 100 ụbọchị na oke.Brenhouse et al., 2008). N'ime nyocha nke Cunningham na ndị ọrụ ibe ya (Cunningham et al., 2002), a hụkwara ụkpụrụ innervation linear na njikọ glutamatergic dị n'etiti amygdala na PFC, nke na-aga n'ihu site na ọmụmụ ruo n'oge uto / ntorobịa (ụbọchị 60) na oke. Esemokwu afọ na njikọ synapti bụkwa qualitative nke ọma. Dịka ọmụmaatụ, neurons glutamate kpụrụ axo-dendendritic (36.5%), axo-spinous (7.7%), na axo-somatic synapses (5.8%) na neurons GABAergic, mana 17.3%, 30.8% na 1.9% na akwara ndị na-abụghị GABAergic. Nhazi nke kọntaktị ndị a na-agbaso ụkpụrụ curvilinear n'ogologo afọ.

N'ụzọ dị iche, ụfọdụ ụkpụrụ nke innervation na-egosi usoro ọmụmụ na-abụghị nke linear na trajectory ha. Dịka ọmụmaatụ, PFC medial (ma mpaghara prelimbic na infralimbic) amụma na amygdala basolateral ka kwụsiri ike n'etiti ụbọchị 25 na 45 n'ime oke, mana belata ihe dịka 50% n'etiti 45 na 90 (XNUMX na XNUMX).Cressman et al., 2010). A na-ahụ nchọpụta ndị yiri ya na ụmụ oke. Afferents sitere na dorsalmedial thalamus ruo cortex n'ihu na-abawanye ruo ụbọchị 13, na-esote mbelata 67% n'ime izu nke atọ nke ndụ, mgbe ha ji nke nta nke nta na-abawanye ruo n'oge uto wee guzosie ike (Rios na Villalobos, 2004). Ejikọtara usoro izizi nke innervation nke mbụ na nhazi ọrụ nke neurons oyi akwa III, na-atụ aro na ntinye glutamate na-ebugharị synaptogenesis. Dopamine neurons na-agbaso ụkpụrụ atụ nke innervation na primate cortex (mpaghara 4, 9, 46): dopaminergic axon na oyi akwa III mụbara okpukpu atọ tupu ọnwa 5-7 dị afọ, na-enweghị mgbanwe dị mma na ọkwa 1 na V.Erickson et al., 1998). Ụdị varicosities akpọrọ aha na-aga n'ihu na-abawanye, na-eru n'ogo kachasị elu (n'ime okpukpu isii karịa nke obere enwe) n'ime anụmanụ 2-3 afọ (oge uto) tupu ọ daa na ọkwa okenye kwụsiri ike (na-eto eto).Rosenberg na Lewis, 1995; Woo et al., 1997). Gerbils na-egosipụta ụkpụrụ yiri nke ahụ. Dopamine innervation n'ime amygdala na-abawanye izu atọ mbụ na ndụ na gerbils, tupu ntakịrị mbelata nke njupụta n'oge ntorobịa nke na-eme ka ọ kwụsie ike ruo n'oge okenye.Brummelte na Teuchert-Noodt, 2006). Ya mere, o yikarịrị (ma ọ bụchaghị ekpuchiri ya nke ọma na nyocha a) na sistemụ neurotransmitter ndị ọzọ na-egosi mgbanwe yiri nke ahụ na usoro innervation.

N'oge a, amabeghị ihe kpatara usoro innervation dị iche iche (dịka, linear versos inverted U-shaped) ji eme n'ígwé cortical dị iche iche (Nyocha 2). Enwere ike nke mbụ dabere na nlele nke afọ, ebe nkwụsị dị egwu nwere ike ịdị na-akọwapụtaghị nke ọma. Nke abụọ ekwe omume dabeere na ọdịdị / ọrụ nke mpaghara na-innervated. Anyị ebulitela okwu a na mbụ n'ihe gbasara ndị na-anabata dopamine (Teicher et al., 1995) na ndị ọzọ maka innervation (Erickson et al., 1998). Kpọmkwem, mpaghara dị iche iche na-etinye aka na ọrụ ndị na-achọ mmelite mgbe niile nwere ike irite uru site na mmụba linear nke na-eme n'oge ndụ (tupu oge uto). N'ụzọ dị iche, mpaghara ndị na-etinye aka na mmụta nke ọrụ ogologo ndụ, dị ka àgwà, na-erite uru site na nhazigharị nke jikọtara ya na ịkwachaa. Ihe nke atọ nwere ike ime bụ na innervation na-egosi ụkpụrụ afọ kpọmkwem na nzukọ laminar, na oyi akwa III na cortex na-egosipụta ọdịdị U-atụgharị, na omimi na elu elu na-egosipụta usoro na-aga n'ihu. Ejikọtara ọnụ, njikọta pụrụ iche dị n'ime ime na nke dịpụrụ adịpụ na-enyere aka n'ịkpụgharị sekit neuronal n'oge uto (Benes, 2009).

Nyocha 2 

a) Eserese nke cortical lamination na vetikal cross-ngalaba nke Santiago Ramon y Cajal na-eso Nissl (aka ekpe, n'etiti) n'ime okenye yana ntụpọ Golgi (n'aka nri) n'ime nwa ọhụrụ dị 1½ ọnwa. B) ụkpụrụ nke mgbanwe synapti na-eme n'oge mgbanwe ...

3.2.1 Myelination

N'ime mmepe niile, ọtụtụ n'ime uru zuru oke na ụbụrụ ụbụrụ na-enweta site na myelination nke traktị eriri (fiber).Benes et al., 1994). Myelination na-abawanye ọsọ nke mgbanwe ozi, ma ọ dịkarịa ala na-ahụ maka mpụta nke omume omume mammalian bara ọgaranya (Ubi, 2005). Myelination na ụbụrụ mmadụ dị iche site na mmekọahụ na mpaghara (Benes et al., 1994; Giedd et al., 1999b). Myelination na-eji nke nta nke nta na-abawanye na ntozu oke na nwoke na nwanyị abụọ, dabere na ọmụmụ post-mortem (Benes et al., 1987) na MRI ọmụmụ nke na-enyocha mgbanwe ndị dị otú ahụ site n'ichepụta ihe ọcha na isi awọ (Paus et al., 1999) ma ọ bụ site na iji ihe onyonyo tensor diffusion (DTI) (Paus et al., 1999). Ihe ka ọtụtụ n'ime ihe a maara banyere mgbanwe mmepe na myelination dabeere na ọmụmụ nke corpus callosum, traktị myelin kasị ukwuu na ụbụrụ (dịka, (Keshavan et al., 2002; Teicher et al., 2004)). N'adịghị ka mgbanwe isi awọ, usoro rostral - caudal nke ihe ọcha na-aga n'ihu na-abawanye ogo corpus callosal n'ime oge ntorobịa (Giedd et al., 1996a). Mgbanwe ndị metụtara afọ na-eme na ngalaba azụ (Paus et al., 1999). Traktị ndị ọzọ na-acha ọcha, ya bụ n'ime capsule na aka ekpe arcuate fasciculus, na-aga n'ihu na myelinate na maturation. Myelination na-egbu oge nke njikọ frontocortical nke na-eme n'ime afọ iri nke abụọ na nke atọ n'ime mmadụ nwere ike jikọta ya na ụkpụrụ omume na nkwalite mkpali na-apụta mgbe ntorobịa gasịrị.Luna et al., 2010; Paus, 2005).

