Ukuqonda okungokwemvelo ekusebenzisaneni kwemizwelo nokuqashelwa kwe-prefrontal cortex (i-2011)

Uhlobo lokugcina oluhleliwe lomshicileli luyatholakala at Neurosci Biobehav Rev

Bona ezinye izihloko ku-PMC ukuthi ubalula isihloko esishicilelwe.

Ukucwaninga ngokwengqondo kukhombisa ukuthi izinqubo ezingokomzwelo ziyaxhumana nezinye izici zokucabanga. Ucwaningo luveze bobabili amandla wethonya lomoya ukuthonya uhla olubanzi lwemisebenzi yokusebenza kwengqondo, kanye nekhono labantu lokusebenzisa izindlela zokuqondisa eziqonda phezulu ukulawula izimpendulo zemizwelo. Izingxenye ze-cortex yangaphambili zibonakala zidlala indima enkulu kulokhu kusebenzisana. Kodwa-ke, indlela okwenziwa ngayo lokhu kusebenzisana kusalokhu kucacisiwe ngokwengxenye. Ekubuyekezweni kwamanje sichaza ukuxhumana okukhona phakathi kwezindawo zangaphansi nezendawo yangaphambi kwangaphakathi kanye nokuxhumeka kwazo nezifunda ezinamandla. Ukusethwa kwezindawo zangaphambi kokuqala kuphela ezingathonya ngqo ukucubungulwa kwe-amygdalar, futhi njengamamodeli wokulawulwa kwangaphambilini kwezimozamamodeli namamodeli wokulawulwa okungokomzwelo okufanele ancishiswe ezindleleni ezinamandla zothonya. Siphinde sigxile ekutheni iphethini ethile yokuxhuma okuphakelayo nokuphendula phakathi kwalezi zifunda ingaluchaza kanjani uhlobo lokuhamba kolwazi phakathi kwezindawo zangaphakathi nezendawo yangaphansi yangaphansi ne-amygdala. Lezi zindlela zokuxhuma azihambelani nokucatshangelwa okumbalwa okuvezwa ngesimo sokuxhumana phakathi kwemizwelo nokuqonda.

I-Topography

I-PFC ihlukaniswa kaningi izifunda ezibanzi ze-6, dorsolateral, ventrolateral (VLPFC), i-frontopolar (FP), OFC, i-ventromedial (VMPFC), ne-dorsomedial (DMPFC) (bona Umfanekiso we-1). Imingcele eqondile yendawo yalezi zifunda isetshenziswa ngokuhlukile ngabaphenyi, kepha ukufakwa kwamagama jikelele kufakazele ukuba wusizo njengohlaka olubanzi lokuhlela lokuqonda ukwakheka nomsebenzi we-PFC.

  

Umfanekiso we-1

Izifunda ezijwayelekile ze-PFC ebantwini. Izifunda ezinemibala zibonisa ukulinganiselwa okuningiliziwe kwezindawo ezibanzi ze-PFC. Kokubili ukubuka okulandelayo (kwesobunxele) kanye nokubuka okungokwesimo (kwesokudla), izifunda zimbozwe endaweni ebanzi “ekhuphukile ngokwengxenye” ...

I-Phylogeny neCytoarchitecture

I-PFC iqukethe izindlela ezimbili zokwakha izakhiwo ezihlukanisiwe, ezinokwehlukaniswa phylogenetically (IBarbas, i-1988; ISanides, 1969; I-Yeterian & Pandya, ngo-1991). Umkhuba we-basoventral usuka endaweni eyisisekelo ye-ophodoory (i-rendortical) ngokusebenzisa i-OFC bese usakazeka ungaphandle kwesigxobo esingaphambili esingaphambili, futhi kamuva uye ku-VLPFC (ephela endaweni yeBrodmann (BA) V46). Ngokuphambene nalokho, indlela esetshenziswayo yokuxhumana phakathi kuqala i-corpus collosum, iqhubekela phambili odongeni lwendondo ye-lobe yangaphambili bese igoqa umngcele ophakeme we-lobe ungene kwi-DLPFC (ephetha kwi-BA D46). Unembe ngamunye welezi zimo ukhombisa iphethini yezigaba ezilandelanayo zokwakheka kwe-cortical eboniswe ekuthuthukisweni nasekunabiseni kongqimba lwe-granular IV. Ingxenye yakudala yokuziphendukela kwemvelo yalezi zindlela i-agranular emvelweni, kanti izindawo ezincane ezivele ngokuguqukayo zinesendlalelo esiminyene nesichazwe kahle sogogo. Ngokwesimo se-basoventral, lokhu kuqhubekela phambili kwe-cortical kuqala endaweni engemuva ye-OFC (i-agranular insula isebenzisa isigama sikaCarmichael noPrint (UCarmichael & Intengo, 1994)) elandelwa yi-dysgranular (buthakathaka granular) cortex ezindaweni eziphakathi ne-OFC, iqhubekela eulaminate I cortex nge-granular ungqimba olungafaniyo lwe-IV njengoba eyodwa ihamba ngaphandle noma kamuva, futhi ekugcineni ifinyelele eortaminate II cortex enobubanzi obukhulu be-IV ne-supragranular eqinile izendlalelo lapho uhamba uqonde esifundeni sangaphambi kwesifunda kanye nasezifundeni ezihamba ngezinyawo (IBarbas & Pandya, 1989; UCarmichael & Intengo, 1994; I-Petrides & Mackey, 2006; Intengo, i-2006a). Umkhuba we-Mediodorsal ukhombisa ukuqhubekeka okufanayo kwe-cytoarakhiwa. Ukuqala nge-periallocortex cortex eduze kwe-rostral corpus collosum, umkhuba uba dysgranular ku-cingate (kufaka phakathi izindawo ezingaphezulu, ezingaphambi kwesikhathi, kanye nezindawo eziphezulu), u-eulaminate I njengoba umuntu eqhubekela phambili odongeni lwendabuko noma ngokudlulele kwi-gyrus ephambili yangaphambili, futhi ekugcineni ibe yinqolobane I-II ezifundeni ezi-dorsolateral (BA 8 ne-46).

