Neurosci Biobehav Rev. 2012 Pherekhong; 36(1): 479-501. E phatlalalitsoe inthaneteng 2011 August 25. doi: 10.1016 / j.neubiorev.2011.08.005
Rebecca Hohle1 'me David H. Zald2
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
Patlisiso ea kelello le kelello e ntse e bonts'a hore mekhoa ea maikutlo e sebelisana le likarolo tse ling tsa kelello. Boithuto bo bonts'itse bokhoni ba susumetso ea maikutlo ho susumetsa ts'ebetso e pharalletseng, le bokhoni ba batho ba ho sebelisa mekhoa e laoloang ho laola maemo a maikutlo. Likarolo tsa cortex ea pele li bonahala li bapala karolo ea bohlokoa litšebelisanong tsena. Leha ho le joalo, mokhoa oo litšebelisano tsena li kenngoeng ka ona o lula o hlakisitsoe ka mokhoa o fokolang. Thutong ea hona joale re hlalosa khokahano ea anatomical lipakeng tsa libaka tse ka pele le tse kahare tsa pele le kopanyo ea tsona le libaka tsa maoto. Ke karolo e ka tlase feela ea libaka tse tlang pele e ka susumetsang ts'ebetso ea amygdalar, 'me joalo ka mehlala ea taolo ea maikutlo le mehlala ea taolo ea maikutlo e lokelang ho qobelloa ho ba le tšusumetso ea tšusumetso ea tšusumetso. Re shebisisa hape hore na mokhoa o ikhethileng oa khokahano ea feedxp le karabelo lipakeng pakeng tsa libaka tsena o ka laola mofuta oa phallo ea tlhahisoleseling lipakeng tsa libaka tse kenang ka hare ho sebaka sa dorsal pele le amygdala. Mekhoa ena ea khokahano e hanana le maikutlo a 'maloa a hlalosoang ka tlhaho ea puisano lipakeng tsa maikutlo le kelello.
1. Selelekela
Patlisiso e fumanehang e lekola hore na maikutlo a ikopanya joang le likarolo tse ling tsa kelello. Mosebetsi o joalo o hlakisitse litsela tseo tlhahisoleseling e netefalitsoeng ka maikutlo e ka lebisang tlhokomelo ea leeme kapa e sa rateheng (Ahman et al. 2001; Boholo et al. 2005; Mathews & Wells, 1999), le ho susumetsa mekhoa ea liqeto (Knutson et al. 2008). Ka nako e ts'oanang, lingoliloeng tse ntseng li hola li bonts'a hore mekhoa ea ho lemoha e kang ho lekola botjha e ka laola karabelo ea maikutlo (Jackson et al. 2000; Kim le Hamann, 2007; Ochsner et al. 2002; Ochsner, Ray, et al. 2004; Ray, Wilhelm le Gross, 2008). Kannete, tšebelisano lipakeng tsa mesebetsi e hlalosoang e le ea maikutlo kapa e le tlhalohanyo e matla e lekane ho tsosa karohano e arohaneng hangata pakeng tsa libaka tsena (bona ka mohlala Pessoa, 2008). Leha ho le joalo, karohano e fana ka boleng ba 'nete ka hore ts'ebetso ea maikutlo e na le litšobotsi tse ikhethang tsa ts'ebetso tse ka khetholloang ho libaka tse ling tsa kelello ka mokhoa o ts'oanang moo lits'ebetso tsa tlhokomelo kapa tsa memori li nang le litšobotsi tse fapaneng mme li ts'oaroa ka mokhoa o fapaneng ka marang-rang (leha ho le joalo ka linako tse ling li fetellana) ea libaka tsa boksi.
Tsela eo maikutlo le maqhubu a mang a kelello a sebelisanang ka eona e se e le bohareng ba mehlala ea psychopathology. Mohlala, conceptualizations ea mathata a ho tšoenyeha khafetsa e tsepamisa maikutlo ho maikutlo a leeme a lebisang tlhokomelong e tšosang (Bishopo, 2007; Cisler & Koster, 2010; Ouimet, Gawronski le Dozois, 2009; Williams et al. 1996). Ka mokhoa o ts'oanang, liphoso tsa ho sebelisa taolo e phahameng ea maikutlo holim'a maikutlo li ntse li talingoa e le setsi sa mafu a kelello a hlahang ho tepelletseng maikutlo (Fales et al. 2008; Johnstone et al. 2007; Almeida et al. 2009; Taylor Tavares et al. 2008), ho moeli oa bofokoli ba botho (E ncha et al. 2008).
Libaka tsa pele li hlahella haholo mehlaleng ea methapo ea kutlo lipakeng tsa maikutlo le likarolo tse ling tsa kutloisiso. Leha ho le joalo, likarolo tsa anatomical tsa libaka tse fapaneng tsa pele pele li fuoa tlhokomelo e matla feela ha ho nahanoa bonnete ba mefuta e joalo. Ho isa tekanyong ea anatomy, hangata ho buisanoa feela ka maemo a pharaletseng a hore na sebaka se na le khokahano efe kapa efe e amanang ka kotloloho le libaka tsa maoto, joalo ka amygdala kapa hypothalamus. Leha ho le joalo, lintlha tsa likhokelo tsena li bohlokoa ho utloisisa ts'ebelisano ena ea tikoloho. Mohlala, mohlala o bontšang hore dorsolateral prefrontal cortex (DLPFC) ka kotloloho ts'ebetso ea amygdalar e ka utloahala feela haeba e bonts'oa hore DLPFC e romella likhakanyo tse lekaneng tsa amygdala. Haeba likhakanyo tse joalo li le bonolo kapa li le sieo, mehlala e meng e itšetlehileng ka libaka tse bohareng e tla hlokahala ho hlalosa tšusumetso ea DLPFC mabapi le likarabo tsa amygdalar.
Likarolo tsa libopeho tsa libaka tse fapaneng tsa pele le mokhoa oa laminar oa khokahano ea tsona li ka fana ka leseli le tebileng lipakeng tsa tšebelisano lipakeng tsa maikutlo le mekhoa ea kelello e kopantsoeng ke preortal cortex (PFC). Ka ho khetheha, likarolo tsa tikoloho ea libaka tse fapaneng tsa cortical li laola tsela eo li sebetsanang le tlhaiso-leseling ka eona le ho sebelisana le libaka tse ling. Boemo bona ba bobeli ba ho hlahlobisisa ha bo a kena lipuisanong tsa tšebelisano ea maikutlo le maikutlo, leha e na le litlamorao tse kholo bakeng sa ho utloisisa lits'ebetso tsena.
Tokomaneng ea hona joale, re leka ho bontša likarolo tsa puisano e fapaneng pakeng tsa libaka tse fapaneng tsa PFC, le litšebelisano tsa bona le amygdala. Re shebisisa haholo phapang pakeng tsa orbital le dorsolateral PFC ka lebaka la mekhatlo ea nako e telele ea orbitofrontal cortex (OFC) lits'ebetsong tsa maikutlo (Zald & Kim, 1996) le mokhatlo o tšoanang oa nako e telele oa DLPFC ho ea maemong a phahameng a hlokofatso (Fuster, 1989; Stuss & Benson, 1986). Re boetse re hlalosa karolo ea likarolo tsa karolo ea pele ea anterior cingulate (ACC) / medial frontal litšebelisanong tsena, kaha lintlha tse ntseng li eketseha li bontša hore meaho ena e fana ka khokahano e matla lipakeng tsa maikutlo le likarolo tse ling tsa ho utloisisa.
2. Togamaiso le likarolo tsa moetlo oa sebopeho sa PFC
Tebello ea litaba
Hangata PFC e arotsoe libaka tse pharalletseng tsa 6, dorsolateral, ventrolateral (VLPFC), frontopolar (FP), OFC, ventromedial (VMPFC), le dorsomedial (DMPFC) (bona Setšoantšo sa 1). Meeli e nepahetseng ea topographical ea libaka tsena e sebelisoa ka mokhoa o fapaneng ke bafuputsi, empa kakaretso ea kakaretso e ipakile e le sebopeho se hlophisehileng sa ho utloisisa sebopeho le tšebetso ea PFC.
Phylogeny le Cytoarchitecture
PFC e na le mekhoa e 'meli ea karohano e arohantsoeng, phylogenetly e arohaneng (Barbas, 1988; Sanides, 1969; Yeterian & Pandya, 1991). Mokhoa oa basoventral o atoloha ho tloha sebakeng sa polokelo ea mesebetsi ea tlhaho e fanang ka bokhabane ba mesebetsi ea tlhaho e fanang ka mokhoa oa "olgory" (")" ho phatlalla le OFC mme o hasana ka kotloloho ho poleiti e ka pele ea monyako, 'me hamorao ho VLPFC (e qetellang sebakeng sa Brodmann (BA) V46). Ho fapana le moo, mokhoa oa ho kenella lipakeng o qala ho latela Corpus collosum, e tsoelapele ka lebota la medial la lobe ea bokapele ebe e phutha moqomo o phahameng oa lobe ho DLPFC (e qetellang ka BA D46). E 'ngoe le e' ngoe ea mekhoa ena e bonts'a mekhahlelo ea tatellano ea meralo ea cortical e bonts'itsoeng ntlafatsong le katolosong ea granular layer IV. Karolo ea khale ka ho fetesisa ea mekhoa ena e agranular ka tlhaho, athe libaka tse nyane ka ho iphetola ha lintho li na le granular e boreleli ebile e hlalositsoe hantle. Ka ts'ebetso ea basoventral, tsoelo-pele ena ea cortical e qala ka kamoreng e ka morao ea OFC (ts'ebetso ea agranular e sebelisang poleloana ea Carmichael le Price (Carmichael & Theko, 1994)) e lateloa ke dysgranular (e fokolang granular) cortex libakeng tse bohareng ba OFC, e fetela ho eulaminate I cortex e nang le mela e ikhethileng ea granular IV ha e le ngoe e tsamaea ka ntle kapa ka morao, 'me qetellong e fihla eulaminate II cortex e nang le bophara ba IV le grragranular e matla. le dikarolo ha e le 'ngoe e leba nqeng e ka pele le polasetering ea libaka (Barbas & Pandya, 1989; Carmichael & Theko, 1994; Petrides & Mackey, 2006; Theko, 2006a). Tloaelo ea mekhoa ea bohanyetsi e bontša tsoelo-pele e ts'oanang ea ts'ebetso ea tikoloho. Ho qala ka periallocortex cortex e haufi le rostral corpus collosum, mokhoa o fetoha oa sethala ho cingate (ho kenyelletsa libaka tse haufi, tsa pele, le tse phahameng), eulaminate I ha e le 'ngoe e itsamaela kantle ho lebota la medical kapa ka mokhoa o fetisisang o kenella ka pele ho gyrus e ka pele,' me qetellong e fetoha e phahameng II libakeng tsa dorsolateral (BA 8 le 46).
Bakeng sa ho qoba pherekano, re hlokomela hore ts'ebeliso ea polelo ea methapo ea methapo ha ea lokela ho ferekanngoa le sebaka sa DMPFC se boletsoeng ho Setšoantšo sa 1. Mokhoa oa bohloeki o kenyelletsa DMPFC, empa o boetse o kenyelletsa libaka tsa VMPFC 25 le 32, le likarolo tsa BA 10 leboteng la medial (sebaka sa 10m ka nomenclature ea Ongur et al. (2003); Setšoantšo sa 2).
Sebopeho sa nts'etsopele ea cytoarological ha e le 'ngoe e tloha ho agranular ho ea ho eulaminate II cortex e tsamaisana le keketseho ea palo e felletseng ea li-neurons (boholo ba lisele), boholo ba lisele tsa pyramidal ho mela II le V, le boemo ba myelination (Barbas & Pandya, 1989; Dombrowski et al. 2001; Setšoantšo sa 3), tseo ka bonngoe li hlahisang litšobotsi tse fapaneng tsa ts'ebetso ea tlhahisoleseling libakeng tse fapaneng. Likhohlano tse ling tse kholo lipakeng tsa libaka tsa pele li hlaha ho latela maemo a nalane, hangata li bonts'a likarolo tse fapaneng tsa li-interneuron. Carmichael le Theko (Carmichael & Theko, 1994) arola Macaque OFC le medial PFC ka li-sub subions tse ngata ho latela likarolo tse joalo (bona Setšoantšo sa 4), mme tse ngata tsa likarolo tsena li ka bonoa ho batho (Ongur et al., 2003). Likarolo tse fapaneng tsa tšebeliso ea litaba li bonoa likarolong tse ka pele tsa pele li ama litšobotsi tse ikhethang tsa ts'ebetso ea tlhahisoleseling e fihlellelitsoeng ka sub suballel (Wang et al., 2004; Zald, 2007), empa li fetisa boholo ba pampiri ena. Ka mokhoa o makatsang, likarohano tse hlalositsoeng ka kotloloho tsa PFC li na le mekhoa e fapaneng ea khokahano kahare ho PFC le likarolo tse ling tsa boko le likotlo tse fapaneng.
Bohlale ba tikoloho ho Humans
Leha ho na le thuto ea mahlale ea bohlokoa tikolohong ea batho ba mehleng ea khale le tikoloho ea batho ka hare ho li-lobes tse ka pele, 'me mekhoa ea phylogenetic e arolelanoang mefuta eohle ea lintho tsa khale, ho hlaha mathata a' maloa pakeng tsa data ea motho le liphoofolo. Taba ea pele, lithuto tsa neuroimaging tsa batho hangata li bua ka libaka tsa Brodmann (Brodmann, 1914), empa ha li bonts'e tsoelo-pele e bonts'ang libaka tse amanang le cytoarological le meeli ea mabatooa e bileng teng ho tloha ha Brodmann a ntse a bula maliboho hoo e ka bang lilemong tse lekholo tse fetileng. Taba ea bobeli, ts'ebeliso ea li-labels tsa sebaka sena hangata e thehiloe ho Atlase ea Talairach (Talairach le Tournoux, 1988), empa atlas ena e haufi haholo, hobane tlhahlobo ea tikoloho ha e ea etsoa kelellong e theang motheo oa atlas. Taba ea boraro, ho na le ho se utloane pakeng tsa mabitso a liphoofolo le mabitso a batho kahare ea lebopo le ka pele, ka hore tlhaiso-leseling ea phoofolo e sebelisa lintho tse fapaneng moheleng e entsoeng ke Walker (Walker, 1940), eo bangoli ba bang ba seng ba e fetisitse ho batho (Petrides & Mackey, 2006; Ongur et al., 2003), ha bahlahlobisisi ba bangata ba neuro ba ntse ba sebelisa mabitso a Brodmann. Ka bomalimabe, ka linako tse ling ha ho hlake hore na bafuputsi ba tloaelo ea ho ngolla mabitso ba bua ka eng ha ba fana ka tlaleho ea seo ba se fumaneng. Sena se hlahisa ho hloka botsitso ho ikhethang ho OFC / VLPFC, moo bafuputsi ba batho ba atisang ho bua ka BA 47, empa lingoliloeng tsa liphoofolo li bua ka sebaka sa 12. Label 47 / 12 e se e amohetsoe ke litsebi tse ling tsa neuroanatom ho hlalosa sebaka sena ho batho, leha meeli ea medical ea sebaka sena e lula e ngangisana ka li-neuroanatomists tse etellang pele (Petrides & Mackey, 2006; Ongur et al., 2003). Ka mokhoa o ts'oanang, libaka tsa 13 le 14 li khethollotsoe ka ho hlakileng ho litšoene, 'me libaka tsa maiketsetso li bonoa ho batho, empa ha li nkuoe ke Brodmann kapa Talairach, ea sebelisitseng lebitso la sebaka sa 11 likarolo tse ka morao le tsa ka ntle tsa medical OFC. Ha re hlalosa datha tsa batho tse fanang ka maikutlo a ntlafatsang, ka kakaretso re bua ka mokhoa o pharaletseng oa ho ngola mabitso o hlalositsoeng ke Petrides le Mackey (2006), ho e-na le likhokahano tsa Talairach ho sebelisa monyetla oa tlhaiso-leseling ho tsoa lithutong tsa morao-rao tsa batho bao e seng batho.