DTI na-eme atụmatụ na ntule nke mmegharị mmiri, site na nha nke pụtara diffusivity (MD) na fractional anisotropy (FA). N'ime voxel e nyere, usoro FA dị iche site na 0 (mgbasa isotropic zuru oke) ruo 1 (mgbasa anisotropic zuru oke), ma kpebisie ike site na dayameta eriri na njupụta, ịdị n'otu na ogo myelination.Basser na Pierpaoli, 1996). FA na-enyocha ogo ntụziaka nke mgbasa mmiri. Ntugharị mmiri n'otu ụzọ, dị ka ihe na-eme n'akụkụ traktị, nwere uru FA dị elu. Enwere ike ịhụ nkọwa zuru oke banyere otú MD na FA si agbanwe n'ofe afọ (5-30 afọ) na mpaghara ụbụrụ dị iche iche na akụkọ Lebel et al.Lebel et al., 2008na Qiu et al (Qiu et al., 2008). N'ime mpaghara ndị e ji mara na akwụkwọ akụkọ Liebel et al, ihe kachasị njọ nke MD na-eme na caudate nucleus n'oge uto ebe splenium nke corpus callosum na-erute n'uju ya (~ 8%) tupu 15 afọ. Agbanyeghị, nha FA na-egosipụta karịa myelination, ma gụnye atụmatụ nke ndịiche dị na ọdịdị nke traktị eriri n'onwe ha (dịka ọmụmaatụ, ntinye n'otu n'otu nke axon n'otu n'otu na ngwugwu “njupụta” ha; Paus, 2010). Ya mere, e mere atụmatụ mgbanwe na myelination dabere na FA jikoro kwesịrị iburu n'uche ma myelin na axon dayameta. Oke `g' (dayameta axon: dayameta axon + ọkpụrụkpụ n'ọbọ myelin) ka emepụtala maka dayameta axon na dayameta eriri. Ebe ọ bụ na dayameta axon na ọkpụrụkpụ myelin na-emetụta ọsọ nke iduzi mana ha anaghị abawanye n'otu ogo mgbe ha tolitere, oke 'g' nwere ike gosipụta mgbanwe mmepe na ihe ọcha na ihe omume (Paus na Toro, 2009). Ịtụle ogo nke myelination na mmekọrịta ya na dayameta axonal chọrọ microscopy eletrọn. N'ime oke, usoro stereological na-enweghị isi na-egosi na ọnụ ọgụgụ nke mkpụrụ ndụ glial na-agbanwe n'ụzọ dabere na mpaghara. Nọmba cell Glia kwụsiri ike na ventromedial PFC n'etiti oge uto na ntozu, mana ọ na-abawanye ihe fọrọ nke nta ka ọ bụrụ 40% na ntozu oke na PFC dorsal (Markham et al., 2007). Ya mere, mgbanwe na DTI na-egosipụta ma glia na axonal mgbanwe mgbanwe.

Ụzọ ọzọ isi chọpụta mgbanwe na myelination bụ inyocha mkpụrụ ndụ ihe nketa. N'ikwekọ na nha anatomical a nụchara anụcha, mkpụrụ ndụ ihe nketa jikọtara na myelination na-abawanye nkwupụta n'oge uto n'ime mmadụ.Harris et al., 2009). Dịka ọmụmaatụ, mkpụrụ ndụ ihe nketa gụnyere MBP (protein nke myelin), MOG (myelin oligodendrocyte glycoprotein), na MAG (glycoprotein metụtara myelin) na-abawanye nkwupụta ha na ntozu okè. Ọ bụ ezie na MBP na MOG metụtara mgbanwe nhazi na myelin, MAG na-etinye aka na njikọta axonal caliber (ọrụ) na ogo nke myelination (ọrụ).Mee et al., 1998). N'ịchịkọta ọnụ, njupụta ọcha na-abawanye na ejiji na-aga n'ihu, nke dị n'ahịrị nke dị iche na ntụgharị U-ọdịdị isi awọ ntozu nke na-egosipụtakarị oge uto.

3.2.2 Ndabere mmekọahụ nke myelination

Esemokwu mmekọahụ na-eme na myelination ma na-ahụ n'oge mmalite nke ntozu okè. Ọtụtụ ọmụmụ na-egosi mmụba dị ukwuu na myelination nke ọtụtụ mpaghara ụbụrụ n'oge ntorobịa ruo n'oge nwoke, mana ọ bụghị ụmụ nwanyị (Blanton et al., 2004; Leussis na Andersen, 2008; Paus, 2010). Kama nke ahụ, myelination na-apụta na mbụ na ụmụ nwanyị. Dịka ọmụmaatụ, esemokwu mmekọahụ na hippocampal myelination nke mmadụ na-apụta mgbe afọ 5 gasịrị, yana nkezi nke 37% nke myelination dị ukwuu na ụmụ nwanyị karịa ụmụ nwoke.Benes et al., 1994). A na-ahụ ndịiche dị ka nwoke na nwanyị n'ofe ụdị (dịka ọmụmaatụ, mmadụ, oke (Kodama, 2008)). Site na okenye, myelination na corpus callosum ka ukwuu na ụmụ nwoke, ọ bụ ezie na ụmụ nwanyị nwere obere mkpụrụ ndụ glia na-enye aka (Nunez na Juraska, 1998; Kim et al 1997). N'otu aka ahụ, oke PFC nwere 15% obere mkpụrụ ndụ glia n'ime ụmụ nwanyị karịa ụmụ nwoke site na ntozu oke, nke nwere ike itinye aka na ọdịiche nwoke na nwanyị na olu na mpaghara ahụ.Markham et al., 2007).