Ukuze ugweme ukudideka, siphawula ukuthi ukusetshenziswa kwegama le-Mediodorsal mkhuba akufanele kudidaniswe nesifunda se-DMPFC esichazwe Umfanekiso we-1. Isimo se-Mediodorsal sifaka i-DMPFC, kodwa futhi sifaka izindawo ze-VMPFC i-25 ne-32, kanye nezingxenye ze-BA 10 odongeni lwezindondo (indawo eyi-10m kusiqalo se Ongur et al. (2003); Umfanekiso we-2).

  

Umfanekiso we-2

Amathrendi we-basoventral ne-Mediodorsal phylogenetic. Kuzo zombili lezi zindlela, i-cortex ihlukaniswa kancane kancane. Umdwebo uguqulwe nemvume evela UBarbas noPandya 1989. Izifinyezo: Pro procosex; I-PAII limbic periallocortex; D dorsal; ...

Iphethini yentuthuko yemvelo njengoba eyodwa isuka ku-agranular iye eulaminate II cortex ihambisana nokwanda kwenani eliphelele lama-neurons (cellensens), usayizi wamaseli we-pyramidal ezingqimbeni II no-V, kanye nezinga le-myelination (IBarbas & Pandya, 1989; Dombrowski et al. I-2001; Umfanekiso we-3), okuholela ekuthini kube nezimpawu ezahlukahlukene zokulungiswa kolwazi kuzo zonke izifunda ezihlukile. Eminye umehluko omkhulu phakathi kwezifunda zangaphambi kokuvela ngokwemibandela yokufakwa emlandweni, imvamisa ibonisa izici ezihlukile zokuhlangana. UCarmichael noPrint (UCarmichael & Intengo, 1994) hlukanisa i-Macaque OFC ne-PFC yokwehlukaniswa ngokwezigatshana eziningi ngokususelwa kuzimpawu ezinjalo (bheka figure 4), futhi eziningi zalezi zici zingakhonjwa kubantu (Ongur et al., 2003). Izici ezihlukanisayo zokuhlangana okubonwe ngokwahlukaniswa kokuqala kwangaphambilini zithinta izici ezithile zokucutshungulwa kolwazi okufezekiswe yangaphambilini kwangaphambilini (U-Wang et al., I-2004; I-Zald, 2007), kepha bangaphezu kobubanzi baleli phepha. Ngokudabukisayo, ukwahlukaniswa okuchazwe ngendlela ehlelekile kwe-PFC kunamaphethini ahlukahlukene wokuxhuma ngaphakathi kwe-PFC kanye nezinye izifunda zobuchopho kanye nezengqondo zobuchopho.

  

Umfanekiso we-3

Amazinga alandelanayo womehluko kumasendlalelo we-cortical ngaphakathi kwe-PFC. Kanye nokuvela kogqinsi lwe-granular cortical IV, kukhona ukwanda kwesibalo samaseli, kanye nosayizi we-pyramidal neurons kuzendlalelo III no-V. Umdwebo oguqulwe nemvume. ...

  

Umfanekiso we-4

Imephu yefulethi ekhombisa ukwahlukaniswa kwezakhiwo kwe-PFC eMacaque. Kulesi sifanekiselo semephu esiyicaba, i-cortex isikwa kumgomo we-sulcus (phansi nangaphezulu wezo zibalo). Isikimu nezinombolo zokulebula zenziwe kabusha kusuka ku UCarmichael noPrint, ...