3. Khokahano
Boholo ba data bo teng mabapi le likhokahanyo tsa pele bo tsoa liphuputsong tsa liphoofolo. Leha ho le joalo, ha ho fanoa ka thuto ea mahlale a amanang le cytarate ho pholletsa le li-primates (Petrides & Mackey, 2006; Ongur et al., 2003), ka kakaretso ho nahanoa hore khokahano ea libaka tsena e bolokiloe haholo-holo ka mefuta ea mefuta-futa. Kahoo, hoa utloahala ho sebelisa lingoliloeng tsa boleng bo holimo ho batho e le mokhoa oa ho lekola tšebeliso ea batho. Re shebisisa mefuta e 'meli ea khokahano mona: khokahano ea amygdala-PFC, le khokahano lipakeng tsa libaka tse fapaneng tsa PFC.
Tlhahiso ea Amygdalar ho PFC
OFC le PFC ea moriana e amohela litlatsetso tse ngata ho tsoa ho amygdala (Amaral et al., 1992; Carmichael & Theko, 1995; Barbas & Zikopoulos, 2006). Sena se fapana ka ho fapana le DLPFC, e fumanang merero e nyane e tobileng ho tsoa ho amygdala. Tlhahlobo ea lingoliloeng e bonts'a hore tse ling tsa likhakanyo tsa li-PFC tsa moea le tsa moea li fapana ho ea ka khokahano kapa subnucleus ea tšimolohoAmaral & Theko, 1984; Barbas & De, 1990; Amaral et al., 1992; Carmichael & Theko, 1995). Leha ho le joalo, lintlha tsena li ka nqane ho leqephe lena, 'me setšoantšo se matla sa khokahano se hlaha ka nqa e fapaneng ho tsebisa puisano e akaretsang ea mekhoa ea khokahano. Setšoantšo sa 5 e bonts'a merero e akaretsang ea merero ea amygdalar (e hlahisoang ke li-nuclei tse ngata tsa amygdalar) bokaholimo ba medial le ventral bokong ba macaque bo sebelisang lebitso la Carmichael le Price. Palo e bontša hore bokaholimo ba orbital ha bo ts'oanehe khokahanong ea eona ea kopanelo le amygdala. Habohlokoa ka ho khetheha ke ho ba sieo ha phepelo e kholo libakeng tsa 13m, 13l, 12m, 11l, le 10o holima orbital. Lerako la medial le boetse le fumana lipehelo tse matla tsa amygdalar, empa hape ha e ts'oane, kaha ha ho sebaka sa 10m kapa sebaka sa 9 ha se fumane tlhahiso ea bohlokoa ea amygdalar.
Ho ka etsoa liqeto tse peli tse tsoang mothating ona oa ho kenya maikutlo. Taba ea mantlha, karolelano ea amygalar ho PFC e hlophisitsoe ka tsela e hlophisehileng 'me e tsepame libakeng tse tsoetseng pele haholo ka tikoloho. Sena se bonts'a hore e tla ba phoso ho tšoara mofuta oa OFC kapa mofuta oa moruo ka mokhoa oa tlhaho joalokaha eka o hokahane le amygdala. Ho fapana le moo, tlhokomelo e mabapi le sebaka se kahare ho OFC le Puo ea Matsoho ea morafo e eletsoa pele ho ho hula maikutlo a mabapi le khokahano ea amygdalar. Taba ea bobeli, DLPFC le FP li amohela ho kenya letsoho ka mokhoa o fokolang oa amygdalar (ka mokhoa o hlakileng feela ke mekhoa e hlokolosi haholo e bonts'ang bopaki ba tlatsetso ea amygdalar). Ka lebaka leo, litšusumetso tsa amygdalar ho DLPFC le ts'ebetso ea FP li kanna tsa se tsamaellane, ekaba li fetisoa ka libaka tsa cingulate kapa tsa poso tsa OFC (kapa ka mekhoa e meng e akaretsang, joalo ka modula ea li-system tsa neurotransmitter).
Tlhahiso ea pele ho amygdala
Liphetho tsa PFC ho amygdala le tsona li hlophisitsoe ka tikoloho (Theko, 2006b; Ghashghaei et al., 2007; Stefanacci le Amaral, 2002; Stefanacci le Amaral, 2000). Ka kakaretso, libaka tse tlang pele tse amohelang likhakanyo ho tsoa ho amygdala li romella li-amygdala, ha libaka tse sa fumaneng tlatsetso e kholo ea amygdalar (joalo ka DLPFC le FP) li na le lipehelo tse fokolang tsa matla ho amygdala. Botebo ba likhakanyo bo bonts'a boholo ba cytoarchitectonics, le ho fokola ha methapo ea likhakanyo ha motho a tloha libakeng tse agranular ho ea ho isocortex e hlophisitsoeng ka sebopeho. Sebopeho sena se bonts'a libaka tsa isocortical (DLPFC le FP) li ke ke tsa fana ka tšusumetso e matla holima amygdala, mme ho fihlela moo li susumetsang amygdala, tšusumetso e kanna ea ba e sa tobang. Sena ha se bolele hore ha ho na likhakanyo tse tobileng tse tsoang ho DLPFC ho ea ho amygdala, ka ha liphuputso tse ngata li hlile li hlokometse likhakanyo tse tobileng ho tsoa sebakeng sa 9 le 46 (Stefanacci le Amaral, 2002; Aggleton et al., 1980; Amaral & Insausti, 1992). Ho fapana le hoo, likhakanyo li bobebe haholo ho fana ka molao o felletseng oa tšebetso ea amygdalar.
Le ha kakaretso ea cytoarchitectonics e fana ka tlhophiso e matla ea tlhophiso ho latela khokahano ea pele ho sebaka sa amygdala, phepelo ea potoloho ea liphetho le litholoana ha e tšoane.Ghashghaei et al., 2007). Haholoholo, phello e phahameng ka ho fetisisa ea amygdalar ho PFC e sebakeng sa agranular insularular haufi le poso ea OFC, athe sebaka se seholo ka ho fetisisa se hlahisoang ke amygdala se hlaha tikolohong ea morao-rao ea "cingulate" ea BA 25 ). Ka kakaretso, libaka tsa mabota a medial li na le tlhahiso e phahameng ho feta ho tsoa ho amygdala, athe libaka tsa kamorao ho OFC li na le tlhahiso e phahameng ho feta tlhahiso. Ho khahlisang ke hore libaka tsa morao-rao tse amanang haufi le li-PFC, libaka tsa DLPFC (BA 24, 8 le dorsal 9) li na le karolo e kholo ho tsoa ho tlhahiso ho amygdala, athe mohlala o khutlisetsoa kahare ho VLPFC. Tabeng ena, sebaka sa morao-rao sa sebaka sa 46l ka har'a VLPFC se fana ka likhakanyo tse itekanetseng ho amygdala, e leng sona feela sebaka sa morao-rao sa PFC se nang le tlatsetso ea bohlokoa ho amygdala. Ho joalo, likhakanyo tsena li matla ho feta se bonoang libakeng tsa anterior orbital, tse arolelanang likarolo tsa 12l e kholo ho feta tlhahiso, empa e bonts'a maemo a fokolang a khokahano ho feta sebakeng sa 12l.
Tse 'maloa tsa li-subnuclei tse fapaneng tsa amygdala li amohela ho kenya letsoho ha PFC. Basal and accessory basal and accessial nuclei e fumana lipallo tse matla haholo, hammoho le ho amohela likhakanyo tse tsoang ho libaka tse pharalletseng tsa libaka tsa PFC, athe li-nolo tsa motheo le tse bohareng li amohela lipehelo tsa PFC, empa ka tekanyo e tlase le e pharalletseng (Stefanacci le Amaral, 2002). BA 25 ea hlokomeleha hobane e sa romelle merero e boima haholo ho amygdala, e boetse e romella likhakanyo tse ngata tsa li-nuclei, kaha li-subnuklease tsohle tsa amygdala tse boletsoeng ka holimo li fumana ho tsoa ho BA 25. Leha e hlaha e le bobebe kholomo ea B ea Setšoantšo sa 6, ho bohlokoa ho hlokomela hore BA 32 e fana ka khakanyo e hlalositsoeng hantle ho amygdala. Ka litsela tse ngata BA32 e bonahala e le Homologous ho prelimbic cortex ka litoeba (Theko, 2006a). Ka litoeba, merero ea "cortex" ea prelimbic e ea likarolo tsa karolo ea basolatal le bohareng ba "amygdala"Vertes, 2004). Maemong a khale a se nang botho, likhakanyo li boetse li bonoe ho tloha BA 32 ho ea karolong e senotsoeng ea "basal nucleus" e fumanehangChiba et al., 2001). Ka hona, le ha e na le letsoalo le tlase haholo ho feta likhakanyo tse tsoang ho BA 25, BA 32 e hlaha e le boemong ba ho sebetsana le lits'ebetso tse khethiloeng tsa amygdalar.
Barbas le Zikopoulos (2006) ba pheha khang ea hore litlhahiso tsa pele tsa medial le tsa OFC ho amygdala li kanna tsa ba le litlamorao tse fapaneng mosebetsing oa amygdalar. BA25 ka lehlakoreng la medical e fana ka tlhahiso e matla haholo ea likarolo tse tlatselletsang tsa amygdala, tseo le tsona li fanang ka maikutlo a nyakallo ho hypothalamus. Ka lehlakoreng le leng, posterior agranular ea OFC e lula e le kahare ho maselese a kopaneng a pota-potileng bokahare ba basal (Bona. Feie. 7). Matšoele a kopaneng a fana ka tšitiso ea ho kenella hare khutleng e bohareng. Ha e hlohlelletsoa, maqhubu a kopaneng a emisa tsela ea thibelo ea tonic ho tloha sebakeng se bohareng ho isa ho hypothalamus, ka hona a baka tlhahiso ea hypothalamus. Litebello tse bobebe tsa thabo e kholo li boetse li fihla khubu e bohareng ka kotloloho ho tsoa mokokotlong oa OFC, e lumellang OFC hore e eketse ka bobeli kapa e fokotse ho thunya ho hoholo ha methapo.
Merero e tlang pele ho hypothalamus le brainstem
Libaka tsa OFC le li-PFC tse nang le maikutlo a pusong ea amygdala le tsona li hlahisa libaka tsa hypothalamus le autonomic brainstem / periaqueductal grey (Et al., 1998; Barbas et al., 2003; Theko, 2006b; Rempel-Clower & Barbas, 1998), ho fana ka bokhoni bo tobileng ba ho susumetsa libaka tsa boikemelo tsa autonomic tse amanang le tlhahiso ea maikutlo (bona Setšoantšo sa 8). Lintho tsena li bonahala li le matla haholo ho tsoa mehahong e meng ea lerako, empa hape li hlaha joalo ka sebaka se khubung ea orbital moo karolo ea amygdalar e leng kholo. Joalo ka ho haella ha eona ea ho fihlella amygdala ka kotloloho, DLPFC le FP ha li na lipalo tse tobileng libakeng tsena. Ho feta moo, likarolo tse ling tsa ka ntle tsa OFC li bonts'a tlhahiso e fokolang ho litsi tsena tsa tsamaiso.
Khokahano kahare ho lobe ea pele
Joalokaha ho boletsoe ka holimo PFC e ka aroloa ka mekhoa e 'meli e meholo ea phylogenetic. Likhokahanyo tse phahameng ka ho fetisisa sebakeng ka seng li na le libaka tse tšoanang, haholo libaka tsa boahisani tseo e seng karolo e le 'ngoe ea ntlafatso hole le sebaka seo ho buuoang ka sona. Ka hona, mohlala, libaka tsa "agranular insularular" tse ka morao ho OFC li na le khokahano e kholo ho libaka tse ling tsa agranular le dysgranular orbital, empa hangata ha li na khokahano le libaka tse se nang sebaka joalo ka 46 ea sebakeng sa heno. Moo likhokahanyo tsa maqhubu a pakeng tsa methapo e fapaneng li atang ha li tlohe maemong a fetang bonngoe a nts'etsopele ea meralo. Mohlala, sebaka sa isocortical ventral 46 se hokahane ka matla le isocortical dorsal sebakeng sa 46 mokhoeng oa methapo, empa ha e hokahane le libaka tse mpe tse tsoetseng pele tsa morako tse kang cgenrate ea subgenual (BA 46). Libaka tse eketsehileng tsa kahare le tsa morao tsa OFC li na le khokahano e kholo ea sebaka sa 25 sa ventral le sebaka se haufi sa 46, empa likhokelo tse hokahanyang sulcus e ka sehloohong karolong ea sebaka sa 45 ha li fumanehe hangata.
Leha ho le joalo, libaka tse 'maloa tsa OFC li bonahala li na le khokahano e tobileng le DLPFC. Libaka tse ikhethileng tsa 11m, 12o, 13a le 14r e na le likhokahano le DLPFC ka kotloloho. Gyrus rectus (e kenyeletsang sebaka sa 14r) e kanna ea talingoa e le karolo ea tloaelo ea lipakeng kapa e le sebaka sa phetoho lipakeng tsa mekhoa, ka hona likhokelo tsa eona ha li emele ho tlola ha maemo. Leha ho le joalo, libaka tsa 11m, 12o, le 13a li hlophisitsoe e le karolo ea mokhoa oa basoventral, ka hona, lihokela tsa tsona le DLPFC li emela likhokahano tsa mekhoa ea lipakeng. Ho utloisisa boemo bo boholo ba marang-rang a libaka tsena, ho bohlokoa ho hlahloba mokhoa o mong oa ho khetholla libaka tsa libaka tsa bophelo bo botle le tsa bongaka. Sebakeng sa mehlala ea phylogeny, Carmichael le Theko (1996) arola lebota la OFC le marangrang ho marangrang a marang-rang le a marang-rang a amanang le marang-rang a thehiloeng ka thata ho matla a khokahano lipakeng tsa libaka (Bona Feie. 9). Morero oa mofuta ona oa mekhahlelo o bonts'a pharalletsano e kholo pakeng tsa karohano ea phyolgenetic lipakeng tsa mekhoa ea basoventral le mediodorsal, e leng ho sa makatseng ha ho fanoa ka mokhatlo o hlophisitsoeng oa likhokahano tse seng li ntse li tšohloa. Leha ho le joalo, litsamaiso tse peli tsa karohano ha li tšoane ka botlalo. Ho khahlisang ke hore, libaka tsohle tsa orbital tse hokahanang le DLPFC ekaba karolo ea sebaka sa marang-rang sa Carmichael le Price, kapa se nkuoa e le mothusi lipakeng tsa marang-rang. Mohlala, sebaka sa 11m se nkuoa e le karolo ea sebaka sa marang-rang sa marang-rang, hobane se na le likhokahano tse kholo le libaka tsa lebota la medical ho fapana le tsa OFC kaofela. Carmichael le Price li khetholla libaka tsa 12o le 13a e le libaka tsa khokahano hobane li na le lihokela tse boima libakeng tsa medical le orbital. Mefuta ena e fapaneng ea khokahano e tiisa hore ho tla ba le matšoao a khethollo ea tikoloho, kapa esita le a sa lekanyetsoang, a bokhoni ba OFC ba ho sebelisana le libaka tse ling tsa pele. Haholo-holo, rectus ea gyrus, hammoho le 11m, 12o, le 13a li boemong ba ho ikopanya le libaka tse peli tsa mabota a medial (mohlala, cingulate) le libaka tsa DLPFC, athe libaka tse ling tsa Orbital ha li na kamano e tobileng ena.