Mgbe ekewa nyocha DTI n'ime trajectories nke FA na MD, profaịlụ dị iche iche dị n'etiti usoro, n'ofe mmekọahụ, na n'ofe mpaghara.Asato et al., 2010). Traktị eriri nke arcuate fasciculus (nke jikọtara mpaghara Wernicke na mpaghara Broca) na nke dị ala fronto-occipital fasciculus (nke jikọtara sensorimotor na mpaghara ihu) gosipụtara FA na ụmụ agbọghọ, mana belatara FA na ụmụ nwoke n'etiti afọ 6-20. ; Achọpụtaghị ọdịiche mmekọahụ maka MD (Ashtari et al., 2007; Schmithorst et al., 2008). Mgbanwe ndị a emetụtala IQ na nhazi okwu dị elu na ụmụ nwanyị na-eto eto karịa ụmụ nwoke (Ashtari et al., 2007; Schmithorst et al., 2005). N'ụzọ dị iche, traktị ndị ọzọ adịghị egosi mmụba afọ metụtara afọ a tụrụ anya na FA, ebe MD belatara (Eluvathingal et al., 2007). Usoro ndị na-egosipụta mmụba nke FA na enweghị mgbanwe na radial diffusivity (nke nwere ike ịbụ index demyelination) nwere ike igosi mgbanwe site na mbelata nhụsianya gaa na nzukọ fiber axonal ka ukwuu (ma ọ bụ eriri kwụ ọtọ) n'oge oge ntorobịa (Ashtari et al., 2007). Nhazi karịa nke ọma ga-abụ amụma amụma nke mgbanwe ndị dị otú ahụ.

Ọkwa testosterone metụtara mgbanwe `g' n'ime ụmụ nwoke.Perrin et al., 2008). 'g ratio' na-abawanye na ụmụ nwoke, mana ọ na-agbanwe agbanwe na ụmụ nwanyị (Paus na Toro, 2009). Axonal caliber na-agbanwe n'oge mmepe ma nwee ike ịkọwa mmụba nke DTI na ụmụ nwoke, ebe mgbanwe ụmụ nwanyị na DTI nwere ike igosi myelination nke ọma (Perrin et al., 2009) Nnyocha ndị bụ isi na-egosi na corpus callosum nke nwanyị na-enwe mmetụta nke hormones na-eto eto, na ovariectomy na ụbọchị 20 na oke na-ebelata ọnụ ọgụgụ nke axon myelinated ma e jiri ya tụnyere njikwa.Yates na Juraska, 2008); ngụkọta ọnụ ọgụgụ nke axon na ọmụmụ ihe a emetụtaghị, na-atụ aro na mgbanwe ndị a bụ n'ihi myelin ma ọ bụghị mkpụrụ ndụ. Otu nkọwa nwere ike ịbụ na esemokwu mmekọahụ dị n'oge ndụ nke oligodendrocytes, ebe mkpụrụ ndụ na-anwụ ngwa ngwa na ụmụ nwanyị na-eto eto karịa ụmụ nwoke.Cerghet et al., 2006). Ohere ndị ọzọ na-agụnye mmetụta estrogenic nke na-emegharị hormones gonadal ndị ọzọ (dịka ọmụmaatụ, progesterone), hormones na-akpata nchekasị, ma ọ bụ ọbụna ihe ndị na-eto eto na-eme ka myelination pụta (ntụle na Yates na Juraska, 2008). Nnyocha ndị ọzọ ga-ejupụta oghere ndị na-efunahụ ihe na-efu na ka estrogen si emegharị myelination.

Anyị na-amalite ịghọta ka synaptogenesis na kwachaa na-emekọ ihe na usoro myelinating na ụbụrụ na-arụ ọrụ iji zụlite omume ntorobịa (Paus et al., 2008). Myelin na-arụ ọrụ dị mkpa na mmepe, mana nke ka mkpa, n'ịhazi ọsọ nke ntinye dị iche iche site na ebe dị iche iche gaa na mpaghara enyere. Ngosipụta mmekọrịta bụ ihe kacha mkpa maka mmepe nkịtị ịga n'ihu (Ubi, 2005), na mgbanwe nke myelination na-emetụta n'ọtụtụ ọrịa uche.

4.0. Mmepe mgbanwe ọrụ

Ntụleghachi a ekpuchiwo mgbanwe nhazi nke na-eme n'oge nwata na ntụgharị ndị okenye, ma mgbanwe ọrụ nwere ike igosi usoro nke ha. Ụbụrụ na-eto eto na-eji nhazi ya na akụrụngwa na-agbanwe agbanwe (dịka ọmụmaatụ, glucose metabolism) iji kparịta ụka n'etiti na n'ime ụlọ iji metụta omume. Kedu ka mpaghara ụbụrụ si arụ ọrụ dị iche iche na nzaghachi nye mkpali enyere nwekwara ike ịgwa anyị ka ejikọtara ha na-arụ ọrụ. Na ngalaba a, "njikọ ọrụ" dị ka atụpụtara site na MRI na-ezo aka na mmekọrịta mmekọrịta dị n'etiti mpaghara abụọ.

4.1. Iji ike eme ihe

Mgbanwe ọdịdị ọdịdị nke akọwara n'elu na-ebutekarị mgbanwe arụ ọrụ n'ime ụbụrụ. Ọmụmụ ihe izizi gbasara mgbanwe ọrụ ejirila PET imaging nke glucose mee ka eserese ike ojiji n'ime nhazi akụkụ (nke a na-emepe emepe).Chugani, 1998; Feinberg, 1988). Iji glucose eme ihe n'ime mmadụ na-erute ọkwa okenye site na afọ abụọ nke ndụ.Chugani et al., 1987) ma na-ebili na 4-5 afọ ma na-edobe ala dị larịị ruo afọ 10 tupu ịkwachaa site ~ 50% site na 16-18 afọ (Chugani, 1998). A na-egosipụta mkpụrụ ndụ ihe nketa metụtara glucose metabolism, dịka, gene acyl coA dehydrogenase (ACADSB), n'ogo dị elu n'oge uto, n'agbanyeghị na amabeghị ọrụ ọrụ ha n'oge a.Harris et al., 2009).

Ihe nrịbama ndị ọzọ nke ọrụ ụbụrụ na-enyocha metabolism nke ụbụrụ, dị ka n-acetylaspartate (NAA; akara nke neurons na usoro), phosphocreatine (PCr; ike ike), na membrane phospholipid metabolism (ya na ndị na-eme sPME na sPDE) ejirila magnetik enyocha ya. resonance spectroscopic imaging (MRSI) iji nye ndepụta mmepe na-adịghị emerụ ahụ. A na-akọwa mgbanwe na akara ndị a na mpekere axial nke ụbụrụ n'ofe nwoke na nwanyị 6-9.5, 9.5-12, na 12-18 afọ na n = 106 isiokwu (Goldstein et al., 2009). Ntụnyere n'etiti 6-9.5 afọ na 12-18 afọ egosighi ihe dị iche na NAA, nke na-egosi na ọ dịghị akara neuronal mgbanwe. Nlebanya a dị iche na mfu neuronal nke amaara nke ọma site na nha ozugbo na anụ ahụ nwụrụ anwụ (dịka ọmụmaatụ, Huttenlocher, 1979). Otú ọ dị, NAA na-enye acetate maka olidgodendrocytes bụ ndị na-ahụ maka mmepụta myelin. Ya mere, ọ dịghị mgbanwe ọ bụla na NAA n'ofe mmepe ntorobịa nwere ike igosi nguzozi n'etiti ọnwụ neuronal na ụbara myelination. PCr belatara n'ime ndị ntorobịa, mana ọkwa dị elu na pasent isi isi awọ na oke sPME/sPDE, nke na-egosipụta mgbanwe mgbanwe akpụkpọ ahụ phospholipid, dị elu. PCr na pasent isi awọ jikọtara ya na afọ, mana NAA, sPME, sPDE, na sPME/sPDE adịghị. Ọ bụ ezie na mgbanwe ụfọdụ nwere ike ịgbaghara site na ijikọta nwoke na nwanyị, data ndị a na-egosi na MRSI adịghị egosi mgbanwe mgbanwe metabolic dị mkpa nke afọ.