Ukuqanjwa kwamakhompiyutha kuHumans

Yize kunokubaluleka okubalulekile kokuqanjwa kobuntu nokwenziwa kobuntu phakathi kwendawo yokuhlala yangaphambili, kanye nemikhuba ejwayelekile yama-phylogenetic kwabelwana ngayo kuzo zonke izinhlobo zezinto eziphilayo, kunobunzima obuhlukahlukene bokuhamba phakathi kwedatha yomuntu neyezilwane. Okokuqala, izifundo zabantu ze-neuroimaging zivame ukubhekisela ezindaweni zeBrodmann (Brodmann, 1914), kepha angikhombisi intuthuko ekubonakalweni kwezindawo ezisetshenziselwa ubuchwepheshe bemvelo kanye nemingcele yezindawo okwenzekile kusukela umsebenzi wokuphayona kaBrodmann cishe eminyakeni eyikhulu edlule. Okwesibili, ukusetshenziswa kwalawa amalebula endaweni kuvame ukususelwa ku-Talairach atlas (ITalairach neTournoux, ngo-1988), kepha le atlas iyisilinganiso esingcono kakhulu, ngoba ukuhlaziya ubuchwepheshe be-cytoarological asenzelwanga ebuchosheni obakha isisekelo se-atlas. Okwesithathu, kukhona ukungaboni ngaso linye phakathi kwamalebula ezilwane namalebula abantu ku-patral frontal lobe, ngoba imininingwane yezilwane isebenzisa okuhlukile kwilebula eyenziwe ngu-Walker (Walker, 1940), abanye abalobi manje abaye bangeza kubantu (I-Petrides & Mackey, 2006; Ongur et al., 2003), ngenkathi iningi labacwaningi be-neuroimaging besebenzisa amalebula we-Brodmann. Ngeshwa, kwesinye isikhathi akukacaci ukuthi yibaphi abacwaningi bohlelo lwe-neuroimaging abakhombisa lapho bebika abakutholile. Lokhu kukhiqiza ukuqondisisa okuthile kwi-lateral OFC / VLPFC, lapho abacwaningi abangabantu bavame ukubhekisela kwi-BA 47, kepha izincwadi zezilwane zibhekisela endaweni ye-12. Ilebula 47 / 12 manje selisemukelwa ngabathile be-neuroanatomists ukuchaza le ndawo kubantu, yize umngcele we-medical wale ndawo uhlala uphikiswa ngabaphambili be-neuroanatomists (I-Petrides & Mackey, 2006; Ongur et al., 2003). Ngokufanayo, izindawo i-13 ne-14 zidalulwe ngokucacile ezinkawu, futhi izindawo ezifunda ekhaya zibonwa kubantu, kepha azitholakalanga nguBrodmann noma uTalairach, ofake ilebuli ejwayelekile yendawo ye-11 kuzo zombili izigaba ezingemuva nezingaphandle zedical OFC. Lapho sichaza idatha ye-neuroimaging yabantu, ngokuvamile sibheka uhlelo olubanzi lokulebula oluchazwe yi I-Petrides neMackey (2006), kune-Talairach atlas ukuze usizakale ngemininingwane evela ezifundweni zamanje ezingezona ezomuntu.

Okokufaka kwe-Amygdalar ku-PFC

I-OFC ne-PFC yezokwelashwa bathola okokufaka okuvela kuma-amygdala (I-Amaral et al., 1992; UCarmichael & Intengo, 1995; IBarbas & Zikopoulos, 2006). Lokhu kuphambene kakhulu ne-DLPFC, ethola ukuqagelwa okuncane okuvela kuma-amygdala. Ukubuyekezwa kwezincwadi kukhombisa ukuthi okunye ukuqagelwa kwe-PFC ye-ventral kanye ne-medial kuyahlukahluka ngokuya nge-nucleus noma i-subnucleus yemvelaphi (Amaral & Intengo, 1984; IBarbas & De, ngo-1990; I-Amaral et al., 1992; UCarmichael & Intengo, 1995). Kodwa-ke, le mininingwane ingaphezu kobubanzi baleli phepha, futhi isithombe esinamandla soxhumaxha siphuma kuyo yonke i-nuclei ehlukahlukene ukwazisa ingxoxo ejwayelekile yamaphethini wokuxhumana. Umfanekiso we-5 ibonisa isikimu esivamile sokucatshangelwa kwe-amygdalar (kusuka ezincwadini eziningi ze-amygdalar) singene endaweni yezengqondo ne-ventral yobuchopho be-macaque besebenzisa i-Carmencell kanye ne-Price nomenclature. Lesi sibalo sikubeka sobala ukuthi ingaphezulu le-orbital alifani ekuxhumaneni kwalo ne-amygdala. Okuqaphelekile ikakhulukazi ukungatholakali okufakiwe okuningamelayo ezindaweni ze-13m, 13l, 12m, 11l, kanye ne-10o ebusweni be-orbital. Udonga lwe-medial libuye lufake okokufaka kwe-amygdalar okukhulu, kepha futhi alufani, ngoba indawo 10m noma indawo 9 ayitholi okokufaka okubalulekile kwe-amygdalar.

  

Umfanekiso we-5

Amamephu we-cytoarchitectural of orbital surface. Ukulebulwa kobumbeko kwe-cytoarological kwe-lobes yangaphambili kuguqulwe kusuka Brodmann (1914) (kwesokudla), I-Ongur, Isikebhe Nentengo (2003) (maphakathi) no I-Petrides neMackey (2006)(kwesobunxele). Qaphela umehluko omkhulu phakathi ...