Khokahano ea marang-rang ea marang-rang e laola tsela ho amygdala
Bakeng sa libaka tse haelloang ke tlhahiso e matla ea amygdala, bokhoni ba ho susumetsa ts'ebetso ea amygdalar bo tlameha ho its'etleha litseleng tse sa tsejoeng, 'me litsela tsena li tla laoloa haholo ke maemo a tsona ka har'a marang-rang a mantlha a pele. Ha ho fanoa ka matla a lipono tsa subgenual cingulate's (BA 25) ho amygdala, e ka fana ka thekiso ea bohlokoa ka ho khetheha eo likarolo tse fapaneng tsa PFC li susumetsang amygdala. Joalokaha ho ka bonoa ho Setšoantšo sa 9, BA 25 e fumana likhakanyo tse kholo tsa libaka tsa marang-rang tsa medial le libaka tse sebakeng sa orbital tse amanang le marang-rang a medial. Ho fapana le seo, likhakanyo tsa dorsolateral tse ngata li haella. Vogt le Pandya (1987) hlokomela hore BA 25 e amohela likhakanyo tse tsoang ho DLPFC, 'me u hlalose ka kotloloho tlatsetso e tsoang sebakeng sa 9 karolong e phahameng ea DLPFC. Leha ho le joalo, matla a khokahano ena a bonahala a fokola, 'me ha a so ka a bonoa hantle lithutong tse ling (Barbas & Pandya, 1989). Leha ho le joalo, BA 9 e hokahane hantle le BA 32 haufi le lebota la medical, leo ka lehlakoreng le leng le amanang haholo le BA 25 (Carmichael & Theko, 1996; Barbas & Pandya, 1989), mme ka hona e fana ka tsela e khonehang e sa sebetseng bakeng sa ts'ebetso ea DLPFC ho susumetsa BA 25. Ka mokhoa o ts'oanang, dorsal BA 46 ha e na khokahano e kholo le BA 25, mme e kanna ea tlameha ho kenya letsoho BA 32, kapa mohlomong likarolo tse ling tsa cingex cortex, ho buisana le BA 25.
Dorsal ACC (BA 24) e boetse e fana ka sebaka sa bohlokoa sa tlhahiso ho amygdala. Sebaka sena se na le litlatsetso tse ngata tse tsoang ho PFC (Carmichael & Theko, 1996; Vogt & Pandya, 1987; Barbas & Pandya, 1989). Sena se kenyelletsa kenyelletso e kholo ho tsoa ho BA 9, le ho isa tekanyong e tlase ea BA 46 ho DLPFC, likarolo tsa BA 32, le BA 10 leboteng la medical, le libaka tse 'maloa tsa OFC (haholo-holo libaka tsa marang-rang tsa methapo / pakeng tsa 13a le Iai, le 12o ). Kahoo, dorsal ACC e hlaha e le boemong bo matla haholo ba ho kopanya likarolo tsa ts'ebetso ea PFC libakeng tse ngata.
Leha libaka tsa anterior OFC le VLPFC li bonahala li na le keketseho e kholo ea tlhahiso ho amygdala ho feta tlhahiso e tsoang amygdala, hobane liporojeke tsena li fokola haholo, libaka tsena tse kantle ho naha li ka sebelisa litsela tse sa tobang ho kenella amygdala. Bakeng sa libaka tse ka thoko tsa OFC, sena se kanna sa lebisoa ho litereke tse ka morao tsa OFC. Ka lehlakoreng le leng, bakeng sa libaka tse kenang ka hare ho koloi, sebaka sa kamorao ho 12l se ka fana ka tsela e ts'oanang ea ho kenyelletsa amygdala, ha e fuoa maemo a eona a ikhethang har'a marang-rang a pele le a pele-a-amygdala.
4. Moetso oa metheo
Karolo ea tikoloho ea tikoloho ea cortical e susumetsa haholo tsela eo tikoloho e sebelisanang le likarolo tse ling tsa boko. Haholo-holo, boemo ba granularity le laminar nts'etsopele e ama boemo ba eona ba phepelo le tlhahiso ea maikutlo (Barbas & Rempel-Clower, 1997; Barbas 2000). Kahare ho mohlala o hlahisitsoeng ke Barbas, ho nahanoa ka mokhoa o hlakileng ho hlalosoa e le ho tloha meheleng e meholo le ho kenella ho likarolo tse tebileng tsa cortex. Ka lits'ebetso tsa maikutlo, methati ea pele ea ts'ebetso ea ts'ebetso e fana ka tlhahisoleseling ho methati e latelang ka mofuta ona oa projeke ea provo (Rockland & Pandya, 1979; Pandya, 1995). Ka har'a lits'ebetso tse nang le karolo e hlakileng ea phallo ea tlhaiso-leseling, joalo ka methapo ea kutlo, menyetla ea phepelo ea lijo le eona e ka hlalosoa e nyoloha ha e tloha sebakeng sa mantlha ho ea maemong a holimo mohloling oa ts'ebetso (mohlala V1 ho V2). Mabapi le lits'ebetso tsa kelello, ho fetisa tlhahisoleseling joalo ho lumellana le seo ba setso ba reng ke "theology" ke ts'ebetso e tlase.Kastner & Ungerleider, 2000).
Ka lehlakoreng le leng, menahano ea maikutlo e qala ka likarolo tse tebileng tsa cortex ebe projeke e ea holimo ho cortex (bona Setšoantšo sa 10). Ka lits'ebetso tsa methapo ea kutlo tse nang le sebopeho se hlakileng sa hierarchical, maikutlo ana a karohano a ka hlalosoa a theoha, ha a ntse a tsamaea ho tloha mehatong ea morao-rao ho isa nakong e fetileng ea molapo oa ts'ebetso ea sensory (mohlala, V2 ho V1). Maikemisetso a ho fana ka litlatsetso a sebetsa ho fetola kapa ho khetholla lipheo tse etsoang mehatong ea pejana ea ts'ebetso (Raizada le Grossberg, 2003). Mohlala, likhakanyo tsena tsa karabelo li thusa ho nyolla likarabo tsa lisele tse fumaneng lintho kapa libaka, ha li ntse li hatella kapa li hatella likarabo linthong tse sa lebelloang (Mehta et al., 2000; Saalmann et al., 2007). Maikutlo a joalo a thusa ho li-process tsa motheo tse hlakileng joalo ka khethollo ea lipalo (Domijan le Setic, 2008; Roland et al., 2006), hammoho le ho lumella taolo e tlase-tlase ea se sebetsitsoeng ho molapo oa tlhahisoleseling (Grossberg, 2007). Ka mantsoe a kelello, taolo ena e tlase-tlase e lumella ho feto-fetoha ha ts'ebetso ho latela litebello, lipheo tsa hona joale le tlhokomelo e lebisitsoeng (Glibert le Sigman 2007).
Bakeng sa ho hlaka, ho bohlokoa ho khetholla polelo ea ho nyoloha / ho theoha, ho fepa / ho fana ka maikutlo, le ho theoleha / holimo-tlase, hobane li bolela lintho tse fapaneng (bona Penny et al. 2004 bakeng sa puisano). Ho nyoloha le ho theoha ho nahanoa ho supa likarolo tse ikhethang tsa hierarchical, 'me polelo ea polelo e bohlokoa ka ho khetheha maemong a melatsoana e sebetsitsoeng hantle. Re sebelisa poleloana e ka tlase-holimo le tlase ho hloaea lits'ebetso tsa kelello, ka ho reroa ka boits'ebetso tse itlhahelang feela, joalo ka likarabo tse tsamaisoang ke maikutlo a ts'usumetso, le holimo ho bua ka mekhoa e lumellang. bakeng sa modular e feto-fetohang ea ho sebetsa ho kopane ka lipheo tsa hona joale le litebello. Mantsoe a felletseng le karabelo moelelong ona a na le moelelo o kopaneng, kaha o hlalosoa ke likarolo tse ikhethang tsa laminar, empa li boetse li fana ka maikutlo a likarolo tsa ts'ebetso ea tlhahisoleseling.
Hobane ho fana ka maikutlo le mehopolo e felletseng e hlalosoa ke likarolo tsa laminar, ho ka tsosoa kameho mabapi le mekhoa e ikhethang ea laminar e sebelisoang ke Barbas le basebetsi mmoho bakeng sa ho tšoaea merero e le feedxp kapa karabelo ka ntle ho melapo ea ts'ebetso ea maikutlo. Mefuteng ea sisteme ea pono, ho hlalosoa ka mokhoa o hlakileng ho hlalosoa ka mokhoa o ikhethileng mabapi le ho emisoa ha lits'ebetso tsa IV, ka litheko tse nyolohang tsa litheko tse phahameng le ho khaotsa ka mokokotlo oa IV (ho fapana le likarolo tse tebileng ka botlalo; Felleman & Van Essen, 1991). Ho fapana le moo, Barbas e sebelisa tlhaloso e pharalletseng e sa khetholle ka kotloloho lipakeng tsa mela ea IV le mela ea infragranular. Katoloso ena e holim 'a sebaka se loketseng ka ha ho na le libaka tsa pele tse sa na le likarolo tse matla tsa granular, le mekhoa ea ho felisoa ea laminar e hlokometsoeng libakeng tsena. Leha ho le joalo, lits'ebetso tse sebetsang tsa tlatsetso ena li lula li hlakisitsoe.
Ho hlaha bothata bo fapaneng hole le tlhaloso ea maikutlo a lebelo. Tlhaloso ea Barbas mabapi le karabo e shebisisa feela mehopolo ea ho tsoa lipakeng tse tebileng (infragranular) le ho khaola ka likarolo tse phahameng, tse lumellanang le mosebetsi oa mantlha oa Rockland le Pandya (1979). Leha ho le joalo, Felleman le Van Essen (1991) ba re tse ling tsa litlatsetso tse tlisoang ke ho theoha ha maikutlo li ka ba le tšimoloho ea 'mino ka tšebeliso ea tšimoloho ea litloholo le maemo a phahameng haholo. Ho tloha, Barbas o boloka litlhaloso tse ntlafatsang haholoanyane, ho amohela tlhaiso ea hae ho hlakile. Leha ho le joalo, e lebisa ho monyetla oa hore karolo ea menahano e khethiloeng e le karabelo ho PFC e ka ba e phahameng ka ho sebelisa tlhaloso e lokolohileng haholoanyane.
Taba ea bohlokoa ea sebopeho sa sebopeho ke hore boemo ba likarabo le lipakanyo tse felletseng lipakeng tsa libaka li khethoa haholo ke tekanyo ea ntlafatso ea tikoloho libakeng. Morero ho tloha cortex e arohaneng le ho feta (ke hore, sebopeho se pharaletseng haholoanyane, le sehokelo sa granular) ho ea cortex e sa hlophisehang hantle ea thekiso ea theknoloji e latela mokhoa oo oa phepelo, ha e tlohang boqapi bo tlase bo ntlafalitsoeng ho ea cortex e ntlafalitsoeng ka ho fetisisa ea tikoloho e latela mefuta e fapaneng ea karabo. Sebopeho sena se lumellana le se bonoang litsamaisong tsa maikutlo, empa sebopeho se bonahala se lumellana le lits'ebetso tse ngata. Ho PFC, sebopeho sa sebopeho se bolela esale pele tekanyo ea phepelo ea feedxp le karabelo hoo e ka bang 80% ea nako, ka tekanyo e lekanang ea karabelo le khokahano ea lijo li fetella haholo phapusing e kholo ea nts'etsopele ea cytoarural pakeng tsa libaka tse peli tse botsoang.Barbas & Rempel-Clower, 1997).
Sesebelisoa sa mantlha sa sebopeho sa sebopeho ho ea sehlooho se teng hona joale ke hore se lebisa ho liphatlalatso tse matla mabapi le mofuta oa puisano lipakeng tsa boko esita leha ho se na data e sebetsang ka kotloloho. Ho joalo, qetellong, li-electrophysiological kapa mekhoa e meng e khona ho lekola phallo ea tlhaiso-leseling ea laminar e tla hlokahala ho netefatsa hore lipaterone tsa mochini oa laminar ho PFC li sebetsa ka tsela e ts'oanang le se bonoang ho sensor cortices (ke hore, karabelo e hlalositsoeng ka mokhoa o hlophisitsoeng le projeketso e amanang le eona. thepa e tšoanang ea phallo ea tlhaiso-leseling ho sa tsotelehe mokhoa oo ho buuoang ka ona). Lithuto tse joalo tsa electrophysiological le tsona li ka thusa ho nchafatsa litekanyetso tsa ho hlalosa feedfoward le ho fana ka maikutlo. Khabareng, sebopeho sa sebopeho se fana ka motheo o matla oa boithuto ba hona joale o fumanehang bakeng sa ho bolela mofuta oa phallo ea tlhahisoleseling ho PFC. Haeba sebopeho sa sebopeho se nepile molemong oa tšebeliso ea tlhahisoleseling kahare ho PFC, e na le litlamorao tse kholo bakeng sa meetso ea tšebelisano-maikutlo.
Litaele tsa Laminar le likhokahanyo tse kenellang pele ho eona
E tsamaellana le sebopeho sa sebopeho, tlhahlobo ea mekhoa ea laminar ea likhakanyo e bonts'a hore dysgranular OFC e tšoauoa ka likarolo tse matla tsa ho fana ka maikutlo mabapi le khokahano ea eona le libaka tse ntlafalitsoeng ka moetlo oa PFC (Barbas, 2000). Ka ho tšoana le lits'ebetso tsa methapo ea kutlo, sena se ka bolela hore merero ea OFC e etselitsoe ho khetholla kapa ho fetola lipalo. Ho fapana le hoo, eulaminate DLPFC e na le litekanyetso tse phahameng haholo tsa "proxpidence" tse e lumellang, hore e fepe liphetho le liphumants'o tsa lipokello tsa eona ho libaka tsa boko tse latelang. Mokhoa ona o akaretsang oa ho fepa le ho fana ka maikutlo o boetse o tšoaea khokahano e itseng pakeng tsa OFC le DLPFC. Khokahano ea mantlha ea mantlha ho OFC e tsoa haholo likarolong tse kaholimo tsa cortical (2-3) mme li-axon tsa tsona li emisa haholo litheko tse tebileng (4-6), e tsamaellanang le mohlala o felletseng (Barbas & Rempel-Clower, 1997). Ka lehlakoreng le leng, merero ea OFC ho ea morao-rao ea PFC e simoloha haholo lithutong tse tebileng (5-6) ha li-axon tsa tsona li khaotsa haholo libakeng tse kaholimo (1-3), sebopeho sa karabelo. Mokhoa ona o bonahala o sebetsa ho 70-80% ea likhakanyo. Kahoo, tlhaiso-leseling e tsoang ho OFC ho ea ho PFC ea granular e na le maikutlo a mangata, athe tlhahisoleseling e eang ka lehlakoreng le leng e lumellana haholo le mohlala.