4.2 Njikọ ọrụ dị ka akọwapụtara na MRI

Njikọ ọrụ bụ ụzọ ọzọ a na-eji egosi mmekọrịta dị n'etiti oge n'etiti mpaghara ịgbalite n'oge ọnọdụ ezumike ma ọ bụ n'oge ọrụ fMRI.Fair et al., 2008; Supekar et al., 2009; Thomason et al., 2009; Zuo et al., 2010). A na-akpọkwa maapụ nke njikọta ọrụ dị ka njikọ (Biswal et al., 2010), yana ngwa fMRI na-anọchite anya ngwa mpaghara a na nso nso a.Lichtman na Sanes, 2008). Ụzọ a na-enye ụfọdụ nghọta na mmepe ụbụrụ nke ndị nọ n'afọ iri na ụma, n'agbanyeghị na ọ bụ nanị site na ụfọdụ nleba anya na a na-ahụ 'njikọ ọrụ' n'ebe ndị na-adịghị na ezi njikọ anụ ahụ.Mmanụ a etụ et al., 2009; Koch et al., 2002). fMRI steeti izu ike gbadoro ụkwụ na nlebanya na nnukwu njupụta na-enweghi oke obere ugboro (<0.1 Hz) na-eme (Biswal et al., 2010). Ụzọ iji ghọta njikọta ọrụ gụnyere mkpụrụ dabere (ebe a na-eji aka achọpụta ebe mmalite iji chọpụta ebe mmalite), nyocha akụrụngwa onwe ya (ICA), na nyocha ngalaba ugboro ugboro. Mmepe na-arụ ọrụ nke sistemu ụbụrụ dị iche iche gụnyere nchikota nke mbelata njikọ dị mkpụmkpụ (ya bụ, nkewa) na ịba ụba njikọ ogologo ogologo (ya bụ, njikọta)Fair et al., 2007; Stevens et al., 2009). N'ikwu ya n'ụzọ ọzọ, mmepe na-esi na mpaghara gaa na netwọk kesawanye ka mpaghara dị iche iche na-ejikọta ọnụ (Fair et al., 2009). Njikọ njikọ a abụghị mmekọrịta, kama nke ahụ, mpaghara nke ọ bụla na-ejikọta ya wee jikọta (Supekar et al., 2010).

Ọmụmụ njikọ njikọta ọrụ nke fMRI izu ike na-egosi na “netwọk ndabara” dị na ụbụrụ mgbe ọ naghị arụ ọrụ nke ọma ozi. Netwọk ndabara gụnyere cortex cingulate dị n'azụ, mPFC, lobes medial temporal, na gyrus angular. Ihe owuwu ndị a na-egosipụta ngbanwe, mgbagharị ugboro ole dị ala (0.1 Hz) mgbe onye ahụ nọ jụụ, ọnọdụ izu ike. Ka ụbụrụ na-esiwanye ike n'etiti mpaghara n'etiti nwata na oge uto (Fair et al., 2008), ụbara njikọta n'ime netwọk ndabara na-eme n'oge mgbanwe a (n'etiti 9-12 afọ; Broyd et al., 2009). Netwọk ndabara ka echepụtala ka ọ rụọ ọrụ na imepụta ihe, ebe mbelata n'ime netwọk ndabara ejirila schizophrenia na autism.

Otú ọ dị, netwọk ndị ọzọ na-arụ ọrụ dị na ụbụrụ. N'ime ọmụmụ ihe atụnyere ndị na-eto eto (pụtara afọ 12.5 ± 0.51 [SD] afọ) na ndị na-eto eto (22.2 ± 1.67 [SD] afọ) n'ime otu nwoke na nwanyị agwakọta, 13 netwọk ọrụ na-arụ ọrụ bụ ndị a chọpụtara (Jolles et al., 2010). N'ime netwọk ndị a, asatọ gosipụtara mmụba dị n'etiti mpaghara cortical n'oge uto, abụọ egosighi ihe dị iche na ọrụ, na atọ jikọtara ya na isi ọrụ visual ma ọ bụ sensorimotor (ie, sensorimotor, visual system, na ventral stream networks) ma gosipụta obere ọrụ n'oge uto. karịa ịbụ nwata. Nchọpụta netwọkụ ndị a ga-emezi nyocha n'ọdịnihu maka ihe kpatara ha ji egosipụta mgbanwe ndị metụtara afọ.

5. Ọrụ mmepe nke sekit

N'oge uto, mgbanwe omume dị egwu na-ejikọta na mgbanwe ndị metụtara afọ n'ime ụbụrụ. A na-ahụ nnukwu nlebanya nke omume ndị ntorobịa n'ebe ndị ọzọ (Okwu, 2000), mana anyị na-enye nkọwa dị nkenke banyere otu mgbanwe kpọmkwem na nhazi ọrụ n'oge uto nwere ike isi kọwaa ụfọdụ n'ime omume ndị a. N'ime nhazi nke ịmepụta ụbụrụ, mpaghara ọ bụla nwere oge mmepe nke ya nke ntozu okè (Tau na Peterson, 2010). N'ozuzu, akụkụ cortical na-etolite ma emechaa karịa mpaghara subcortical, dịka a tụlere n'elu. Oge mmepe ma ọ bụ mmepe nke mbụ n'ime oghere nke nhazi netwọkụ neuronal nwere ike ịmalite usoro ihe omume mmepe nke domino nke na-agbanwe ọnọdụ nke mpaghara ụbụrụ ọtụtụ.Ernst na Fudge, 2009; Haber na Rauch, 2010). Site n'echiche a, ọmụmụ ogologo oge ga-enye aka n'ịchọpụta usoro mgbanwe ụbụrụ mpaghara ka ihe omume dị iche iche na-apụta (Gogtay et al., 2006; Sowell et al., 2004). Dịka ọmụmaatụ, Shaw na ndị ọrụ ibe (Shaw et al., 2007) egosila na mmepe cortical lags na ụmụaka nwere ADHD gbasara ndị ọgbọ ha, mana ọ na-enweta site n'ịbụ okenye. N'ụzọ dị iche, mmalite nke schizophrenia nke nwata na-ejikọta ya na ịkwachaa azụ azụ na mbụ karịa ka a na-ahụ na ụmụaka.Rapoport et al., 1999). Ọmụmụ ihe dị ka ndị a dị mkpa maka nsuso usoro nsogbu ahụ, mana n'otu oge na-egosipụtakwa windo mmepe nke nwere ike ịkarị ma ọ bụ na-adịghị emetụta mmetụta ndị ọzọ.