Iziphetho ezimbili zingahle zithathwe kule ndlela yokufaka. Okokuqala, okokufaka kwe-amygalar kwi-PFC kucacisiwe ngokwakhiwa futhi kugxiliwe ezifundeni ezithuthukiswe ngokwakhiwa kwezwe. Lokhu kukhombisa ukuthi kungaba yiphutha ukuphatha ngohlobo lwayo yonke i-OFC noma i -ialial PFC sengathi ixhunywe kakhulu kwi-amygdala. Esikhundleni salokho, ukunakwa kwendawo engaphakathi kwe-OFC kanye ne-medial PFC kuyelulekwa ngaphambi kokudonsela okuphathelene nokuxhumana kwe-amygdalar. Okwesibili, i-DLPFC kanye ne-FP bathola okokufaka okubuthakathaka okuqondile kwe-amygdalar (impela amasu abucayi kakhulu abonisa ubufakazi bokufaka kwe-amygdalar). Njengomphumela, amathonya e-amygdalar ku-DLPFC kanye ne-FP ukucubungula kungenzeka ukuthi angaqondile, kungaba ukuthi adluliselwa ezifundeni ze-cingulate noma ezingemuva kwe-OFC (noma ezinye izindlela ezijwayelekile, njengokushintshwa kwezinhlelo ze-neurotransmitter).

Ukukhishwa kwangaphambi kokuqala ku-amygdala

Imiphumela ye-PFC iye e-amygdala nayo icacisiwe esifundeni (Intengo, i-2006b; I-Ghashghaei et al., 2007; UStefanacci no-Amaral, 2002; UStefanacci no-Amaral, 2000). Ngokuvamile, izindawo ezisekuqaleni ezithola ukuqagelwa okuvela kuma-amygdala zithumela ukuqagela emuva e-amygdala, kuyilapho izindawo ezingatholi okufakwayo okuningana kwe-amygdalar (njenge-DLPFC ne-FP) kunamathuba angatheni aqine kakhulu e-amygdala. Ubuningi bokuqagela bukhombisa ubukhulu be-cytoarchitectonics, ngokuwohloka kokwehla kobuningi beprojekhthi njengoba uhamba usuka ezindaweni ezinamacala aguqukayo uye isocortex eyakheke ngokwengeziwe. Leli phethini libonisa ukuthi izindawo ze-isocortical (DLPFC ne-FP) azikwazi ukunikeza ithonya eliqondile eliqinile ngaphezulu kwe-amygdala, futhi ngezinga lapho bathonya khona i-amygdala, ithonya kungenzeka lingaqondile. Lokhu akusho ukuthi akukho okuqageliwe okuqondile kusuka kwi-DLPFC kuya e-amygdala, njengoba izifundo eziningi ngempela zibonile ukuqagelwa okuqondile kusuka endaweni 9 nase46 (UStefanacci no-Amaral, 2002; Aggleton et al., 1980; Amaral & Insausti, 1992). Esikhundleni salokho, ukuqagelwa ngokuvamile kukhanya kakhulu ukuhlinzeka ngomthethonqubo obanzi wokusebenza kwe-amygdalar.

Yize i-cytoarchitectonics ejwayelekile ihlinzeka ngothishanhloko wokuhlela oqinile ngokuya ngokuxhunywa kwe-amygdala-pre-preal, ukusatshalaliswa kokungenayo nokufakiwe kwesifunda okulinganiselwe akuyona i-symmetric (I-Ghashghaei et al., 2007). Ngokuphawulekile, okokufaka okuphezulu kakhulu kwe-amygdalar ku-PFC kutholakala esifundeni esingagugiyo sangemuva komgwaqo we-OFC, kanti indawo enkulu ekuphumeni esifundeni i-amygdala ivela esifundeni sangemuva sangaphambilini se-cingrate (BA 25) kanye nezingxenye ze-dorsal anterior cingulate (BA 24 ). Ngokujwayelekile, izindawo zodonga lwendondo zinokukhipha okuphezulu kunokufaka okuvela kuma-amygdala, kanti izindawo zangemuva kwe-OFC zinokufaka okuphezulu kunokukhishwa. Kuyathakazelisa ukuthi izindawo ezinamathele kakhulu ezixhunyaniswe ngokwedlule i-PFC, izifunda ze-DLPFC (BA 8, 9 kanye ne-dorsal 46) zinokufaka okukhulu kusuka kokuphumayo kuya ku-amygdala, kanti leli phethini liphindiselwa ngaphakathi kwe-VLPFC. Okuqaphelekile kulokhu, isifunda sangemuva sendawo i-12l ngaphakathi kwe-VLPFC sinikezela ukuqagela okulinganiselwe kuma-amygdala, okwenza kube ukuphela kwesifunda se-PFC esinokufaka okuphambili kwe-amygdala. Ngempela, lokhu kucatshangelwa kunamandla kunokubonwa ezifundeni ezingaphandle kwe-orbital, ezabelana ngezindawo ezinkulu ze-12l zokuphuma kunokufaka, kepha zibonise amazinga angenamandla okuxhuma ngokuvamile kunendawo ye-12l.