Moetso oa sebopeho o tsosa takatso ka hore o fana ka maikutlo a hore sebopeho sa puisano pakeng tsa libaka le naha se ka kenngoa ho ipapisitse le laminar kgokahanyo. Haeba sebopeho sa sebopeho se nepahetse, e re qobella ho sheba mofuta oa puisano le maikutlo mabapi le puisano ea libaka. Mefuta e bontšang hore PFC ea morao-rao e sebetsa haholo kapa ka boits'ebetso ba ts'ebetso ea mekhoa e holimo-tlase ho thata ho boelanya le likarolo tsa eona tse tsebahalang tsa phepelo. Ka mokhoa o ts'oanang, mefuta ea OFC e e bonang e fana feela ka sephetho sa sephetho (ka mohlala oa boleng ba moputso) ho PFC ea morao-rao, e sitoa ho nka bokhoni ba sebaka ka seng ba litakatso tse ts'oaroang ho PFC ea morao-rao. Leha ho le joalo, joalo ka ha ho hlalositsoe hamorao sengolong sena, mekhoa ea ho sebelisana lipakeng tsa PFC, le lipakeng tsa ho ameha maikutlo “maikutlong” khafetsa e hlokomoloha moelelo o ka bang teng molemong oa sebopeho. Ho joalo, mehlala e teng e tloaetse ho etsa mesebetsi ea morao-rao ea PFC, haholo mesebetsi ea DLPFC, mabapi le taolo e tlase-tlase, mme ha ba na taba le monyetla oa hore libaka tse sa ntlafalitsoeng joalo ka OFC li ka fana ka tšusumetso e matla libakeng tse latelang tsa PFC.
Meetso ea Laminar ea likhokahanyo tsa pele-amygdalar
Morero oa amygdala kamorao ho OFC o kenyelletsa likarolo tsohle tsa cortex, ka hona e kanna ea se keng ea lekanyetsoa feela ho ea provo kapa mofuta oa tlhahiso (Ghashghaei et al., 2007). Leha ho le joalo, ho hlakile hore ho na le karolo e matla ea phepelo ho merero ena e thehiloeng ho felisoa ha laminar. Ho fapana le seo, monahano oa OFC ho amygdala haholo o hlaha ho 5, o bonts'a sebopeho sa ona e le tlhahiso ea maikutlo (a fana ka maikutlo a hore ba nka khato ea ts'ebetso ea bias amygdalar ho fapana le ho fana ka tlhaiso-leseling e khethehileng joalo ka litšobotsi tsa maikutlo a susumetsang). Ho khahlisang ke hore, litebello tsa 'mele tse tsoang ho PFC ea morao-rao li lebisitsoe ho 5 ea OFC, e leng karolo ea mantlha ea lits'ebetso tsa OFC ho amygdala.
Na leseli la anatomical le ka thusa lipuisano tsa mesebetsi ea pele? Lilemo tse 'maloa tse fetileng li bone ho phatloha ho hoholo ka tsela eo likarolo tse fapaneng tsa boko li sebetsanang. Thahasello ena e hlahile ka lebaka la ho hlaha ha mekhoa ea ho lekola khokahano e sebetsang le fMRI, e fana ka lekhetlo la pele bokhoni ba ho hlahloba tšebelisano pakeng tsa likarolo tsa boko ho batho ba phetseng hantle. Leha ho le joalo, lipuisano tsa data tsena, le mehlala e hlahang ho datha tsena, ha li lula li qobelloa ke anatomy. Ha mefuta ena e ntse e e-ba le tšusumetso e kholo, re lumela hore ho bohlokoa ho lekola hore na e lumellana hantle hakae le neuroanatomy e hlalositsoeng ka holimo. Re lumela hore mefuta e joalo e hloka ho tsamaisana le mekhoa e tsebahalang e hokahanyang libaka tse fapaneng tsa cortical le subcortical le tlhaiso / mofuta oa phepelo ea lipaterone tsena. Ha meetso e sa lumellane le litšitiso tsena, ha e khone ho fana ka maikutlo, kapa bonyane li hloka tlhaloso ea hore na li ka sebetsa joang ka lebaka la ho se lumellane ha tsona le khokahano e tsebahalang ea boko.
Lingoliloeng tse ntseng li eketseha tsa kelello li leka ho utloisisa tsela eo "kelello ea kelello" e sebelisanang le "maikutlo" a eona. Leha ho na le meeli e hlakileng ea karohano ea maiketsetso lipakeng tsa ts'ebetso ea kelello le ea maikutlo (Pessoa, 2008), phapano e ipakile e na le thuso ho supa mefuta e fapaneng ea boits'oaro bo kang taolo ea maikutlo, tšusumetso, ho nka liqeto tsa moruo le tataiso ea mekhoa ea tlhokomelo. Likarolong tse latelang, re hlalosa datha le mehlala e hlahang bakeng sa molawana oa maikutlo, mohopolo oa ts'ebetso le litšebelisano tsa pele tsa dorsal-ventral, ka ho shebana le ho lumellana ha tsona le data ea anatomical. Re shebisisa haholo lingoliloeng tsa maikutlo, ha lingoliloeng tsena li ntse li eketseha lipuisanong tsa psychopathology le kalafo ea psychotherapeutic.
5. Taolo ea maikutlo
Tekanyetso ea maikutlo e hlalositsoe e le lits'ebetso tse amehang ho fetoleng qaleho, nako, botebo kapa dikahare tsa karabelo ea maikutlo (Gross, 1998; Gross, 2008). Lits'ebetso tsa taolo ea maikutlo li tloha liketsong tse nkuoeng nako e telele pele mohopolo o hlaha, joalo ka khetho ea maemo, lits'ebetsong tse kenyellelitsoeng pele kapa hang ha maikutlo a qalile ho hlaha, joalo ka ho tsamaisoa ka hloko kapa ho tsebisoa bocha (Gross, 1998). Ke mefuta ena ea hoqetela maano a ileng a etsa lipatlisiso mabapi le kamano lipakeng tsa libaka tse amanang le taolo ea kelello ea maikutlo le a amanang le karabelo ea maikutlo a khahlisang haholo. Lipatlisiso tsena li ka hlalosetsa ka mokhoa o hlakileng kapa ka mokhoa o hlakileng mokhoa oa ho laola maikutlo joalo ka ho romelloa ha libaka tse holimo-tlase, tse batang tsa "baridi" tse laolang libaka tsa PFC ho theola maemo a tlase, a chesang, a sebetsang a kenyeletsang likarolo tse tlase tsa maoto le matsoho tse kang amygdala. Ho hloleha ho atlehisoeng ha katleho ea methapo ea taolo ea bohlasoa ba PFC kapa mekhoa e mengata e tlase ea ts'ebetso ea amygdala ho hlahisitsoe ho kenya letsoho mefuteng e 'maloa ea psychopathology (Rottenberg le Gross, 2003; Rottenberg & Johnson, 2007).
Leano la taolo ea maikutlo le fumaneng tlhokomelo e kholo ho lingoliloeng tsa neuroimury ke ho hlahloba bocha. Leano lena la taolo le kenyelletsa tlhahiso-leseling e hlalosang maikutlo ka mokhoa o hlakileng ho fetola karabelo ea maikutlo (Gross, 1998). Reappraisal e akaretsa sehlopha se pharaletseng sa mekhoa e amanang. Mohlala, ho hlahloba hape ho ka tsepamisa mohopolo ho hlalosoeng ha moelelo oa ntho ea maikutlo ho e etsa hore e be ea bohlokoa kapa e sa itlhomphe. Ntle le moo, ho hlahlobisisa ho ka shebisisa ho hlakola sesosa, sephetho, kapa 'nete ea maikutlo a susumetsang ntle le ho fetola likamano tsa motho ho ea tšusumetsong. Mohlala, motho a ka khetha kotsi ea koloi lehlakoreng la tsela ka mohlomong a fella ka hore mokhahlelong kaofela o tlohele ketsahalo e bolokehile. Lithuto tse 'maloa tse sebetsang tsa neuroimaging joale li se li entsoe nakong ea mesebetsi ea ho lekola hape,' me e thathamisitsoe ho Lethathamo 1, le sebaka sa ts'ebetso ea PFC e bonts'itsoeng Setšoantšo sa 11. Ho sebelisa mantsoe a bohlokoa a tsamaiso mokhosi, ho sitisa le ho lekola botjha, lingoliloeng tse matlafatsang tse laolang maikutlo a boithatelo li kenyellelitsoe. Lithuto tsena tsa fMRI li ne li kenyelletsa tlhahlobo ea tlhaiso-leseling e laetsoeng, khatello ea maikutlo le lithuto tse sitisang batho ba bangata bao e seng litleliniki. Lethathamo lena la lithuto tsa taolo ea maikutlo ha le qete; mohlala, ha e kenyeletse mehopolo e amanang le mofuta oa molumo oa maikutlo. Rea hlokomela hore litafoleng tsohle re bolokile nomenclature (li-labels tsa Brodmann, kapa litlhaloso tsa setereke) tse sebelisitsoeng ke bangoli ba lipampiri tsa pele. Ho na le maemo a mang moo lipotso li ka hlahisoang mabapi le ts'ebeliso e khethehileng ea mabitso, empa re haelloa ke "standard standard" e hokahanyang libaka bakeng sa libaka tse tlang pele ho nako, ha re so ka re fetola mabitso, ntle le hore ho sengoloang re ela hloko ts'ebetso ea VLPFC tse lumellanang le karolo e ka morao ea BA 47 / 12. Ho haelloa ke demarcation e hlakileng ea karolo ea BA 47 / 12 e nang le khokahano ea bohlokoa ea amygdala ho batho, re nka karolo ea sebaka seo e leng posterior ho y = 32 ka kakaretso e emelang poso ea morao ea X 47 / 12. Re boetse re supa mangolong ha li-tataiso tsa OFC li lumellana le sebaka sa BA 13 (ho sa tsotelloe sebopeho sa bona sa mantlha).
Pherekano e atileng haholo ea ho ithuta ho bala bocha e botsa litho tsa sehlopha hore ba shebe li-valenced tse sa nepahalang haholo, tse tsosang takatso, litšoantšo tse tsitsitseng (mohlala, ho phumola, ho hlasela, ho bola le ho nyenyefatsa) le ho bapisa ts'ebetso ea neural nakong ea liteko tse hlahisitsoeng bakeng sa tlhahlobo e hlakileng ea kelello le liteko tse hlahisitsoeng bakeng sa ho shebella feela (Eippert et al., 2006; Kim et al., 2007; Ochsner et al., 2002; Ochsner et al., 2004; Phan et al., 2005; Urry et al., 2006; Van Reekum et al., 2007). Ha ho ntse ho e-na le phapang lintlheng tsa litaelo tsa ho hlahlobisisa ho tloha ho ithuteng ho ea ho ithuteng, ba hloka hore barupeluoa ba thehe tlhaloso e ncha ea moelelo, sesosa, sephetho kapa bohlokoa ba setšoantšo nakong ea liteko. Ho qhekelloa hape ha ho bapisoa le ho se shebelloe ha litšoantšo tse mpe ho nka libaka tse pharalletseng tsa PFC, ho kenyelletsa DLPFC le VLPFC (hangata e tlohelloa ka lehlakoreng le leng), le libaka tsa ACC tsa dorsal le / kapa tsa taolo ea PFC joalo ka ho ts'ehetsa likarolo tsa taolo ea boiphetetso. Setšoantšo sa 11 e bonts'a sebaka sa ts'ebetso e amanang le ts'ebetso e latelang (matšoao a cyan bakeng sa ho fokotsa ts'usumetso e mpe ea valenced, le 'mala o moputsoa bakeng sa ho fokotsa tsusumetso e ntle) ho tsoa lithutong tse boletsoeng ka holimo.
Paradigm e amanang le eona e sebelisa litšoantšo tse hlakileng tsa libaesekopo ho e-na le litšoantšo tsa tsona. Boithuto bona bo boetse bo bonts'a ho hira DLPFC ea linaha tse ling nakong ea ha motho a boela a hlahlojoa empa a fapana le hore na libaka tsa ACC le tsa PFC li ntse li hloa ho fokotsa ho hlonama, manyala kapa ho tsosa takatso ea thobalano (Beauregard et al., 2001; Goldin et al., 2008; Levesque et al., 2003, 2004).
Boithuto bo 'maloa ba boithuto bo sebelisang litšoantšo tse hlakileng kapa tse matla, litheko tsa amygdala li ile tsa sebelisoa e le projeke ea phetoho ea botle bo sa rateheng le ho nyakalla ha mmoho le ho fokotseha ha ho haptjoa hoa mekhoa e metle lithutong tse ling (Goldin et al., 2008; Levesque et al., 2003; Ochsner et al., 2002; 2004; Phan et al., 2005). Rea hlokomela hore ho leka-lekanya ho lekanang ha mesebetsi ea amygdalar ka phello e mpe ho na le mathata, ha re fuoa hore 1) li-amygdala li ba maemong maemong a seng a mabe, mme 2) liphihlelo tse mpe tse amanang le litšebelisano li kenyelletsa likarolo tsa cortical le subcortical tse eketsang ka nqane ho amygdala. Leha ho le joalo, ka lebaka la thahasello ea rona lipuisanong tsa boko ba tikoloho, taolo ea tlase ea tšebetso ea amygdalar e fana ka index e sebetsang bakeng sa ho lekanya tšebelisano-leseling ea maoto le maqhubu ho sa tsotelehe hore na ts'ebetso ea eona e tsamaisana le efe e amehang. Boholo ba lithuto li fumana ho fokotseha ho amygdala e setseng, le khafetsa li-amygdalae, ha li sebelisa boiphumano ho theola taolo e mpe. Ke lithuto tse 'maloa feela tse ileng tsa hlahloba ho hlahlojoa hape ha maikutlo a khothalelitsoeng. Ha motho a botsoa hore a hlahlobe bocha kapa a theole melaoana e khothatsang kapa e tsosang takatso ea thobalano, boemo ba ts'ebetso ea amygdala le letona bo fokotsehile (Beauregard et al., 2001; Kim le Hamann, 2007). Sena se ka phahamisa mohopolo mabapi le semelo sa morao-rao sa maikutlo, empa ka kakaretso, lithuto tse hlahlojoang bakeng sa litšebelisano tse hlophisitsoeng le amygdala latous li haelloa.
Leano le leng la taolo ea maikutlo le kenyelletsa ho hopotsa litšoantšo tse ntle kapa tse thobang ebang ke tsa tlhaho kapa tsa nakong e fetileng ho nka sebaka kapa ho loants'a tse mpe. Liteko tsa boits'oaro li bonts'a hore ho hopola menahano e sa lokang kapa litšoantšo tse fokotsehang ho ama tse mpe (Erber le Erber, 1994; Joormann, Seimer le Gotlib, 2007; Parrott & Sabini, 1990; Rusting & DeHart, 2000). Lithuto tse peli tse entsoeng ka neano tse holimo li ile tsa bapisa ho laola motho ka ho hopola setšoantšo se khutsitseng kapa mohopolo ka tebello e sa laoloang ea ho makala. Kalisch le basebetsi-'moho (2005) o ile a letsetsa liteko ka lithane tse bonts'a hore na ho na le menyetla ea tšabo litekong tseo kapa che. Litekong tsa molawana, barupeluoa ba ile ba laeloa hore ba tlohele maikutlo a bona a matšoenyeho mme ba nahane ka sebaka se ikhethileng se khethiloeng pejana. Litekong tseo e seng tsa molao, barupeluoa ba laetsoe ho etsa likarabo tsa bona tsa maikutlo. Boithuto ba ROI bo bonts'itse hore mofuta ona oa taolo o hira sebaka sa right anterolateral frontal cortex (MNI: 42, 48, 18) le regulation moo ho nang le matšoenyeho a hapileng libaka tsa mediti PFC le rostral ACC (−4, 46, 28). Thutong e tšoanang, Delgado le basebetsi mmoho (2008b) o sebelisitse lithunya tse mebala ho khetha liteko tseo ho ka etsahalang hore a tsitsinyehe, 'me a kopa barupeluoa ho laola matšoenyeho a bona ka ho hopola sebaka se le seng sa tlhaho se khethiloeng tlhahisong. Tlhahlobo ea bona ea ROI e bonts'a hore ba hopotsa litšoantšo tsa tlholeho ha ba lebella ho tsitsipana ba bokella karolo ea letsoho le letšehali la pele (Talairach: −43, 28, 30). Bokapele ba eona bo neng bo amana le katleho ea melaoana. Taolo e boetse e hlahisitse ts'ebetso leboteng la ventral medial le cingrate ea subgenual (BA 32; −3 36, −8 le BA 25; 0, 14, −11), eo bangoli ba e supang e amana le ho timela (Phelps et al., 2004) le ho fokotseha hoa ts'ebetso ea amygdalar e setseng. Le ha lithuto tsena ka bobeli li sebelisa maemo a tšoanang, mekhoa ea bona ea ho sekaseka ho kenyelletsa khetho ea li-ROI le mokhoa oa ho bapisa litla-morao tsa tonic le phasic e kanna ea ikarabella bakeng sa liphapang tse teng literekeng tse tlalehiloeng bakeng sa ho taka litšoantšo tse ntle kapa tse thobang ho loants'a matšoenyeho a amanang le ho emela ho ka etsahala. ho thothomela.