Mpụta nke psychopathology n'oge oge ntorobịa na ngalaba dị ukwuu nke ụgwọ ọrụ na nhazi metụtara mmetụta abụghị ihe ndabara. Nyere ọnụ ọgụgụ nke mgbanwe dị ịrịba ama na-eme n'oge a, usoro ndị na-aga n'ihu ma ọ bụ na-eduhie na mbụ na ndụ na unmasked site na mgbanwe ndị a (Andersen, 2003; Andersen na Teicher, 2008; Weinberger, 1987; Laviola et al., 2003) ga-egosipụta n'oge a. Mkpa ọ dị ịkọwapụta na ijikwa oge dị nro dabere n'ịghọta nsonaazụ ọjọọ na usoro mmepe. Na mgbakwunye, ọtụtụ nsogbu nwere ihe ndabere na usoro neurodevelopmental agahiere ụzọ. Mkpughe mbụ na nsogbu na-anọchi anya ihe dị oke egwu maka ọtụtụ nsogbu. Dịka ọmụmaatụ, nchọpụta ọrịa na-efe efe egosila na ikpughe na nhụjuanya na-ebute mmụba dị elu na nnukwu nsogbu ịda mbà n'obi (Anda et al., 2006; Anda et al., 2002; Chapman et al., 2004), ọgba aghara akparamàgwà mmadụ, iji ọgwụ eme ihe n'ụzọ na-ezighị ezi (Andersen na Teicher, 2009), na igbu onwe ya, na ịda mbà n'obi dị ka ihe na-esokarị ndị okenye na-emegbu oge mbụ (Putnam, 2003; Zisook et al., 2007).

5.1. Mmepe na-arụ ọrụ nke sekit mmetụta

Mmepe na-arụ ọrụ nke sekit na usoro na ụbụrụ dị mgbagwoju anya, na-enwe ọtụtụ ihe na-akpụ akpụ iji jikọta ọnụ. Dịka ụzọ isi ruo sekit mmepe, anyị na-enye nkọwa ndị a ka ha na-emetụta ma mmetụta na ụgwọ ọrụ n'oge oge ntorobịa. Ụzọ ndị a adịghị agụnye ọmụmụ ihe na-enweghị atụ na nke dị mkpa nke na-enyocha mgbanwe omume na ọgwụ na-eme n'oge uto, ma na-elekwasị anya na ọmụmụ ndị nwere mmekọrịta neuroanatomical na mgbọrọgwụ ha.

Ọtụtụ omume na mkpali mmadụ na-ebilite site na mkpakọrịta ndị enwetara mbụ n'etiti ihe mkpali na-akwụghachi ụgwọ ma ọ bụ nke na-adịghị mma na ọnọdụ ha na-eme (Cardinal et al., 2002). Mkpakọrịta ndị a siri ike, mmụta na-akwalite omume anyị ugbu a na ọdịnihu (Cardinal et al., 2002) ma mee site na usoro nhazi nke Pavlovian (Rosenkranz et al., 2003). A na-ahazi ozi gbasara gburugburu ebe obibi na mmetụta n'ime amygdala basolateral (BLA) (Grace na Rosenkranz, 2002), nke na-etolite mkpakọrịta siri ike n'etiti ihe mkpali na-ebu amụma na ọ ga-eme ihe na-agụ agụụ ma ọ bụ nke na-adịghị mma, ma na-emepụta "mmetụta" n'ime BLA (Cardinal et al., 2002; Laviolette et al., 2005; Schoenbaum, 2004; Lee et al., 2003). Agbanyeghị, ịzaghachi ihe mkpali enyere kwesịrị ịkọwapụta na dabara adaba n'ihe gbasara ọnọdụ, mkpa mmetụta uche, ma ọ bụ nlebara anya dịka ọ metụtara nhọrọ (Paus et al., 1996). Usoro a na-eme na PFC (Cardinal et al., 2003; Rebec na Sun, 2005; Schoenbaum, 2004; Ventura et al., 2007). Noradrenergic na ndị na-anabata dopaminergic na PFC na-edozi usoro nlebara anya, omume na mmetụta site na ime ka njikọ netwọk dị n'etiti neurons na ntinye aka na-ekekọrịta (Arnsten, 2009). N'ime mPFC, a na-ahazi salience nke ozi iji hazie nlebara anya ahọpụtara.

Ya mere, a na-ebufe ozi sitere na BLA na mPFC site na amụma glutamatergic.Bechara et al., 1999; Laviolette et al., 2005; McDonald na Pearson, 1989), ebe a na-edozi ya maka salience (Schultz, 1998) na njehie ndị dị mkpa maka ịkọ ihe ga-eme n'ọdịnihu (Falkenstein et al., 2000; Ahịa, 1999). N'ihi ya, ihe mkpali ndị na-ebu amụma ihe ga-esi na ya pụta nwere ike ịzaghachi ya n'ụzọ dabara adaba nke ọma (Pezze et al., 2003). A na-arụ ọrụ a site na akara dopaminergic na mPFC (Jackson na Moghaddam, 2004), nke na-etinye ozi agbakwunyere nke salience na ihe ọhụrụ na ozi mmetụta uche (Cardinal et al., 2002; Milad na Quirk, 2002) imetụta ihe mgbaru ọsọ na-eduzi, omume mkpali. MPFC na-eziga ozi a na nucleus accumbens ozugbo (Goto na Grace, 2005; Voorn et al., 2004), ma ọ bụ na-apụtaghị ìhè site na amygdala. Na-esote, ọrụ nsonaazụ n'ime mPFC, ozugbo ma ọ bụ na-apụtaghị ìhè, na-emetụta omume mkpali na nucleus accumbens.