Inani lama-subnuclei e-amygdala ahlukahlukene athola okokufaka kwe-PFC. I-basal kanye ne-accessory eyisisekelo ne-mainial nuclei ithola ukuqagelwa okuminyene, kanye nokwamukela ukuqagela okuvela ezinhlakeni ezibanzi kakhulu zezifunda ze-PFC, kuyilapho i-mainic, central ne-cortical nuclei ithola ukuqagelwa kwe-PFC, kepha ngezinga elincane nelibanzi (UStefanacci no-Amaral, 2002). I-BA 25 iyaphawuleka ngoba ayithumeli nje kuphela ukuqagela okunzima kuma-amygdala, futhi ithumela ukuqagela kuhlaka olubanzi lwe-nuclei, njengoba yonke isizinda se-amygdala esishiwo ngenhla sithola okokufaka okuvela ku-BA 25. Yize ivela njengokukhanya kukholamu B ka Umfanekiso we-6, kufanelekile ukuqaphela ukuthi i-BA 32 inikezela ngamanani achazwe kahle kuma-amygdala. Ngezici eziningi i-BA32 ibonakala i-Homologous cortex ye-prelimbic kuma-rodents (Intengo, i-2006a). Kuma-rodents, amaphrojekthi we-cortex we-prelimbic ezingxenyeni ze-basolateral kanye ne-central nucleus ye-amygdala (I-Vertes, 2004). Ezindaweni ezingezona ezomuntu, ukuqagela kubuye kwaqashelwa kusuka ku-BA 32 kuya engxenyeni etholakalayo ye-accessory basal nucleus (I-Chiba et al., 2001). Ngakho-ke, yize incane kakhulu futhi ibanzi kakhulu kunokuqagelwa okuvela ku-BA 25, i-BA 32 ibonakala isesimweni sokusebenzisana nezinqubo zokukhetha ze-amygdalar.

  

Umfanekiso we-6

Izifunda ezithinta i-Amygdala ePFC. Lesi sibalo simelela okuqhamuka ezifundweni eziningi zokulandela umkhondo kulandela imijovo engxenyeni ye-basal nucleus, i-accessory basal, medial, ne-lateral amygdala nuclei. Izindawo ezimpunga zithola okokufaka okubalulekile ...

UBarbas noZikopoulos (2006) baphikisana ngokuthi ukuphuma kwe-medial kuqala noku-OFC ku-amygdala kungaba nemiphumela ehlukile ekusebenzeni kwe-amygdalar. I-BA25 esezingeni lokuphaka ihlinzeka ngokukhipha amandla okugcabha ezingxenyeni zeasolateral ze-amygdala, zona ezinikeza ukuqagela okujabulisayo ku-hypothalamus. Ngokuphambene nalokho, i-poster yangaphandle ye-agranular ye-OFC ifaka ngaphakathi kakhulu amasokisi ahlanganyelwe azungeze i-basal nucleus (Bona I-Fig. 7). Amasokisi ahlangane ahlinzeka ngokufaka kokuvinjelayo ku-nucleus ephakathi. Lapho ivuselelwa, izinxuku ezihlangane zimisa umzila we-tonic inhibitory kusuka ku-nucleus ephakathi kuya kwi-hypothalamus, ngaleyo ndlela zibangele ukuphazamiseka kwe-hypothalamus. Ukuqagela okujabulisayo okulula kufinyelela naku-nucleus ephakathi ngqo kusuka ngemuva kwe-OFC, okuvumela i-OFC ukuthi ikhule noma inciphise ukudubula kwe-nucleus central.

  

Umfanekiso we-7

A) Ukufaka i-Amygdala ku-PFC; B) Ukukhishwa kwangaphambi kokuqala ku-amygdala; kanye ne-C) Ukulinganisa kokufaka kusuka kokuphuma kokuphuma kuya ku-amygdala. Amanani atholakala ekulebula kokucwaninga kwezifundo zobuningi nge IGhashghaei et al. (2007). Ubuningi be-Projection kanye nezilinganiso zikhonjiswa kuma-lateral, ...

Ukuqagela kokuqala kwe-hypothalamus kanye ne-brainstem

Izindawo ze-OFC kanye ne-PFC enezinqubo ezilandelwayo ezilungiselelwe i-amygdala nazo zivame ukufeza esifundeni se-hypothalamus kanye ne-autonomic brainstem / periaqueductal grey (I-et al., 1998; UBarbas et al., 2003; Intengo, i-2006b; I-Rempel-Clower neBarbas, ngo-1998), ukuhlinzeka ngekhono eliqondile lokuthonya izifunda ze-autonomic effector ezihambisana nokukhipha okungokomzwelo (bona Umfanekiso we-8). Lokhu kuqagela kubonakala kuqinile ikakhulukazi kuzakhiwo zodonga ezi-medical, kepha futhi kuqhamuka endaweni efana ne-crescent lapho kukhona okokufaka kwe-amygdalar. Njengokuntuleka kwayo kokufinyelela okuqondile kuma-amygdala, i-DLPFC ne-FP kakhulu ngaphandle kokuqagelwa okuqondile kulezi zingosi. Ngokwengeziwe, ezinye izingxenye zangaphandle ze-OFC zibonisa okuncane okukhiphayo okuqondile kulezi zikhungo ezizimele.