E ts'oanang le leano la pele la taolo ea maikutlo, ho sitisoa ho kenyelletsa ho boloka leseli le sa nepahalang le mohopolong oa motho ea sebetsang. Patlisiso ea boitšoaro e bontša hore ho etsa joalo ho fokotsa likotsi tse mpe bathong ba dysphoric le ba nondysphoric (Fennell, Teasdale, Jones, & Damle, 1987; Lyubomirsky, Caldwell, & Nolen-Hoeksema, 1998; Teasdale & Rezin, 1978). Ka ho nka matla a memori a sebetsang le likhopolo tse sa lokang tsa mehopolo, menahano e metle ea moea e thibetsoeng ho fihlella lisebelisoa tsa tlhokomelo (Minaer, 2005). Lithuto tse feto-fetohang tsa tšitiso li sebelisitse lipapatso tse peli tse fapaneng. Ea pele, e hiriloeng ke Kalisch et al. (2006), o sebelisitse tebello ea tšohanyetso ea tšabo, ntle le hore moemeli a ka hopola mohopolo o monate kapa o bolokehileng, ho bile le taelo e bulehileng eo morupeluoa a khothaletsoang ho nahana ka eng kapa eng ntle le ho tsitsipana. Paradigm ena e bontšitse sebaka sa PFC e setseng (MNI: −56, 30, 22) e neng e le mafolofolo haholoanyane litekong tseo barupeluoa ba neng ba laetsoe hore ba itšilafatse ho feta litekong tse sa sitiseng letho. Phapang ea bobeli ea tšitiso e kenyelletsa mosebetsi o itseng oa ho khelosa (Sternberg memory memory) moo morupeluoa a tšoereng letoto la mangolo a memori ea ts'ebetso ha a ntse a shebile litšoantšo tse mpe kapa tse sa nke lehlakore ebe o latela setšoantšo se ts'oanang o tlameha ho arabela hore na lengolo le le leng le ne le le ho sete eo ba neng ba e hopola. McRae et al. (2009) e tlaleha hore ho etsa mosebetsi oa ho hopola ha o ntse o shebella lits'oants'o tse mpe ha ho bapisoa le ho shebella feela ho eketsa karabelo ea BOLD ka letsohong le letšehali le ka lehlakoreng le ka pele la bo-gyri (MNI: BA6; −6, 10, 62 le −56, −4 , 48 le 48, 42, 32; BA 9; −42, 22, 30 le 42, 30, 34; BA 10; −36, 62, 12 le 38 and 64 and 14 47p; 12, 36, −20).
Litlaleho tse ngata tse fanang ka maikutlo mabapi le taolo ea maikutlo li hlahisa ka ho hlaka libaka tsa DLPFC e le karolo ea taolo e itseng ebile li seli ka ho re likarabo tsa nako e tšoanang tsa likarabo tsa amygdala li hokahana ka kotloloho le amygdala. Tabeng ea ho lekola bocha hape le litšitiso, tlhokomeliso ena e bohlokoa ka ho khetheha hobane ts'ebetso tsena li hlahisa sepheo se fetisoang ho PFC (Setšoantšo sa 11). Joalokaha ho boletsoe pejana, sebopeho sa lipalo tsa anatomical tse tsoang cortex li bontša hore litsela tse otlolohileng tse tsoang libakeng tsa DLPFC ha li na monyetla oa ho fana ka taolo e matla ea ts'ebetsong ea amygdala. Libaka tsa PFC tse nang le menahano e metenya ka morao ho PFC li fumaneha feela karolong e nyane ea VLPFC, haholo-holo libakeng tse kamorao ho BA 47 / 12. Ka bomalimabe, joalo ka ha ho boletsoe pejana, lebitso le sebelisitsoeng ho tlaleha ts'ebetso sebakeng sena lithutong tse ngata li baka ho hloka botsitso ha ho tluoa mabapi le lipotso tsa khokahano le amygdala. Boithuto ba boithuto bocha, ho hopola ka nepo kapa ho nka karolo ha litšoantšo le tšitiso hangata li tlaleha ts'ebetso libakeng tse akaretsang tsa VLPFC le medial OFC (Eippert et al., 2007; Goldin et al., 2008; Kim le Hamann, 2007; Lieberman et al., 2007; McRae et al., 2009; Ochsner, Ray et al., 2004). Ka ho khetheha, lithuto tse ngata tsa boipatlisiso li tlaleha ts'ebetso ea naha e 'ngoe ea BA 47 / 12 ha e fokotseha maikutlo a mabe kapa a nepahetseng. Joalokaha ho boletsoe kaholimo, BA 47 / 12 ke sebaka se seholo ebile se na le mathata ebile ke libaka tse ka morao tsa BA 47 / 12 ke libaka tsa lipalo tsa bohlokoa tsa amygdalar. Ka hona, lipolelo tse matla mabapi le tšusumetso e tobileng ea kelello ho amygdala e utloahala haholoanyane lithutong tseo ka ts'ebetso karolong ena e ikhethang ea BA 47 / 12.
Libaka tsa kalafo tsa PFC hangata li nkuoa e le monyetla o motle oa ho fihlella libakeng tse tlase tse joalo ka amygdala. Leha ho le joalo, ho ea ka sehokelo sa malk se kopantsoeng ka ho toba le amygdala, ke libaka tseo feela tse nang le kamano e ntle ea methapo (BA 25) le dorsal ACC (BA 24) tse nang le khokahanyo e tobileng le amygdala. Ke lithuto feela ka Delgado le basebetsi mmoho (2008a, 2008b) tlaleha ho tsepamisa maikutlo holim'a sebopeho sa medical se literekeng tse nang le tšusumetso e kholo ho amygdala. Ha re fuoa lintlha tsa anatomical, ho ka utloahala ho makatsa hore ts'ebetso ea BA25 ha e hlahe khafetsa lithutong tsena. Leha ho le joalo, ho a utloahala hore letšoao la ho theoha setulong se ka morao ho VMPFC le thibetse lithuto ho bonts'a ts'ebetso e ts'oanang ea tikoloho ena. Khafetsa, lithuto tsa inhibition / khatello, tharollo le tlhaiso-leseling li tlaleha feela tse hlahang ho BA 32, tse ka bonts'ang phetoho e hlakileng ea amygdala, ha e fuoa mofuta o potolohileng oa tlatsetso ea BA 32 ho amygdala.
Lithuto tsa moea mabapi le ho qaptjoa hoa amygdala
Bakeng sa ho utloisisa ka botlalo haholoanyane hore na melaoana ea maikutlo a holimo-le-moea e sebelisana joang le amygdala, lithuto tsa melaoana ea maikutlo li fetile haholo ho feta karohano ea mesebetsi le taolo ho fuputsa likhakanyo tse ikhethang tsa ho fokotseha hoa tšebetso ea amygdala (Bona. Lethathamo 2). Ka mantsoe a mang, ho fapana le ho botsa hore na ho na le likarolo life tse etsang mosebetsi o tsejoang ka ho laola tšebetso ea amygdala, ba ile ba leka ka nepo hore na ho na le khokahano pakeng tsa amygdala le kelello eohle nakong ea ts'ebetso ea taolo ea maikutlo. Ntle le moo, lithuto tse ling tse amanang le amygdala lia fokotseha ka libaka tse seng li tsebahalitsoe pele ho lipapatso tse ka sehloohong tsa molao. Lithuto tsena li bonts'a hore litheko tsa amygdalar li amana hampe le libaka tse ngata tsa ts'ebetso ea PFC. Ntho ea bohlokoa ke ts'ebetso ho VMPFC, ho kenyelletsa BA 11m / 14r (5, 37, −12; −6, 46, −20: Urry et al., 2006, Ochsner et al., 2002 ka ho latellana). Ntle le moo, libaka tse tlolang pele ho nako le tsa tlhaho li ile tsa bonoa li sa tsamaellane hantle le mosebetsi oa amygdala nakong ea taolo. Mohlala, Urry le basebetsi-'moho (2006) e tlaleha tikoloho ea BA 32 / 10 (e phahameng ho −23, 43, −10) e ileng ea atoloha ka moea le kahare. Delgado et al. (2008b) hape o tlaleha kamano e fapaneng pakeng tsa mosebetsi oa BA 32 (0, 35, N8) le amygdala e fokotseha. Libaka tsa Posterior (BA 13) tsa OFC le tsona li amana hampe le amygdala deactivation (−24, 28, −14; 26, 24, −22: Banks et al, 2007: −30, 22, X16, 34, x 24: Ochsner, Ray et al., 2004). Libaka tse tlase tsa PFC ho BA 47 (34, 54, 12) le BA46 (−54, 12, 12: Urry et al., 2006; −54, 42, 12: Ochsner et al., 2002), hape e hlahile lithutong tsena. Boithuto bo habeli bo hokahanya likarolo tse ikhethileng tsa DLPFC le libaka tsa medial, tse neng li tsamaisana le ho fokotseha ha karabelo ea amygdala. Thutong ka Urry et al. (2006), tlhahlobo ea lipakeng e bontšitse khokahano lipakeng tsa amygdala, BA 10 (3, 63, 18) le sebaka sa DLPFC (−50, 23, 19). Delgado et al. (2008b) ka mokhoa o fapaneng o sebelisitse sebaka sa medial BA 32 e le peo bakeng sa tlhahlobo ea bona ea PPI e ileng ea supa sebaka sa amygdala le setereke sa DLPFC. Habohlokoa, liphuputso tsena li supa libaka tse tsamaisanang le phapang ea amygdala tseo le tsona li boletsoeng kaholimo molemong oa projeke ea amygdala joalo ka dorsal anterior cingulate, cgenergen ea morao-rao le cortex ea morao-rao.
Likarolong tse tlalehiloeng ho tsoa ho litekolo tsena tsa tlhahlobo kapa tlhahlobo ea khatello ea methapo, palo e lekanyelitsoeng ea tsona e na le khokahano e tobileng ea amygdala. Libaka tse atileng haholo tse sa tsamaellanang hantle le karabelo ea amygdala ke libaka tsa kamora bokhabane ba OFC le cingmbre e ikhethileng le VLPFC (Setšoantšo sa 12). Likarolong tse ka pele tsa pele feela karolo e ka morao ea BA 47 / 12 e na le lipalo tse matla ho amygdala. Mabatooa a anterior BA 32 a boetse a bonoa tlhahlobisong ea bohlophisi, a ka bonts'ang likhakanyo ho "testal" le basal lateral nucleus ea amygdala (Cheba et al, 2001).
Mehlala ea melao ea tsamaiso
Ho fihla joale, mohlala o hlophisitsoeng haholo oa tsamaiso ea maikutlo o tsoa thutong ea ho lekola botjha ke Wager le basebetsi-'moho (2008). Phapang ea lithahasello ke phetoho ho tse amehang hampe tse mpe. Mokhoa o kopanetsoeng oa sebopeho o sebelisitsoe ho datha e nang le neuroimaging ho tsoa paradigmiti ea boithuto e ts'oanang le e sebelisitsoeng ke Ochsner et al. (2002; 2004). VLPFC e nepahetseng e ile ea khethoa e le sebaka sa ho qala sa tlhahlobo, 'me e hokahane le lihokela sebakeng se kenyelletsang karolo ea bokantle ba sebaka sa 47 / 12 e reriloeng ho amygdala. Bangoli ba qalile ba sebelisa mokhoa oa ROI ho lekola karolo ea amygdala le li-nucleus accumbens e le babuelli pakeng tsa VLPFC e nepahetseng mme ba fokotse tšusumetso e mpe e ileng ea tsebahala e le metriki ea mantlha ea katleho ea katleho. Thutong ena ea ROI lits'ebetso tsena ka bobeli li bonts'itsoe ho loants'a likamano lipakeng tsa VLPFC le tokelo ea ho its'oara e theohile ka tšusumetso e mpe (bona Setšoantšo sa 13).
Bangoli ba ile ba sebelisa tlhahlobo ea kholo ea kholo ea kelello le tšusumetso ea maikutlo ho khetholla marang-rang a mabeli ka moo ho ka khonehang babuelli ba kamano pakeng tsa VLPFC le liphetoho tse tlalehiloeng tse mpe tse amehang (bona Setšoantšo sa 14). Khokahano e le 'ngoe e na le leeme le sa tobang mabapi le ho eketsa phetoho liphatsong tse mpe. Marang-rang ana a kenyelletsa likarolo tsa li-nucleus accumbens, subgenual cingulate (BA 25), SMA ea morao-rao, precuneus, DMPFC (MNI: 24, 41, 40), le "front" ea "gy" (24, 21, 58). Har'a libaka tsena, li-nucleus tse bokellanang le li-cingrate tse tlisoang ke tlhaho li na le kamano e matla haholo le amygdala. Khokahano ea bobeli e khethiloeng e na le leeme le sa tobang ho fokotsa phetoho e mpe le ho fokotsa katleho. Marang-rang ana a kenyelletsa rostral dorsal ACC, amygdala (bilatal) le posterior-lateral OFC (48, 24, −18). Mosebetsi oa nako e tlang o tla tlameha ho hlakisa hore na likarolo tsa li-network li sebelisana joang le hore na likhokahano tsena li ikhethile ho mofuta ona oa leano la taolo ea maikutlo.
Bafuputsi ba bangata ba fane ka mehlala ea maikutlo mabapi le mekhoa ea neural e tsamaisang maikutlo. Tse bonolo ka ho fetisisa tsa mefuta ena li fana ka maikutlo a hore libaka tse fokolang li na le tšusumetso e tobileng ho amygdala. Delgado et al. (2008b), Hansel le von Kanel (2008) 'me Quirk le Beer (2006) tlhahiso e 'ngoe le e' ngoe e bonts'ang hore PFC e tsoang ka tlase e laola libaka tsa amygdala. Mefuta ena e bohlokoa e leka ho theha kutloisiso ea rona ea melaoana ea maikutlo a motho ka har'a lingoliloeng tsa liphoofolo tse pharalletseng le khokahano ea PFC e kopaneng le bongata bo kopaneng bo-basy ba ma-basy.Morgan, Baroma le LeDoux, 1993; Likhtik et al., 2005; Quirk et al., 2000). Quirk le Beer (2006) haha ka boteng ba litlamorao tse nyarosang le tse thibelang maikutlo tsa "ventral" ea maikutlo a PFC ho amygdala ho batho le likhoto. Sebaka sa subingual cingrate, BA 25, ho tsekisanoa hore se sitisa le ho feta ha li-dorsal le anterior BA 32 li khothalletsoa ho ba le khokahano e khahlisang le amygdala. Ka bobeli BA 25 le 32 li na le khokahano le amygdala. BA 32, leha ho le joalo, e na le likhokahanyo tse fokolang haholo.