Atụpụtala nhazi na-eto eto n'etiti amygdala na PFC iji mee ka ọ pụta ìhè na-egbu oge nke ọrịa na-emetụta ruo n'oge uto (Ernst et al., 2006). N'ime ụdị triadic nke Ernst na ndị ọrụ ibe ha tụrụ aroErnst et al., 2006), sistemu ịzere ihe metụtara amygdala na-eme ka omume PFC na-etobeghị enyocha ya. Dị ka ihe nlereanya a si dị, nucleus accumbens na-emezi ike nke njikọ dị n'etiti agụụ na ime ihe ike (aversive conditioning).Horvitz, 2002). Ozizi a bụ otu n'ime ole na ole na-adịghị ahụkebe nke na-agụnye ihe a maara banyere neurobiology nke ịda mbà n'obi n'ime usoro mmepe. Otú ọ dị, ozizi ahụ na-egosi na ụmụaka na ndị nọ n'afọ iri na ụma ga-esi na ịda mbà n'obi ha pụta site na ntozu oke cortical na njikọta, nke na-abụghị ikpe.Andersen na Teicher, 2004; 2008).

Anyị enyochala mgbanwe mmepe n'oge na-adịbeghị anya n'oge ntorobịa nke nwere ike ịbawanye ike ịda mbà n'obi (Andersen na Teicher, 2008). Na nkenke, ụmụaka nwere ọtụtụ ọrụ karịa ndị okenye na amygdala na nzaghachi maka mkpali mmetụta uche (Killgore et al., 2001), nke na-akawanye njọ na ụmụaka na ndị nọ n'afọ iri na ụma nwere nsogbu nchekasị nke ọha mmadụ (Beesdo et al., 2009). Otú ọ dị, nucleus accumbens na-etinyekwu aka na nhazi nke agụụ na mkpali na-adịghị mma n'oge uto kama na amygala (Ernst et al., 2005). Mbanye nke PFC na nzaghachi nye mkpali mmetụta uche anaghị eme ruo mgbe ọ bụ okenye (Killgore et al., 2001). Kpọmkwem, nke a dabara na nnwale traktị nke na-egosi mmepe mmepe nke BLA ruo PFC n'oge uto (Cunningham et al., 2002), mana nke kachasị mkpa, ọnụ ọgụgụ kasị elu na ntinye PFC na ntinye BLA n'oge uto (Cressman et al., 2010). Ọnụ, ịba ụba njikọ anatomical nwere ike inye ihe ndabere maka nhụpụta nke mgbaàmà ịda mbà n'obi na igbu oge (na-eto eto) nke na-egosipụta ọnọdụ ntozu oke a ka nchịkwa nchịkwa na-emetụta mmetụta na-etolite (ma ọ bụ na-etolite).

5.2. Mmepe arụmọrụ nke sekit ụgwọ ọrụ

Ọkachamara MRI na ihe ọmụmụ electrophysiologic na-egosipụta ọrụ pụrụ iche nke nkewa n'ime cortex n'ihu na nhazi ụgwọ ọrụ. The mPFC (Broadman mpaghara [BA] 10/12/32 na gụnyere anterior cingulate cortex; BA 24) na-anabata nsonaazụ nke ụgwọ ọrụ: ọ na-arụ ọrụ ma ọ bụrụ na a na-atụ anya ụgwọ ọrụ na-anata na gbanyụọ mgbe enwetaghị (Knutson et al., 2003; Schulz et al., 2004). The orbital frontal cortex (OFC) na-akọwapụta nsonaazụ ndị a na-atụ anya ya na atụmatụ mkpali dabere na ụgwọ ọrụ nwere ike. Ndị OFC na-arụ ọrụ dị mkpa n'ịmụta mgbanwe na nkwado na-egbu oge (Dalley et al., 2004) site na njikọ ya na mpaghara sensory, limbic, frontal na subcortical. A na-ekewa OFC na-arụ ọrụ yana akụkụ etiti na-aza nke ọma iji kwụọ uru ụgwọ ọrụ, ebe akụkụ akụkụ ahụ na-egbochi usoro metụtara ụgwọ ọrụ gara aga.Elliott et al., 2000; Elliott et al., 2003; London et al., 2000).

Accumbens (mpaghara ventral striatal) na-anabata saliency (Ernst et al., 2004), valence (agụụ ma ọ bụ aversive) (Jensen et al., 2003) na amụma amụma nke ụgwọ ọrụ (ụgwọ ọrụ a na-atụghị anya ya na-arụ ọrụ karịa ụgwọ ọrụ amụma)Berns et al., 2001; Elliott et al., 2000)), ma ọ bụghị akụkụ moto (Zink et al., 2004). N'oge uto, accumbens na-anabata ihe karịrị OFC iji kwụọ ụgwọ (Galvan et al., 2005). Ejikọtara ọnụ, data ndị a na-atụ aro na accumbens na-eto eto na-eme mgbanwe na nhazi ụgwọ ọrụ (Galvan, 2010).

Otú ọ dị, ihe àmà nke otú usoro cortical na subcortical si anabata ụgwọ ọrụ mkpali na-egosi na cortex na-arụ ọrụ ọbụna karị na mgbanwe ndị ntorobịa na nhazi ụgwọ ọrụ. Ọmụmụ anụmanụ egosila na ngbanwe nhazi ụgwọ ọrụ n'oge uto site na ịkwachaa na ike ilekwasị anya na netwọk cortical ka netwọk ahụ na-eto eto ma ghọọ ndị okenye.Brenhouse et al., 2008; Crews et al., 2007). Ọmụmụ fMRI nke ụlọ ọgwụ na-atụ aro na ma ventral striatum na mPFC na-arụ ọrụ iji kwụọ ụgwọ mkpali n'oge uto (Bjork et al., 2004). Tupu mgbanwe a, ọrụ BOLD metụtara ụgwọ ọrụ na-emepụta ihe na-agbasawanye ma na-arụ ọrụ nke ọma nke mpaghara ihu na ụmụaka karịa ndị okenye (Durston et al., 2003). Agbanyeghị, ụmụaka na-egosipụta nkwalite dị ukwuu na ventral striatum (accumbens) (Ernst et al., 2005; Galvan et al., 2006). Dị ka anyị maara ntakịrị usoro maka mmepe ụgwọ ọrụ n'ime mmadụ, anyị ga-ebute nyocha nke izizi maka nghọta ka ukwuu.

A na-egbu oge ntozu nke mPFC n'ọtụtụ mpaghara ụbụrụ ndị ọzọ (Andersen et al., 2000; Huttenlocher, 1979) wee rute njupụta synapti kacha nso na okenye (Benes et al., 2000). Mmụba ome nke neurons dopamine (Benes et al., 1996; Kalsbeek et al., 1988; Verney et al., 1982), njupụta nnabata (Andersen et al., 2000; Leslie et al., 1991), na ọrụ sistemụ ozi nke abụọ (Andersen, 2002) mebie na mbanye dopaminergic kwalitere na mPFC n'oge uto. Nchọpụta ndị na-adịbeghị anya gosikwara mmụba metụtara afọ na ntinye D1 nke mkpụrụ ndụ spiking na-adịghị ngwa ngwa na mPFC, nke na-eme mgbe ntozu okè (Tseng et al., 2006), na ọnụ ọgụgụ kasị elu na ọnụọgụ ọkụ nke VTA dopaminergic neurons n'otu afọ a (McCutcheon na Marinelli, 2009). Nkwuwapụta n'elu nke ndị na-anabata D1 na glutamatergic pụta na accumbens na-arị elu n'oge uto n'otu n'otu na omume ịchọ ọgwụ.Badanich et al., 2006; Brenhouse et al., 2008). Ọnụ ọgụgụ ndị nnabata a etinyela aka na nlọghachi azụ ọgwụ, ya mere nkwuwa okwu ya n'oge uto bụ ihe kwesịrị ịrịba ama (Kalivas, 2005). Mgbanwe ndị a na nhazi ụgwọ ọrụ cortical nwekwara ike imetụta nzaghachi subcortical na psychostimulants.