  

Umfanekiso we-8

Izindlela zokuqala zangaphambili zokuguqula ukuphuma kwe-amygdalar ezifundeni ezizimele. Kuguqulwe nemvume evela (IBarbas & Zikopoulos, 2006). Ukuqagela okuvusa amadlingozi kwe-OFC ezinxekeni ezihlangene (IM) (indlela a) kuholela ekubulaweni kwe-hypothalamus ...

Ukuxhumana ngaphakathi kwe-lobe yangaphambili

Njengoba kuphawuliwe ngenhla i-PFC ingahlukaniswa zibe yimikhuba emibili emikhulu yama-phylogenetic. Ukuxhumana okuphezulu kakhulu kwesifunda ngasinye kunezindawo ezisesimweni esifanayo, ikakhulukazi izindawo ezingomakhelwane ezingekho ngaphezu kwesigaba esisodwa sentuthuko kude nendawo okukhulunywa ngayo. Ngakho-ke, ngokwesibonelo, izifunda ezingama-agranular insular ezingaphakathi kwe-OFC zinokuxhumana okuningana kwezinye izifunda ze-agranular nezidysgranular orbital, kepha ngokuvamile azinayo indawo yokuxhuma kwizifunda ezingahlali njenge-46 yendawo engaphakathi kwendlela yazo. Lapho kuhlangana khona ukuxhumana kwendawo ezivamile, akuvami ukuthi kugxume izigaba ezingaphezu kwesisodwa zokuthuthuka kwezakhiwo. Isibonelo, indawo ye-isocortical ventral 46 ixhumeke ngokuqinile endaweni engasendaweni ye-46 ese-isocortical in the Mediodorsal trend, kodwa ayixhumeki ezindaweni ezakhiwe kahle zodonga lwe-medial ezifana ne-cgenate ye-subgenual (BA 46). Izindawo ezingaphezulu nangaphandle kwezindawo ze-OFC zinokuxhumana okuningana nendawo ye-25 yendawo yangaphakathi ne-46, kodwa ukuxhumeka okugxuma isembozo esiyinhloko engxenyeni engaphakathi yendawo 45 akuvamile.

Noma kunjalo, izindawo ezimbalwa ze-OFC zibonakala zinokuxhumana okuqondile ne-DLPFC. Ngokuqondile izindawo i-11m, i-12o, i-13a kanye ne-14r ngayinye inendawo yokuxhumana ne-DLPFC. I-gyrus rectus (okubandakanya indawo ye-14r) ingahle ibhekwe njengengxenye yendawo yokulalela noma njengendawo yezinguquko phakathi kwezitayela, ngakho ukuxhumeka kwayo akuvezi ukweqa okuphakathi. Kodwa-ke, izindawo i-11m, i-12o, ne-13a zihlelwe njengengxenye yomkhuba we-basoventral, ngakho-ke izixhumanisi zazo ne-DLPFC zimele ukuxhumeka kokuphakathi kwezindlela. Ukuqonda isikhundla senethiwekhi enkulu yalezi zindawo, kungasiza ukubheka enye indlela yokuhlukanisa izifunda ezi-orbital nezindondo. Esikhundleni sokusekela amamodeli we-phylogeny, UCarmichael noPaulo (1996) hlukanisa udonga lwe-OFC nodonga lwendondo lube yinethiwekhi yomgwaqo ne-medial ngokuqinile ngokuqinile emandleni wokuxhumana phakathi kwezifunda (Bona I-Fig. 9). Lolu hlobo lokuhlelwa kwesigaba lukhombisa ukunqwabelana okukhulu ngokuhlukaniswa kwe-phyolgenetic phakathi kwe-basoventral ne-Mediodorsal trends, akumangalisi uma kunikezwa inhlangano yokuxhumana esekuxoxwe ngayo. Kodwa-ke, lezi zinhlelo ezimbili zokuhlukaniswa azifani. Kuyamangaza ukuthi zonke izindawo ezi-orbital ezixhuma kwi-DLPFC noma ziyingxenye yenethiwekhi yezindondo kaCarmichael, ne-Price, noma zithathwa njengezibandakanya phakathi kwamanethiwekhi. Isibonelo, indawo eyi-11m ithathwa njengengxenye yenethiwekhi yezindondo, ngoba inokuxhumana okuningi nezindawo zodonga lwendabuko kunayo yonke i-OFC. I-Carmichael ne-Price bahlukanisa izindawo i-12o ne-13a njengezifunda ezisebenzayo ngoba ziqukethe ukuxhumana okuningana kwezindawo zezindondo nezindawo ezihamba phambili. Lezi zindlela ezihlukile zokuxhumana zenza kube sobala ukuthi kuzoba nokuphawulwa kwesifunda, noma okuphathelene nalokhu okuhlukile, emandleni we-OFC wokuxhumana nezinye izindawo ezingaphambili. Ngokukhethekile, i-gyrus rectus, kanye ne-11m, i-12o, ne-13a zisesimweni sokuxhumana nazo zombili lezi zindawo odongeni lwendondo (isib. I-cingate) nezindawo ze-DLPFC, kanti ezinye izindawo ezi-orbital zingenabo lobu budlelwano obuqondile.