Phillips et al (2008) ba hlahisitse mofuta oa potoloho o lekang ho hlalosa ho hlaka hoa neural ha mefuta e mengata ea taolo ea maikutlo (bona Setšoantšo sa 15). Moetso o na le likarolo tsa DLPFC, OFC, VLPFC, DMPFC le ACC. Ho khahlisang le ho feta, bangoli ba khetholla lipakeng tsa likarolo tse amehang taolong ea maikutlo e ikemetseng (ho subgenual and rostral ACC) le libaka tse hiretsoeng melao ea boithaopo ea boithaopo (DLPFC le VLPFC). Li tšoaea libaka tsena tsa ho qetela e le tse ncha tsa phylogenetically le ho fana ka maikutlo lits'ebetsong tsa maikutlo a khale tsa maikutlo. TheCC, DMPFC le ACC, ka lehlakoreng le leng ke libaka tsa khale tsa phylogenetically tse hlalosoang li sebetsa ka lits'ebetso tsa phepiso ea ho fetisetsa tlhahisoleseling ea kahare ho DLPFC le VLPFC. Bangoli ba beha DMPFC e le eona mosola oo OFC e bolokang tlhahisoleseling ea bohlokoa ho ea libakeng tse sa sebetseng tsa bokong bakeng sa ho etsa liqeto.
Karolo e 'ngoe e ikhethang mohlaleng ona ke ho fana ka litlhaloso tse hlakileng tsa mekhoa ea phepelo le maikutlo. Moetso ona o hohela ka mokhoa o makatsang ebile o lumellana hantle le maikutlo a setso mabapi le DLPFC e nang le taolo ea taolo e tlase holim'a libaka tse "nang le maikutlo" haholo. Leha ho le joalo, ho thata ho lumellanya conceptualization ena le sebopeho sa sebopeho, ka lebaka la kabo ea laminar ea khokahano ea PFC (Barbas & Rempel-Clower, 1997; Barbas, 2000). Ho joalo, mohlala oa sebopeho o fana ka maikutlo a hore tlhahisoleseling e lipakeng tsa DLPFC le OFC ehlile e ka lehlakoreng le fapaneng le lits'ebetso tse simolohang ho OFC le ho ea DLPFC e tšoauoa haholo joalo ka maikutlo, le ba hlahang ho DLPFC le ho ea OFC boholo e tšoauoa ka ho fepa.
Phillips et al. mohlala o boetse oa bohlokoa ha o beoa libaka tse bitsoang "othomathiki tsa tsamaiso" joalo ka setereke se ikatisang le OFC joalo ka tsela ea mantlha eo libaka tse ncha tsa phylogenetically li amang libaka tse matla joalo ka amygdala. Sena se lumellana haholo (haholo tikoloho e ikhethileng ea cingulate) le litokisetso tsa netweke tse hlalositsoeng kaholimo. Ho ka nahanoa, leha ho le joalo, hore ho ka ba le tsela e fetang e le 'ngoe eo libaka tse laolang maikutlo a boithaopo li ka amang ts'ebetso ea amygdala. Haholo-holo, VLPFC ea ka morao e ka khona ho ama ka kotloloho lits'ebetso tsa amygdala ntle le hore e hloke ho kenella ho e 'ngoe ea libaka tsa "medical regulation" tse fanoang ka kotloloho, ka lebaka la mats'oaetso a eona a tobileng ho amygdala nuclei.
Ka kakaretso, letlotlo le lengata la data le bonts'a ho kenella ha libaka tsa PFC nakong ea melao ea tsamaiso ea maikutlo, le ts'ebetso sehlopheng se khethiloeng haholoanyane sa libaka (BA 47 / 12, BA25 le BA 32) tse bonts'ang mekhatlo e nang le bokhoni ba ho laola liketso tsa amygdala. Ho 'nile ha etsoa tlhahiso ea meetso e rarahaneng ea ho hlalosa lintlha tsena. Ho hlaha ha mefuta ena hoa ipiletsa, joalo ka ha ho na le ngongoreho e bonts'itsoeng ke bangoli ba bona mabapi le ts'ebetso ea likhokahano tse kopaneng. Rea hlokomela, leha ho le joalo, hore ha ho na mehlala ho fihlela joale e amohetseng ka ho hlakileng mokhoa oa likhokahanyo pakeng tsa libaka tse fapaneng tsa PFC. Mohlala, Wager et al (2008) e fana ka mohlala o rarahaneng ka ho fetisisa bakeng sa moralo o itseng oa taolo ea maikutlo, empa ha o bue ka mofuta oa phallo ea tlhahisoleseling lipakeng tsa likarolo tsa karolo. Phillips et al. kenya ka ho hlaka mohopolo oa tlhahiso ea tlhahiso le litlhaloso, empa u se ke oa hokahanya maikutlo ana le mokhoa o bonts'itsoeng oa ho fana ka maikutlo le lipakanyo tse latellanang libakeng tse amehang. Re lumela hore ho boelanya litaba tsena ho fana ka a mang a mathata a bohlokoa bakeng sa bafuputsi ba lekang ho utloisisa likarolo tse tlase tsa maikutlo.
6. Taolo ea kelello ea ho sitisoa ha maikutlo
Ha boholo ba tlhahlobo ea rona bo shebile lithutong tsa taolo ea maikutlo, ho hlaha litaba tse ngata tse tšoanang ha ho hlahlojoa lingoliloeng ka taolo ea kelello. Ka botlalo, taolo ea kelello e bua ka lits'ebetso tsa maemo a phahameng tse ntlafatsang ts'ebetso e loketseng, ha e ntse e thibela ts'ebetso e sa sebetseng. Polelo ena e sebelisoa haholo ho sebetsa mesebetsing e hlokang tlhokomelo e hlokoang ea tlhaiso-leseling e nepahetseng ea sepheo le ho thiba lintlha tsa maikutlo tse sa utloahaleng, le khetho e tsamaeang le likarabo tse khothatsang tsa sepheo le khatello ea likarabo tse sa hlolleng sepheo. Ts'ebetso e joalo ea khetho hangata e hlahisoa ka mokhoa o hlakileng ho latela maemo a holimo le tlase le ho khetha mekhoa e metle. Lithuto tsa taolo ea maikutlo le maikutlo li ka talingoa e le karoloana e ikhethileng ea taolo ea kelello e shebileng ho fetolela karabelo e amanang le eona. Ho fapana le hoo, mefuta e meng e mengata ea lithuto tsa taolo ea kelello e kenyelletsang maikutlo ho matla a ho hlola litšitiso tse bakoang ke ho susumetsoa ke maikutlo. Ka lebaka la litšobotsi tsa bona tsa tlhaho (hangata tse ikatisang) tse hapang tlhokomelo (Boholo et al., 2005; 2007; Pessoa, 2008), maikutlo a susumetsang maikutlo hangata a baka tlhokahalo e matla ea taolo ea kelello ho boloka khetho e nepahetseng ea tlhaiso-leseling e nepahetseng. Tlhokahalo ena ea ho qoba ho khelosoa ke maikutlo a susumetsang maikutlo a hlaha ka nako e le ngoe le maikutlo a mang, a sa lumellaneng le litlhoko tse ling tsa mosebetsi, kapa nakong ea mesebetsi ea memori, moo ho sitisoa ho ka sitisang ts'ebetso ea tlhahiso-leseling e fumanehang marang-rang. Re lekola lithuto tsena ka bokhutšoaane ho hlakisa ho hokahana ha tsona le lingoliloeng tse tsamaisang maikutlo. Bakeng sa tlhahlobo e batsi haholo ea babali ba lingoliloeng tsena ho boleloa Banich et al. (2009).
Ho hatelloa hoa maikutlo nakong ea mesebetsi ea kelello
Lithuto tse ngata li sebelisitse paradigms moo barupeluoa ba tlamehang ho araba karolo e sa amaneng le tšusumetso ea mosebetsi (joalo ka 'mala) mme ba se ke ba sitisoa ke litaba tsa maikutlo (ke hore, mantsoe a maikutlo), kapa ho ea ho tšusumetso e seng ea tlhaho (e leng ntlo) ha a ntse a hlokomoloha ts'usumetso ea maikutlo (sefahleho se tšabehang). Mohlala, libaka tsa rostral (dorsomedial, pregenual and dorsal ACC) le libaka ka bobeli tsa DLPFC le VLPFC kaofela li hlokometsoe maemong a maikutlo a Stroop moo lithuto li lokelang ho qoba ho khelosoa ke litaba tsa mantsoe (Whalen et al., 1998; Compton et al., 2003; Herrington et al., 2005; Mohanty et al., 2007). Bakeng sa tlhahlobo e batsi ea hore na taolo ea tlhokomelo le taolo ea maikutlo li ka kenyelletsa babali ba li-subocates ba li-neurocognitive ba bua ka tsona joang Blair & Mitchell (2009) 'me Mitchell (2011).
Tekanyetso e fetolelang boholo ba lipapatso tsena e hlaha leha ho le joalo ho sa hlake hantle hore na libaka tsena li kenella hobane li fana ka taolo ea kelello, ho shebella likhohlano, li kenella ka lebaka la khohlano / tšitiso ntle le ho laola khohlano / tšitiso, kapa ba mpa ba arabela boikutlong ba maikutlo. Ka mohlala, Mohanty le basebetsi-'moho (2007) bonts'a ka mokhoa o hlakileng hore sebaka sa pele sa "cingrate" (hoo e ka bang BA 24 / 32) se bonts'a ts'ebetso e eketsehileng nakong ea mosebetsi oa Stroop ka mantsoe a maikutlo, le hore sena se lumellana le nako e eketsehileng ea karabelo mosebetsing. Sena se ka fetoleloa ho latela rACC e kenang molemong oa ho fana ka taolo ea kelello ho ba sitisang maikutlo. Le ha ho le joalo, kaha ts'ebetso ea ts'ebetso ea sebaka sena e lumellana le nako e kholo ea ho arabela, boemo ba eona ba ts'ebetso ha bo bonahale bo amana le katleho e thibelang ba sitisang. Ntle le moo, e bonts'itse ts'ebetso e kopantsoeng le amygdala, e hlakileng e sa tsamaellaneng le khopolo ea hore racC e ne e tsamaisa melaoana ea litheko tsa Amygdala. Ehlile hoa hlokomeleha hore bangoli ba fana ka maikutlo a hore, ho fapana le ho bonts'a taolo ea racC ea amygdala, khokahano e phahameng nakong ea ho pepesehela lintho tse khelosang maikutlo e ka bonts'a taolo ea amygdalar kapa tlatsetso ho RACC, ho fapana le tsela e 'ngoe.
Har'a tse ling tse khahlang tsa bopaki ba taolo ea taolo ea bokhoni ba taolo ea maikutlo ho amygdala e tsoa thutong ke Etkin et al. (2006), moo barupeluoa ba ileng ba etsa mosebetsi o kang oa Stroop moo ponahalo ea sefahleho se khahlang kapa e sa sebetseng ka mantsoe a fanang ka maikutlo. Moralo oa thuto e ne e le ntho e rarahaneng kaha bangoli ba ne ba sa shebe ho bapisoa ha liteko tsa maikutlo kapa tsa ho se nke lehlakore kapa liteko tse se nang 'nete, empa ba ile ba hlahloba litlamorao nakong ea liteko tse senang' nete tse neng li latela ka kotloloho liteko tse hlakileng kapa tse fosahetseng. Ho khahlisang, DLPFC, e leng sebaka sa DMPFC sebakeng se phahameng sa 'mankhonthe, le rostral (pregenual) ACC e bonts'itse ts'ebetso nakong ea liteko tse senang' nete tse neng li its'epahalla hore na nyeoe ea pele e ne e le ntle kapa che. DLPFC (le DMPFC) e arabetse ho feta litekong tse senang 'nete tse ileng tsa latela nyeoe e phethahetseng, athe ea Rostral ACC e ile ea arabela kholo ho liteko tse latelang teko e' ngoe e sa sebetseng. Boithuto bona ke bo bong ba lithuto tse 'maloa tse fumanehang ho lingoliloeng tsa taolo ea kelello tse ileng tsa hlahloba ka kotloloho kamano ea libaka tsa pele tsa cortical le ts'ebetso ea amygdala (sebelisa tšebeliso ea kelello ea ts'ebetso ea kelello. Friston et al. 1997). Ho makatsang ke hore ts'ebetso e kholo ho rostral ACC e ne e tsamaisana hantle le ts'ebetso e nepahetseng ea amygdala. Ho latela mohlala oa likarabo tsa amygdala, bangoli ba pheha khang ea hore tšebetso ea amygdala e hokahane le tekanyo ea khohlano nyeoeng e fanoeng, le ka ho hatella tšebetso ea amygdala rostral ACC e fana ka taolo ea likhohlano tsena. Ts'ehetso ea mohopolo ona e tsoa ho datha ea boits'oaro hobane ba bonts'itseng ts'ebetso e kholo ea ts'ebetso litekong tse sa lokang ba bontšitse tharollo e kholo ea likhohlano joalo ka ha ho lekantsoe linako tsa karabelo mosebetsing. Thutong e latelang Etkin et al. (2010) o hlokometse hore khatello ena ea ts'ebetso ea amygdala e bonahala e fokola ho bakuli ba nang le khatello ea maikutlo e akaretsang e amanang le taolo e phetseng hantle, e fana ka khalemelo e ka bang teng ea neural ea bothata ba ho laola ho sitisoa ha maikutlo kapa likhohlano ho bongata bona.
Ho na le morero oa bohlokoa o ka re thusang mabapi le lingoliloeng tsena. Taba ea pele, lithuto tsa sehlopha sa Etkin ha li khothaletse ho ba teng ha thibelo ea tonic lefats'e ea amygdala ke libaka tsa PFC nakong ea maikutlo a loantšanang a maikutlo, kapa ho ikamahanya kamehla le "libaka tse laoloang ke kelello" empa ho e-na le hoo ke mosebetsi o ikhethileng o ipapisitseng le boemo ba khohlano lipakeng tsa hang-hang pele ho susumetso. Haeba ho nepahetse, bokhoni ba ho bona kamano e fapaneng pakeng tsa preingual cingrate (kapa libaka tse ling tsa PFC) le amygdala e kanna ea sebetsa haholo le ho hlahlobisisa.
Mefuta e meng ea bopaki e boetse e phahamisa monyetla oa hore libaka tse ling tsa pele, haholo-holo dorsal ACC, li ka fana ka taolo ea thibelo ho amygdala. Mohlala, thutong e sebelisang paradigm e tšoanang le Etkin et al. (Chechko et al., 2009), bakuli ba nang le ts'abo ea ho tšoha ba bontšitse ho lieha ho feta ho laoloa ka bophelo bo botle nakong ea liteko tse sa sebetseng maikutlo, hammoho le amygdala e phahameng, empa ts'ebetso e tlase ea dorsal ACC / DMPFC, e lebisang tlhahiso ea hore tšabo ea ho tšoha e tšoauoa ka taolo e sa lekaneng ea DMPFC / dorsal ACC. Ka mokhoa o ts'oanang Hariri et al., (2003) e bone kamano e mpe lipakeng tsa amygdala le dorsal ACC (le VLPFC) ha lithuto li tlameha ho ngola mangolo le litšoantšo tsa maikutlo (le mosebetsi oa amygdala o ntseng o eketseha bakeng sa boemo ba papali, le VLPFC le dorsal ACC e ntseng e eketseha nakong ea maemo a label). Ho boetse ho fanoe ka maikutlo a hore dACC e ka fana ka taolo ea taolo holim'a amygdala leha ho se na likhohlano tse itseng kapa maikutlo a itseng a etsang mosebetsi. Pezawas et al. (2005) o bone likamano tse fapaneng pakeng tsa dACC le tšebetso ea amygdala nakong ea mosebetsi o tšoanang oa ho sokela. Ho ka boela ha hlokomeloa hore ACC ea morao-rao thutong ea Pezawas e ne e tsamaisana hantle le ts'ebetso ea amygdala, ho fana ka maikutlo a sebapali se ikhethileng pakeng tsa libaka tse fapaneng tsa cingrate le amygdala, le ho fana ka tlhahiso e eketsehileng, joalo ka ha ho le joalo ka Monhaty et al. (2007) pampiri, eo racC, bonyane maemong a mang e ntle, ho e-na le hore e seng hampe, e kopantsoe le amygdala.