N'ụzọ dị iche, ọkwa basal nke extracellular dopamine na nzaghachi dopaminergic maka ihe na-akpali akpali adịghị agbanwe nke ọma n'etiti ndị nọ n'afọ iri na ụma na ndị okenye na accumbens.Frantz et al., 2007) ma ọ bụ mPFC (Jezierski et al., 2007). Agbanyeghị, oke dị n'etiti cortical:accumbens ngosipụta nke mkpụrụ ndụ mmalite mmalite ozugbo c-fos na nzaghachi na-akpali akpali na-abawanye n'etiti oge uto na okenye (Andersen et al., 2001). Ọzọkwa, amphetamine na-emepụta subcortical> cortical activation patterns of c-fos na ụmụaka.Andersen et al., 2001), ma cortical > subcortical ịgbalite na ndị nọ n'afọ iri na ụma (Cao et al., 2007). N'ịchịkọta ọnụ, data ndị a na-egosi na ndị na-eto eto dị iche na ndị nọ n'afọ iri na ụma, bụ ndị toro eto karịa, na nzaghachi ha na-akpali akpali na subcortically. N'ikwu ya n'ụzọ ọzọ, o yikarịrị ka iji ihe eme ihe na-ebili nke ukwuu n'oge uto na-esite na mmetụta kpọmkwem ma ọ bụ na-apụtaghị ìhè nke usoro cortical na ọrụ subcortical.

5.3. Ọrụ mmepe nke cognition

Usoro nnwale dị ka Stroop, Simon, Flanker, Go/No-Go, na ọrụ nkwụsị-ịkwalite chọrọ mbelata omume akpaka karịa iji rụọ obere akpaaka. Usoro nlebara anya, nkwụsị nzaghachi, na esemokwu na nleba anya njehie bụ usoro ọgụgụ isi nke na-arụ ọrụ nke njikwa ọgụgụ isi na ịrụ ọrụ nke ọma. Ịrụ ọrụ ndị a niile na-akawanye mma n'oge mmepe niile, mana ọ naghị abịaru nso n'ogo ndị okenye ruo mgbe ọ dịkarịa ala n'oge nwata ma ọ bụ n'oge uto (Bunge et al., 2002; Casey et al., 1997; Davidson et al., 2006; Luna na Sweeney, 2004; Rubia et al., 2000). Dị ka ọ dị na ebe nchekwa na-arụ ọrụ, ikike ịchịkwa onwe ụmụaka nwere ike ịkabiga ya ngwa ngwa site n'ịbawanye mkpa ọrụ. N'ime ndị okenye, nhazi onwe onye na-adabere na mpaghara cortical sara mbara dị ka mpaghara moto mgbakwunye, oghere ihu ihu, cortex cingulate cortex, dorsolateral PFC, ventralPFC/lateral orbitofrontal cortex, yana mpaghara oge, na mpaghara parietal niile nwere njikọ na striatum na subcortex (Leung et al., 2000; Marsh et al., 2007).

Ịzaghachi nke ọma maka mkpali gburugburu ebe obibi chọrọ nlebara anya nhọrọ na ntụzịaka mkpali, yana nkwụsị nke omume ndị a na-achọghịzi ma ọ bụ ndị na-ekwesịghị ekwesị. A na-atụ nhịahụ a na nnwale site na mgbochi nzaghachi, nke gụnyere usoro atọ jikọrọ ọnụ, dịka Barkely tụrụ aro (Barkley, 1997): 1) igbochi nzaghachi mbụ nke nwere ike, 2) ịkwụsị nzaghachi na-aga n'ihu ma ọ bụ ịzaghachi na-egbu oge, na 3) igbochi nnyonye anya ma ọ bụ ihe ndọpụ uche n'oge oge igbu oge. Basal ganglia na PFC na-etinye aka na usoro ndị a (Casey et al., 2008). N'ozuzu, ebe basal ganglia na-achịkwa mgbochi nke omume na-ekwesịghị ekwesị (Mink, 1996), PFC na-eme ihe iji gbochie nnyonye anya na ozi dị mkpa site na ozi asọmpi (Miller na Cohen, 2001).

N'ụzọ dị iche na nsoro-ịzere, nke na-achọ njirimara salience na-akpali akpali ma na-agbaziri ya nke ukwuu site na imekọ ihe ọnụ nke PFC, striatum, na amygdala (nyochaa (nyocha)Ernst na Fudge, 2009)), mgbochi nzaghachi na-ewe ndị ọrụ sekit nke na-ahazi atụmatụ moto na oge (oge)Deiber et al., 1999). Ọrụ bụ isi nke netwọk fronto-striatal na-enye onwe ya na profaịlụ mmepe dị iche karịa nke mkpali na usoro nlebara anya nhọrọ.

5.4. Mmepe nke mgbochi nzaghachi

Ọ bụ ezie na ndị nọ n'afọ iri na ụma nwere ike ịrụ ọrụ ọgụgụ isi ọkaibe, ikike ime nke a na-aga n'ihu na-aga n'ihu n'oge uto na n'ime oge ntozu, Mmụba nke usoro a n'oge mmepe niile na-egosi na nkwado neurobiological nke cognition na-agbaso usoro ọganihu n'otu aka ahụ. Ụmụntakịrị na-egosi nrụ ọrụ dị elu karịa ndị okenye na mpaghara ihu lobe (Bunge et al., 2002) gụnyere gyrus n'ihu etiti etiti na akụkụ etiti nke gyrus ihu dị elu nke abụọ (Booth et al., 2003). Nke a kwekọrọ n'ọdịiche metụtara afọ na izi ezi na oge mmeghachi omume n'ọrụ aga/agaghị aga n'ofe nwata. N'ụzọ na-akpali mmasị, ọmụmụ ihe nkwonkwo DTI na fMRI nke Stevens na ndị ọrụ ibe mere (Stevens et al., 2009) kọrọ njikọ kpọmkwem n'etiti mgbanwe ndị metụtara afọ na njikọta ọrụ n'etiti frontopolar bilateral, right parietal cortex na caudate ziri ezi, myelination na-abawanye, na ịrụ ọrụ nke ọma na ọrụ Go / No Go. N'ime ọmụmụ DTI ọzọ, mmechi nzaghachi na 7-13 afọ na-ejikọta ya na FA dị elu na MD dị ala na ma gyrus n'ihu nke ọma na cortex moto pre-supportary (Madsen et al., 2010). Usoro mmepe ahịrị ahịrị nke myelination nke a tụlere n'elu kwekọrọ na mmepe ahịrị ahịrị pụtara akara njikwa, n'ihe metụtara ntụgharị ntụgharị U-ụdị mmetụta mmetụta na nhazi ụgwọ ọrụ. Ụmụntakịrị na-egosipụtakwa ike ịrụ ọrụ ka ukwuu karịa ndị okenye na oghere caudate aka ekpe n'oge aga/no-go (Booth et al., 2003) na kwụsị (Rubia et al., 1999) ọrụ. Atụbere basal ganglia ka ọ tinye aka na mgbochi nke omume na-ekwesịghị ekwesị (Casey et al., 2001), na basal ganglia yiri ka ọ toro n'ahịrị site na nwata ruo n'oge.