  

Umfanekiso we-9

Amanethiwekhi wokuxhuma kwe-Orbital ne-Medial njengoba kuchazwe ngu-Price nozakwabo. Kuguqulwe nemvume evela (Intengo, i-2006b). Inothi Lentengo alibandakanyi i-46 yendawo ephakathi nendawo, yize idatha ikhombisa ukuthi izifunda ezingaphezulu ziyakhombisa ...

Ukuxhuma kwenethiwekhi yangaphambili kulawula izindlela eziya e-amygdala

Ezindaweni ezingekho okukhishwayo okuqondile okuphuma ku-amygdala, amandla okuthonya ukucubungula i-amygdalar kumele ancike ezindleleni eziqondile, futhi lezi zindlela ziyonqunyelwa kakhulu isikhundla sazo ngaphakathi kwamanethiwekhi amakhulu angaphambili. Ngokunikezwa amandla okuqanjwa kwe-cgenate cingulate's (BA 25) yama-amygdala, kunganikeza ukubuyiswa okubaluleke kakhulu lapho izifunda ze-PFC ezihlukene zithonya khona i-amygdala. Njengoba kungabonakala kusuka Umfanekiso we-9, I-BA 25 ithola ukuqagela okukhulu okuvela ezindaweni zenethiwekhi yezindali nezindawo ezisebusweni be-orbital ezihambisana nenethiwekhi yezindondo. Ngokuphambene nalokho, ukuqagela okwengeziwe kwe-dorsolateral kunqabile. UVogt noPandya (1987) qaphela ukuthi i-BA 25 ithola ukuqagelwa ku-DLPFC, futhi ichaze ngokuqondile okufakwayo okuvela endaweni 9 engxenyeni ephakeme ye-DLPFC. Noma kunjalo, amandla alesi sixhumanisi abonakala ebuthakathaka, futhi awakaze abonakale ngokucacile kwezinye izifundo (IBarbas & Pandya, 1989). Noma kunjalo, i-BA 9 ixhumeke kahle ne-BA 32 odongeni lwendondo, nayo exhunywe kakhulu ne-BA 25 (UCarmichael & Intengo, 1996; IBarbas & Pandya, 1989), futhi ngaleyo ndlela ihlinzeka ngendlela enokwenzeka yokungaqondile ye-DLPFC ukuthonya i-BA 25. Ngokufanayo, i-dorsal BA 46 ayinakho ukuxhumana okukhulu ne-BA 25, futhi kungenzeka ihlanganyele i-BA 32, noma mhlawumbe ezinye izingxenye ze-cingex cortex, ukuze ikwazi ukuxhumana ne-BA 25.

I-dorsal ACC (BA 24) futhi ihlinzeka nendawo ebalulekile yokuphuma ku-amygdala. Lesi sifunda sinezindlela eziningi zokufaka ezivela ku-PFC (UCarmichael & Intengo, 1996; IVogt & Pandya, 1987; IBarbas & Pandya, 1989). Lokhu kufaka okokufaka okubanzi okuvela ku-BA 9, naku-BA 46 okuncane kakhulu kwi-DLPFC, izingxenye ze-BA 32, ne-BA 10 odongeni lwe-medial, kanye nezifunda eziningana ze-OFC (ikakhulukazi izindawo zenethiwekhi ze-medial / eziphakathi ne-13a ne-Iai, ne-12o ). Ngakho-ke i-dorsal ACC ivela esimeni esiqinile sokuhlanganisa izici zomsebenzi we-PFC ezifundeni eziningi.

Yize ezifundeni ezingaphandle kwe-OFC ne-VLPFC zibonakala zinesilinganiso esikhulu sokuphuma kwe-amygdala kunokufaka okuvela kuma-amygdala, ngoba lezi zindatshana zilingana kakhulu, lezi zifunda zangaphandle zingangeza ngokungeziwe izindlela eziqondile ukwenza i-amygdala. Ezifundeni ezingaphandle kwe-OFC, lokhu kungenzeka ukuthi kuqondiswe esifundeni esingasemuva se-OFC. Ngokuphambene, kwezifunda ze-ventrolateral, indawo yangasemuva i-12l ingahlinzeka ngendlela ecacisiwe yokubandakanya i-amygdala, inikezwe isikhundla sayo esiyingqayizivele ngaphakathi kwamanethiwekhi we-intra-pre-preal ne-pre-pre-prealal-amygdala.