Sehopotso sa ho Sebetsa
Tlhahlobo e 'ngoe e ikhethileng ea liteko tsa taolo ea kelello e bua haholo ka bokhoni ba ho hatella tšitiso ea maikutlo nakong ea mesebetsi ea memori e sebetsang. Hobane palo ea tlhaiso-leseling e ka tšoaroang le ho qhekelloa inthaneteng e lekantsoe (Cowan, 2010), ho bohlokoa hore batho ka bomong ba behe pele ka nepo hore na ke tlhahisoleseling efe e kenang lebenkeleng leo la marang-rang. Ka mokhoa o nepahetseng, re lokela ho boloka tlhaiso-leseling e nepahetseng e amanang le tlhaiso-leseling e sa bohlokoa, empa hape re khone ho lahla litaba tsa mohopolo ha tlhahisoleseling ea bohlokoa e feta merero ea pele. Ka hona, mohopolo oa ho sebetsa o fana ka sebaka se ka thusang bakeng sa ho lekola tšebelisano-maikutlo, haholo-holo ha a fuoa karolo ea bohlokoa ea DLPFC le VLPFC lits'ebetsong tsa memori tse sebetsang (Badre et al., 2005; Blumenfeld et al., 2010; Curtis & D'Esposito, 2004; Jonides et al., 2005; Levy & Goldman-Rakic, 2000, Nee le Jonides, 2010; Postle, 2006; Thompson-Schill et al., 2002).
Litlaleho tse peli tsa Dolcos le basebetsi mmoho ba sebetsa hantle ha li ntse li hokahanya ts'ebetso ea ts'ebetso ea boko le ts'ebetso e atlehileng kapa ho kenella litabeng tsa khokahano e sebetsang (Dolcos le McArthy 2006; Dolcos et al., 2006). Litlaleho tsena ka bobeli li ile tsa sekaseka tlhaiso-leseling ho tsoa ho karabelo e bobebe ea sefahleho eo ho eona ho neng ho hlahisoa litšoantšo tsa maikutlo kapa tse sa nke lehlakore nakong ea mosebetsi. Phuputsong ea pele, ba bonts'itse hore ventrolateral cortex (BA 45 / 47) e sebelisitsoe ka kotloloho nakong ea maikutlo le bahanyetsi ba sa nkeng lehlakore. Barupeluoa ba bonts'itseng ts'ebetso e kholo ea maqhubu a ho ba teng ha lintho tse ba sitisang maikutlong ba lekantse lintho tse joalo e le tse sitisang. Phuputsong e ileng ea latela ba bonts'itse ts'ebetso ea BA 45 e setseng (empa e se tokelo ea BA 45) phapano lipakeng tsa liteko tseo ho tsona batho ka katleho ba ileng ba iphapanya le litšitiso (joalo ka ha li bonts'itsoe ke ts'ebetso e nepahetseng kapa e fosahetseng ea lietsahala ea karabelo). Dolcos et al. (2006) e tlaleha ka VLPFC- amygdala khokahano e sebetsang, ka libaka tse peli li eketseha nakong ea maikutlo a amanang le liteko tsa ho se nke lehlakore. Haholo-holo khokahano ena e ka lehlakoreng le letle ebile e ke ke ea hlalosoa e le khatello ea ho thunya hoa amygdalar.
Lithuto tse entsoeng ke Dolcos le basebetsi mmoho le tsona li fana ka bopaki bakeng sa mekhoa e sa amaneng le ts'ebetso le boikemisetso ho pholletsa le libaka tse ka pele. Ka ho khetheha, libaka tse kenang kahare ho metsi li ile tsa eketseha ka litšitiso tsa maikutlo, ha DLPFC (BA 9 / 46) e fokotseha, e fana ka maikutlo a kamano ea boiphetetso pakeng tsa libaka tsena. Kamano ena ea boiphetetso e tšoana le sebopeho se fapaneng sa dorsal vs. ventral se bonoang ke Perlstein et al. (2002) ba neng ba na le lithuto tse etsang mosebetsi oa memori o sebetsang moo ho neng ho hlaha lits'oants'o tse hlakileng tsa maikutlo e le lintlha tse amanang le mosebetsi [ka mokhoa o khahlisang, kamano ea ho iphetetsa e ne e hokahane haufi-ufi le DLPFC e nyolohang ka libaka tse khothatsang le tse fumanehang (BA 10 / 11) tse bonts'ang ts'ebetso e eketsehileng ea mosebetsi bakeng sa tšusumetso e mpe]. Sebopeho se sa lumellaneng pakeng tsa libaka tse kholo tsa dorsal le ventral tsa PFC se boetse se bonoe maemong a mang a ho sebetsa a memori, ka DLPFC e kholo e amanang le ts'ebetso ea kahare ea letsoho e amanang le mojaro o moholo oa ho sebetsa oa memory (Rypma et al., 2002; Woodward et al., 2006), leha libaka tse khethehileng tsa PFC tse amehang lithutong tse joalo li fapana. Sebopeho se totobetseng sa libaka tse kenang ka hare le tse phoroselang li bontša likhohlano pakeng tsa libaka tsena, empa se bonts'a mofuta oa kamano ea kamano. Ranganath (2006) e fana ka maikutlo a sebopeho sa tsamaiso ea methapo ea kutlo eo ho eona libaka tsa PFC tsa caudal / ventral li fanang ka taolo e holimo ea litsi tsa morao-rao, athe dorsal / rostral PFC e fana ka taolo ea libaka tse ka pele tsa pele tsa kahare. Ho latela pono ena, Ranganath e bolela hore lits'ebetso tsa khetho li kentsoe ke rostral / dorsal PFC li kenyelletsa liphetoho tsa ts'ebetso ho caudal / ventral PFC. Leha ho le joalo, joalo ka ha ho hlalositsoe ka tlase, liphetoho tse ka lehlakoreng le leng li boetse li lokela ho nahanoa.
7. Taolo e amehang ea libaka tse hlokolosi
Ha ho fanoa ka mohlala oa sebopeho o boletsoeng likarolong tse pejana, likhakanyo tsa OFC ho PFC ea morao-rao, ho kenyelletsa DLPFC li ka aroloa ka hore e fana ka karabo hangata. Kahoo, likhakanyo tsena li ka fana ka khethollo le taolo ea libaka tse tsoetseng pele ho feta ka tikoloho. Leha e bonahala e hanana le maikutlo a filosofi a behang mabaka holim'a maikutlo, mohopolo oa hore sebaka se amehang ts'ebetsong e nang le ts'ebeliso e ka fana ka mofuta oa maikutlo mabapi le libaka tse amehang litšebelisanong tse ling tsa kelello o lumellana habonolo le maikutlo a mehleng ena a maikutlo a hatisang matla a maikutlo a ho beha pele le ho khetholla ts'ebetso ea tlhahisoleseling e le ho tsamaisa lipheo tsa bohlokoa tsa bioloji le sechabeng. Maikutlo ana a maikutlo a hlalosoa ka mokhoa o hlakileng ke Gray le basebetsi-'moho (Grey, 2001, Grey, Braver, Raichle, 2002), ea emisang mokhoa oo le ho o ntša o re tšusumetso e matla ea mesebetsi e itseng ea kelello, e matlafatsang le ho sitisa mesebetsi e fapaneng ea temoho ho fihlela litlhoko tsa maemo hantle. Bopaki ba ho feto-fetoha ha maikutlo hoo hoa utloahala ha ho etsoa liqeto (Delgado et al., 2003; Grabenhorst & Rolls, 2009; Hardin, Pine le Ernst, 2009; Piech et al., 2010), empa hape e ka bonoa mesebetsing e meng ea pele e bohareng joalo ka memori e sebetsang. Mohlala, ts'ebetso ea memori e sebetsang ka morao-rao e etsoa ka mokhoa o ikhethileng ka ho kenyelletsa maemo a monate a feto-fetoha le maemo a mabe, ka memori ea ts'ebetso ea sebaka e ntlafatsoa ke maemo a feto-fetohang le maemo a sa sebetseng le memori e sebetsang ea polelo e bonts'a phello e fapaneng (Grey, 2001). Ntle le moo, tlhaiso-leseling e nepahetseng le e fosahetseng ea maikutlo e fokotsa tšitiso ea mohopolo ka ho bapisoa le tlhaiso-leseling e sa nke lehlakore (Levens & Phelps, 2008; 2010). Ka ho le letona la OFC (33 24 N8) le ka ho le letšehali anterior insula (−32 21 2) e arabela haholoanyane tharollong ea maikutlo.
Ka mokhoa o ts'oanang, ts'ebetsong ea ho fetola tšebetso ea boiphetetso, ho kenella ka ts'usumetso e ntle, joalo ka ha ho bapisoa le ho se nke lehlakore kapa ho ama hampe, ho khothalelitse maemo a ho lemoha le ho fokotsa mamello, empa hape ho lebisitse ho distractibility (Dreisbach & Goschke, 2004). Liphumano tse joalo li tsamaisana le bopaki bo ntseng bo hola ba hore maemo a mabe le a fosahetseng a ka phahamisa kapa a tebisa tlhokomelo ho latela matla a mokhoa kapa litšobotsi tsa ho tlohela maemo (mmuso).Fredrickson & Branigan, 2005; Gable & Harmon-Jones, 2008; 2010; Gasper, & Clore 2002).
Ka mokhoa o makatsang, bopaki bo ntseng bo eketseha bo supa hore litlamorao tse susumetsang li ama likarabo tse BOTLENG tsa DLPFC (BA 9) nakong ea mosebetsi oa memori o sebetsang (Gray, Braver, & Raichle, 2002; Savine & Braver, 2010). Ka 'nete, Puseletso Seema (2010) bonts'a hore kahare ho DLPFC (BA 9), meputso ea chelete e khothalletsoang e ntlafalitsoe ka kotloloho mabapi le ts'ebetso e amanang le cue, mme ts'ebetso ena e boletse esale pele hore na nyeoe e tla etsoa hantle. Ha ho kopanngoa hammoho, lithuto tse joalo li hloka hore ho hlakisoe maikutlo a sa hlakileng mabapi le kamano lipakeng tsa ts'ebetso ea kelello le ea maikutlo.
Lithuto tse le 'ngoe tsa lisele li fana ka leseli le eketsehileng nakong ea puisano pakeng tsa libaka tse kenang kahare le tse kenang hanyane ka hanyane mabapi le moputso. Boitsebiso bo tsoang litlalehong tsa pele bo fana ka maikutlo a hore libaka tsa orbital khoutu ea boleng bo hlakileng ba moputso ho feta libaka tse ling tse ka pele, le hore OFC e fana ka leseli lena la boleng ba libaka libakeng tse ka pele tsa pele (Hikosaka le Watanabe, 2000; Wallis & Miller, 2003; Rushworth et al, 2005). Habohlokoa, li-neuron tsa OFC li bonts'a likarabo tsa ho putsa tlhahisoleseling e tlang pele ho likarabo tse amanang le moputso ho DLPFC (Wallis & Miller, 2003). Sena se fana ka mohopolo oa hore OFC e qala ho beha boleng ba moputso ebe e fepa leseli lena libakeng tse ka hokahanyang tlhahisoleseling ena le liketso kapa tlhahisoleseling e ngoe e hlokahalang ho fumana moputso. Rea hlokomela, leha ho le joalo, ha ho hlake hore na tlhahisoleseling ena ea tšusumetso e fihla ho DLPFC ka kotloloho hakae ho latela mofuta oa maikutlo, kapa ho ka nkooa e le lijo ka tlholeho, joalo ka ha ba bang ba 30% ea OFC, DLPFC e ka nahanoa tlhaho (Barbas & Rempel-Clower, 1997). Ho latela sebopeho sa sebopeho phapano ena e tla tseba hore na tefo ea moputso oa lisele tsa DLPFC e bonts'a mofuta oa maikutlo oa DLPFC kapa e bonts'a phetiso ea tlhahisoleseling mabapi le theko eo DLPFC e ka sebetsang ho eona. Ka kopo sheba Mitchell (2011) bakeng sa tlhahlobo mabapi le hore na likarolo tse nyane tsa moputso li ka fetellana joang le melaoana ea maikutlo.
Mohopolo oa hore ts'ebetso ea ho sebetsana le maikutlo o susumetsa ts'ebetso ea kelello le eona e ka ba molemo ha o nahana ka tšebelisano e sebetsang pakeng tsa libaka tsa amygdala le tsa pele. Joalokaha ho hlokometsoe pejana, khokahano e ntle ea ts'ebetso lipakeng tsa libaka tsa PFC (haholo-holo Cgenelie le VLPFC) e hlokometsoe lithutong tse fetileng (Pezawas et al., 2005; Dolcos et al., 2006). Re fana ka maikutlo a hore maemong ana amygdala e kanna ea ba moqapi, ka hore e kanna ea etsa hore boemo bo be pele, mme ba fetise tlhahisoleseling kapa ba leka ho laola libaka tsa PFC ho latela tlhaiso eo ho fapana le tsela e 'ngoe e potileng. Leha ho le joalo, ho tla fihlela kajeno ho entsoe liteko tse 'maloa ho bontša hore na ho bakoa ke eng.
8. Puisano
Re lumela hore tlhahlobo e kaholimo e bonts'a tlhokeho ea ho hlokomela lintlha tsa khokahano ea 'mele ho PFC le kamano ea bona ho amygdala ha ba nahana ka tšebelisano ea maikutlo le maikutlo. Ho sitoa ho etsa joalo ho ka lebisa mehlaleng eo ho leng thata ho e lumellanya le anatomy, ka hona e ka bonahala e sa nepahala. Ho fapana le moo, ho sheba lintlha tsa ts'ebetso ea methapo ea kutlo ha ho ka fana feela ka mehlala e hlakileng ea tšebelisano lipakeng tsa mekhoa ea maikutlo le ea kelello, empa hape e ka senola likarolo tsa ts'ebetso tseo ho seng joalo li neng li ke ke tsa tsotelloa.
Leseli bakeng sa Molao oa Emotion
Ho latela mofuta o ikhethileng oa litsela tsa neuroanatomical lipakeng tsa PFC le amygdala, mehlala e hlakileng ea ho fetoloa ha PFC ka tlhokahalo e tlameha ho kenyelletsa molumo oa, kapa o fetoleloa ka, dorsal anterior cingulate, sebaka se seholo se pharalletseng ho rectus ea gyrus, kapa karolong e ka morao ea sebaka. 47 / 12. Mothating ona oa lebaleng, lipolelo tse bonolo tsa hore PFC e ameha taolong ea maikutlo e fana ka lintlha tse sa lekaneng hore li ka ba molemo, mme maemong a mangata e kanna ea khelosa, kaha boholo ba libaka tsa PFC li haelloa ke mantsoe a matla ho amygdala. Ho hlaha ha mehlala ea tsela e lebisang tlhokomelo ho li-node tsa bohlokoa tse hlahisang amygdala, joalo ka mehlala e hlahisitsoeng le ho lekoa ke Wager et al. le Phillips et al. ke nts'etsopele e khothatsang ntlheng ena. Re belaela hore bakeng sa tsoelo-pele e 'ngoe e lokelang ho etsoa ho utloisisa ho nka karolo ha li-PFC maemong a maikutlo, likarolo tsa setho sa dorsal anterior cingulate, posterior 47 / 12 le sebaka se ka tlasa taolo ea amygdala li tla hloka ho tsejoa.