A tụlere neurobiology bụ isi nke sekit ndị a n'elu ma ọ bụ ka amụbeghị ya n'ime ọnọdụ mmepe. Ọ bụ ezie na enwere ọtụtụ data neuroimaging gbara gburugburu ọrụ mgbochi nzaghachi, enwere obere nyocha nke neurochemistry n'azụ sistemu ndị a (maka nyocha zuru oke, lee Eagle et al., 2008). Otu n'ime nsogbu ndị bụ isi jikọtara na nhazi nhazi nke omume ndị a bụ izu ndị a chọrọ ịzụ ụmụ anụmanụ ka ha rụọ ọrụ ndị a, nke na-egbochi ọmụmụ ha n'oge mmepe. N'iburu mkpa ọ dị na njikwa ọgụgụ isi na mkpali iwu n'oge ntolite ntorobịa ruo n'ịbụ okenye, ngalaba a chọrọ nlebara anya karịa ka ọ natara.

6. Ahụmahụ na-akpụzi ụbụrụ ụbụrụ

Ọ bụ ezie na mkpụrụ ndụ ihe nketa na-enye ụkpụrụ iji wuo ụbụrụ, nwee ahụmahụ ọkpụkpụ ụbụrụ iji kwekọọ mkpa gburugburu ebe obibi. Ọdịmma ikpeazụ nke synapse enyere dabere na nkwado ọrụ. Ụbụrụ na-eto eto abụghị nanị ihe pụrụ iche na-emetụta mmetụta gburugburu ebe obibi, ma oge uto bụkwa oge mgbe ahụmahụ mbụ pụtara (Andersen, 2003; Andersen na Teicher, 2008). Netwọk akwara dị mgbagwoju anya na-etolite n'oge uto, ndị a n'aka nke ya na-akpụkwa ya site na ọrụ mberede na ahụmahụ na-eduzi (Ben-Ari, 2002; Francis et al., 2002; Katz na Shatz, 1996; Zhang na Poo, 2001). Nyocha anyị gara aga (Andersen, 2003) tụlere mmetụta dị ukwuu mmetụta gburugburu ebe obibi na-enwe na mmepe ụbụrụ. Akwụkwọ ntụleghachi ndị ọzọ na-atụle mmetụta mkpughe nrụgide na-enwe na mmepe ụbụrụ nke ndị nọ n'afọ iri na ụma (Andersen na Teicher, 2008; 2009). Ngosipụta na ọgwụ psychotropic n'oge mmepe ga-agbanwekwa usoro nke trajectory, yana mmetụta na-apụta n'oge uto (Brenhouse et al., 2009; Ansorge et al., 2008).

Summary

Ọdịdị na oke mgbanwe nke ndị nọ n'afọ iri na ụma n'ime ụbụrụ neuroanatomy na-agbanwe mgbe niile ka ngwá ọrụ nyocha anyị na-aghọwanye nke ọma. Enwere ike inwe ekele zuru oke naanị mgbe a na-amụ mpaghara n'ime nkewa arụ ọrụ (dịka ọmụmaatụ.Gogtay et al., 2006)), na oge zuru oke nke njirimara, na mgbe ahụmahụ mbụ (Andersen na Teicher, 2008) na ihe ndị ọzọ (dịka ọmụmaatụ, mmekọahụ, Tanner ogbo) na-echebara echiche. Usoro oge ezughị ezu na ọmụmụ ihe mbụ emeela ka e nwee nkwubi okwu na-ezighi ezi banyere oge ntozu okè (a tụlere na McCutcheon na Marinelli, 2009) na ma ahụmahụ ndị mbụ na-emetụta mmepe n'ezie. Nyocha a na-enye nkọwa nke nghọta anyị ugbu a banyere mgbanwe ndị ntorobịa na ụbụrụ n'oge mgbanwe ya site na nwata ruo n'oge. Usoro a dị ịrịba ama na-agbanwe agbanwe nke ukwuu n'ihi plasticity nke na-enye ohere ka usoro anụ ahụ na-eme ka ọ kwekọọ na mkpa nke gburugburu ebe obibi ya.

Na 

Nyocha 1 

Usoro iheomume nke usoro mmepe n'ofe mmadụ na òké. Ogwe pink na-anọchite anya usoro iheomume maka ụmụ nwanyị, nke na-aga n'ihu nke ụmụ nwoke, na-anọchi anya na ogwe anụnụ anụnụ. Ngosipụta nke ndị na-anabata ya ("ectopic") na-apụta n'oge ndụ na ...

Isi

  • Anyị na-enyocha oge uto dị ka ọkwa mmepe dị iche iche.
  • A na-ejikọta mgbanwe neuroanatomical na mmetụta gburugburu ebe obibi na ihe ndị chọrọ.
  • Usoro mmepe na-emekọrịta ihe na mgbanwe dabere na mmekọahụ.
  • Anyị na-eji ngwa nyocha na-enwewanye mmetụta na-ekwurịta mgbanwe nke nghọta.

Ihe odide ala

Nkwupụta Nkwupụta Onye Nkwusa: Nke a bụ faịlụ PDF nke ihe odide a na-akwadoghị nke akwadora maka mbipụta. Dịka ọrụ nke ndị ahịa anyị, anyị na-enye nsụgharị mbụ nke ihe odide ahụ. Ihe odide a ga-enwe ngbanwe, nhazi, na nyochaa ihe kpatara ya tupu e bipụ ya na nke ikpeazụ ya. Biko rịba ama na n'oge mmepụta usoro njehie nwere ike ịchọta nke nwere ike imetụta ọdịnaya ahụ, ihe niile gbasara iwu na-emetụta akwụkwọ akụkọ ahụ.

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