Potso ea bohlokoa e boetse e sala mabapi le ts'ebetso e atileng haholo ea ts'ebetso ea PFC e hlahang nakong ea taolo ea maikutlo e amanang le libaka tsena tsa bohlokoa, hobane ke lithuto tse fokolang feela tse ileng tsa hlahloba kahare ho tumellano ea tšebetso ea Intra-PFC. Anatomically, libaka tsena tsa PFC ha li hokahane ka tsela e tšoanang le dorsal anterior cingrate, posterior 47 / 12 kapa sebaka se seholo, ka hona li kanna tsa amahanngoa ka mokhoa o ikhethileng le litsela tse fapaneng tsa amygdala. Re belaela hore ho utloisisa ho nka karolo ha PFC ho taolo ea maikutlo ho tla hloka ho hlakisoa hore na ke libaka tse kae tsa PFC tse se nang likarolo tse hlakileng tsa ts'ebelisano 'moho li ikamahanya le libaka tse ling tsa PFC tse nang le lipallo tse lekaneng ho ntlafatsa tšebetso ea maoto.
Lintlha tse mabapi le tšusumetso ea litšusumetso
Re pheha khang ea hore mefuta e ka sehloohong ea ts'ebelisano ea PF, le likamano tsa PFC-amygdala tse hlalosang taolo e matla e hlakileng ea taolo e tlase e tlase e sa tsamaellaneng le litšobotsi tsa laminar tsa khokahano lipakeng tsa libaka tsena. Khang ea rona e khahlano le mefuta ena ea setso ea likhetho tsa PFC-amygdala le intra-PFC e ts'ehehile haholo mohlaleng o hlalositsoeng ke Barbas le basebetsi mmoho, moo sebopeho sa laminar se hlalosang hore na merero e emela tlhahiso-leseling joalo ka ho rata, phetiso ea tlhahiso-leseling. Haeba ho nepahetse, libaka tse amanang le maikutlo li bonahala li fana ka taolo e holimo-tlase ea ho fana ka tlhahiso-leseling ka sepheo sa ho fetisa tlhahiso-leseling ka tlhahisoleseling ho feta libaka tse tloahelehileng tsa kelello tsa PFC.
Re lumela hore poleloana ea taolo e holimo-tlase e lebisitse mohopolong o hlakileng oa kutloisiso pakeng tsa likamano tsa kelello le ts'ebetso ea kelello. Leeme lena le lumellana le pono ea filosofi ea likarolo tsa "ho lemoha lintho tse amanang le" le "maikutlo" tse behang kutloisiso holima maikutlo a bophoofolo. Empa khethollo ena e ka kena-kenana le bokhoni ba rona ba ho utloisisa ka botlalo tsela eo boko bo sebetsanang le tlhaiso-leseling ka eona. Haeba ts'ebetso ea maikutlo e laola le ts'ebetso e "hlokolosi" ts'ebetso, haholo kapa ho feta ka tsela e 'ngoe, poleloana ea holimo le tlase e ka ba e sa lokelang ho nahana ka tšebelisano ea maikutlo.
Meeli ha e kenyelletsa mosebetsi ho tsoa mohahong
Bohlokoa ba sebopeho sa sebopeho ke hore e lebisa ho likhakanyo tse matla mabapi le mofuta oa puisano ea libaka. Leha ho le joalo, liphoso tse 'maloa li ka phahamisoa hang hang ho hula liqeto tse sebetsang mabapi le likarolo tsa anatomical. Taba ea pele, leha sebopeho sa sebopeho se ts'ehelitsoe ka matla ho latela lipakanyo tsa sona tsa mekhoa ea khokahano ea laminar e thehiloeng ho boiqapelo ba marang-rang, litekanyetso mabapi le ts'ebetso ea lits'ebetso tsena tsa khokahano ea laminar ha li so fumane teko e hlophisitsoeng makaleng ka ntle ho melapo ea ts'ebetso ea kutlo. Le ha ho bonahala ho utloahala ho nahana hore likarolo tse tšoanang tsa ts'ebetso li supa lits'oants'o tsa laminar tsa monahano ho pholletsa le boko, ha ho hlile ha ho joalo. Kahoo, maikutlo a mabapi le ts'ebetso ea likhokahano ho PFC a sebetsa hantle ha litšobotsi tsa tšebetso tsa tšebeliso ea tšebeliso ea marang-rang le litekanyetso tsa karabo li netefalitsoe ho tšoarella ho li-cortices tsa mokhatlo.
Re hlahisitse khokahano e matla pakeng tsa karabelo e sebetsang le taolo e holimo-tlase, le khokahano e ts'oanang pakeng tsa lits'ebetso tsa feedirect le tlase. Litlhaloso tsa litlhaloso le ho fepa li tsoa ho khopolo ea taolo, e lekang ho hlalosa ts'ebetso ea litsamaiso tse matla. Ho amoheloa ha mantsoe ana ke litsebi tsa kelello le litsebi tsa kelello ha ho makatse ha mohopolo oa mekhoa ea ho fana ka maikutlo a fanang ka taolo ea taolo le mekhoa e felletseng e fanang ka phetiso ea tlhahisoleseling libakeng tse phahameng molemong oa ho sebetsa. Leha ho le joalo, equation e bonolo ea taolo e holimo-tlase ka karabo le tlase-up ka feedirect e na le bothata ho isa moo likarolo tsa tlatsetso li hlahisoang ke conceptualizations ea holimo le tlase le tlase. Likarolo tse ling ha li hlahisoe ka mokhoa o hlakileng, empa li ka paka ho le hoholo ha ho phethoa litsebo tsa ts'ebetso ea tlhahisoleseling. Re belaela hore baithutatsi ba bang ba sebelisa mantsoe a holimo ho ea tlase le tlase-tlase ka mekhoa e sa lumellaneng le maikutlo le mekhoa e lumellanang joalo ka ha e hlalositsoe ke khopolo ea taolo, empa ho se lumellane ho joalo ha ho hlalosehe ka mokhoa o hlakileng ka har'a lingoliloeng.
Ha re tšoaea maikutlo le litlatsetso tsa PFC, re hlokometse hore ha re bolele hore likhakanyo tsohle ke tsa mofuta o le mong. Libaka li na le kopanyo ea maikutlo, khokahano ea feedback le ea morao-rao, empa boholo ba likhokelo tsena bo fapana haholo libakeng. Ka hona re supa lits'oants'o tse kholo tsa khokahano, empa sena ha se bolele hore likhokelo tse setseng ha li bohlokoa. Mohlala, libaka tsa PFC tse nang le eulaminate ka sebele li na le lipalo tse lekaneng tsa ho thusa ho laola likarolo tsa libaka tse fokolang haholo, le haeba ena e se mokhoa oa puisano o pakeng tsa libaka.
Ho feta moo, likhakanyo tsa mofuta oa li fepa ka linako tse ling li ka fetolela ts'ebetso libakeng tse fumanehang ho fapana le ho ba le tlhahisoleseling. Mohlomong mohlala o motle oa mofuta ona oa feed moderate o hlaha mefuteng e kopaneng ea tlholisano (Desimone le Duncan 1995; Duncan et al. 1997) eo ho fumana phaello ea moemeli a le mong ho hlahisang khatello ea e mong. Mehlaleng e joalo, ho fepa moemeli o fanoeng ho ka lebisa ho matlafatso ea ts'usumetso ea khothalletso eo, le khatello ea bobeli ea khothatso e 'ngoe (Desimone le Duncan 1995). Ka tsela ena, se feptjoang pele se ka sebetsa ho fetola tšebetso libakeng tse lebelletsoeng. Maemong a ts'ebetso ea PFC, lets'oao la DLPFC le ka fetola tlholisano lipakeng tsa boemeli bo ka bang teng ho OFC ka mofuta ona oa provo. Mofuta ona oa mochini oa tlholisano oa khahla hobane o tla bolela likarolo tse ikhethang tsa computational (Walther & Koch, 2006), tseo ka kakaretso li sa kenyellelitsoeng ho mehlala ea melao ea tsamaiso.
Ha ho nahanoa sebopeho sa sebopeho, ho bohlokoa ho pheta-pheta hore mekhoa e sebelisitsoeng ke Barbas le basebetsi mmoho le bona ho hlalosa feedxp le khokahano ea maikutlo ha e lumellane ka botlalo le litekanyetso tse sebelisitsoeng ke bafuputsi ba bang ba hlahlobe tlhophiso ea sebaka sa tatellano ea merero ea laminar. Haholo-holo, litlhaloso tsa ho fana ka maikutlo le khokahano ea pele hangata li hlalosoa ho buoa ka karolo ea IV, e le hore likhakanyo tsa feedxp (tse nyolohang) li hlalosoa ho felisoa ha tsona ka karolo IV (kapa boholo ba safu ea IV), ha maikutlo (a theoha) a emisoa ka ntle ho mokato. IV. Leha ho khomarela ka tieo molaong oa IV ho kanna ha elelloe hampe, joalo ka ha mekhelo ea mekhoa ena e hlokometsoe (Pandya le Rockland, 1979; Felleman le Van Essen, 1991), tšusumetso ea mekhoa e pharalletseng ea ho lumella likhakanyo tse qetellang libakeng tse infragranular V le VI hore li nkuoe e le likhakanyo tsa lijo li sa utloisisehe ka botlalo. Ka mokhoa o makatsang, lithuto tsa nako ea ho thunya libakeng tse fapaneng tsa PFC li ka rarolla potso ena, empa tlhaiso-leseling mabapi le taba ena ea fokola.
Potso ea mekhoa e baka ho emisa pele u nka hore OFC-DLPFC ehlile e na le mokhoa oo OFC e lokelang ho ts'oaroa joalo ka boemo bo phahameng ho feta DLPFC, 'me ha se morero oa rona ho pheha khang joalo. Leha ho le joalo, ho ka boleloa ka ho hlaka hore mekhoa ea likhakanyo ka sebele ha e lumellane le mokhatlo o ikhethileng moo DLPFC e boemong bo phahameng kaholimo ho OFC, ka mokhoa o ts'oanang le libaka tse phahameng tsa maikutlo tse lutseng ka holim'a libaka tsa mantlha tsa kutlo. Kahoo, mehlala ea mokhatlo oa PFC e ka ba bohlale ho qoba boemo bo atileng ba DLPFC e le ba lutseng kaholimo ho puso ea libaka tsa PFC.
Meetso ea khokahano ea feedirect le karabelo
Ha ho sekasekwa mefuta e teng ea tšebelisano-maikutlo e mabapi le maikutlo a maikutlo ho ke ke ha hlokomeleha hore lithuto tse 'maloa tse phatlalalitsoeng ho tla fihlela kajeno li kenyelitse liteko tse ikhethang tsa hore na khakanyo e bonts'a karabo, menahano e felletseng kapa ea morao-rao Seminowicz et al. 2004). Boholo ba lithuto tse natefisang taba ha li fane ka leseli le ikhethang le ka rarollang bothata bona. Leha ho le joalo, nts'etsopele ea morao-rao mekhoeng ea ho etsa hore puisano e sebetse hantle e fana ka lisebelisoa tse ka sebelisoang ho bontša mofuta le tataiso ea khokahano lipakeng tsa libaka. Mohlala, mohlala oa matla oa ho qobella (DCM) o sebelisa mohlala oa malapa le mohlala oa Bayesian o ka sebelisoa ho lekola li-hypotheses mabapi le tataiso le mofuta oa phallo ea tlhaiso-leseling le ho feto-fetoha ha maemo libakeng tse fapaneng tsa boko (Friston et al. 2003; Chen et al. 2009; Daunizeau et al. 2009; Friston le Dolan 2010; Penny et al. 2010). DCM e kanna ea leka mehlala ea tlholisano joalo ka ho fana ka lihlooho ho bapisa hore na DLPFC e theola li-amygdala ka kotloloho kapa ka sebopeho se bohareng. Ho fihla joale, ke lithuto tse fokolang feela tsa DCM tse amanang le ts'ebetso ea maikutlo tse phatlalalitsoeng (Ethofer et al. 2006; Smith et al. 2006; Rowe et al. 2008; Almeida et al. 2009), 'me tsebong ea rona ha ho na lithuto tse hatisitsoeng ka kotloloho tse sebetsanang le taolo ea maikutlo. Le ha ho le joalo, ts'ebeliso ea mekhoa e joalo e kanna ea ntlafatsa haholo kutloisiso ea rona ea ts'ebelisano-maikutlo e mabapi le maikutlo lilemong tse tlang.
Liteko tse tobileng tsa tšusumetso
Mohlomong tsela e molemohali ea ho theha likamano tse sebetsang pakeng tsa likarolo tsa boko ke ka tlhahlobo ea tikoloho e le ngoe nakong ea taolo ea 'mele ea ho khetha holimo kapa tlase ea sebaka se seng. Mohlala, haeba DLPFC e hlile e sebetsa ho nyahlatsa ts'ebetso ea OFC, motho a ka lebella likarabo tse fetelletseng ho OFC ha DLPFC e tsamaisoa kantle ho naha. Monyetla ona o ka rarolloa ka ho hlahloba mesebetsi ea OFC le fMRI ho bakuli ba nang le liso tsa DLPFC. Ntle le moo, tšusumetso ea matla a khoheli a transcranial (TMS) e ka sebelisoa ho DLPFC ho fetola tšusumetso ea DLPFC ka nakoana mesebetsing ea OFC. Ka 'nete, Knoch et al. (2006) sa tsoa tlaleha hore TMS holim 'a DLPFC e nepahetseng e hlahisitse liphetoho mesebetsing ea poso ea OFC ka mokhoa o ts'ehetsang khafetsa. Ka mokhoa o ts'oanang, e tla ba ho khahlisang ho tseba hore na liso tsa karolo e le ngoe ea cortex ea pele li ama ts'ebetso ho likarolo tse ling tsa marang-rang. Mohlala, haeba OFC e bohlokoa bakeng sa ho bala boleng bo hlakileng ba moputso, ho etsahalang ka libaka tse eketsehileng tsa thobalano ha OFC e tlosoa? Saddoris et al. (2005) ba sebelisitse mofuta ona oa ho hlahloba hore na li-OFC li-alter amygdalar li thunya ka litoeba, empa lithuto tse ling tse sebelisang mokhoa ona li atisang ho ba teng. Lingolo tse ntseng li eketseha mabapi le khokahano e sebetsang li ka etsahala hore li eketse kutloisiso ea rona ea hore na likarolo tsena tsa bohlokoa tsa boko li sebelisana joang. Leha ho le joalo, kutlwisiso e felletseng ea lipuisano tsena e tla fihlelloa feela ka tlhokomelo e hlakileng ea likarolo tse ikhethileng tsa li-circuits tsena.
Lintlha-khōlō tsa Patlisiso
- Khokahano e ikhethang ea pele e laola maikutlo a maikutlo a amygdala
- Mekhoa ea projeke ea Laminar e khetholla phallo ea tlhahisoleseling ho cortex ea pele
- Litekanyetso tse hlakileng le tse fanang ka maikutlo li phephetsa mokhatlo o hlophisitsoeng pele
lumela hore baa fokola
Mosebetsi ona o ne o tšehelitsoe ke lithuso T32MH018931-21, T32MH018921-20, & 5R01MH074567-04 ho tsoa Setsing sa Naha sa Bophelo ba kelello. Re leboha Tawny Richardson ka thuso ea hae ho hlophisa buka e ngotsoeng ka letsoho.
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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