Amaphethini weLaminar nokuxhuma okungaphakathi kwangaphakathi

Ngokuhambisana nemodeli yokuhleleka, ukucutshungulwa kwamaphethini we-laminar wokuqagela kukhombisa ukuthi i-dysgranular OFC inezimpawu zokuphendula eziqinile ekuxhumaneni kwayo nezifunda ezithuthukisiwe ngokwakhiwa kwe-PFC (IBarbas, i-2000). Ngokufana nezinhlelo zemizwa, lokhu kungasho ukuthi ukuqagelwa kwe-OFC kuhloselwe ukulinganisa noma ukulungisa ukubala. Ngokuphikisana, i-eulaminate DLPFC inamazinga aphezulu kakhulu wokuphakanyiswa, okukuvumela ukuthi kondle imiphumela noma umphumela wokuqalwa kwayo kuya ezifundeni zobuchopho ezalandela. Le iphethini ejwayelekile yokudla okuphindwe kabili okuphindwe kabili futhi okubonakalayo nakho ukuxhumana okukhona phakathi kwe-OFC ne-DLPFC. Ukuxhunyaniswa kwangaphambili kwe-OFC kusuka kakhulu ezingxenyeni ezisondele kakhulu ze-cortical (2-3) kanti ama-axons awo aqeda ngokuyinhloko ezingqimbeni ezijulile (i-4-6), ehambelana nephethini yokudla okuphambili (IBarbas & Rempel-Clower, ngo-1997). Ngokuphambene nalokho, ukuqagelwa kwe-OFC ku-PFC yamuva kuvela ikakhulu ezingxenyeni ezijulile (i-5-6) ngama-axons ayo aqeda kakhulu ezingxenyeni ezingaphezulu (i-1-3), isici sokuphendula. Leli phethini lisebenza lisebenza cishe ku-70-80% yemibono. Ngakho-ke, ukugeleza kolwazi kusuka ku-OFC kuya ku-PFC ebumbene kuqukethe impendulo, kanti imininingwane yokugeleza kolunye uhlangothi ifana kakhulu nephethini yokuphakelayo.

Imodeli yokuhleleka ivusa inkanuko ngoba iphakamisa ukuthi ubunjalo bezokuxhumana phakathi kwezifunda bungafakwa ngokususelwa ekuxhumeni kwe-laminar. Uma imodeli yokwakheka ilungile, kuyasiphoqa ukuthi sibheke kuhlobo lokuphakelayo nolokuphendula ngokuxhumana phakathi kwesifunda. Amamodeli aphakamisa ukuthi i-PFC yamuva isebenze ngokuyinhloko noma ngokukhethekile ngokusetshenziswa kwezinqubo ezisezingeni eliphakeme kunzima ukuzivumelanisa nezici zayo ezivelele zokuphakela. Ngokufanayo, amamodeli we-OFC abheka lokho njengokudlulisela imiphumela yekhompiyutha (ngokwesibonelo senani lomvuzo) ku-PFC esetshenziswayo, ehluleka ukubamba amandla wesifunda wokubala okubandayo okwenziwa ku-PFC eyalandela. Kodwa-ke, njengoba kuchaziwe ngokuhamba kwesikhathi kule ndatshana, izindlela ezikhona zokusebenzisana phakathi kwezifunda ze-PFC, naphakathi kwezindawo ezibandakanyeka “ngokomzwelo” ngokuqhathaniswa nokusebenza “kwengqondo” zihlala zingakunaki lokho okungenzeka kuyimodeli yokwakheka. Impela, amamodeli akhona ngokujwayelekile abamba imisebenzi ye-PFC yamuva, ikakhulukazi imisebenzi ye-DLPFC, ngokulawulwa okuphansi, futhi akuvamile ukuthi abheke ukuthi izindawo ezingathuthukiswa ngesakhiwo ezinjenge-OFC zinganikeza ithonya eliphakeme kakhulu ezifundeni ze-PFC ezizayo.

Amaphethini weLaminar wokuxhuma okungaphambili kwe-amygdalar

Ukuqagelwa kwe-amygdala engemuva kwe-OFC kufaka zonke izingqimba ze-cortex, futhi ngenxa yalokho kungenzeka kungagcini ngokuqinile kumaphrojekthi we-feedlelo noma impendulo yohlobo (I-Ghashghaei et al., 2007). Kodwa-ke, kuyacaca ukuthi kunengxenye eqinile yokudla kulokhu ukuqagelwa okusekelwe ekuqedeni i-laminar. Ngokuphikisana nalokho, ukuqagelwa kwe-OFC kuma-amygdala ngokuyinhloko kuqhamuka kungqimba 5, okubonisa isimo sawo njengokuqagelwa kwempendulo (kusikisela ukuthi basebenza ekusekelweni kwe-amygdalar kunokuba kudluliselwe imininingwane ethile efana nezimpawu zomqondo zesisusa). Ngokuthabisayo, ukuqagelwa okuphindaphindekayo okuvela kwi-CCC ye-lateral kuqondiswe kungqimba lwe-5 lwe-OFC, okuyingqimba yokuqala yokuphuma lapho ukuqagela kwe-OFC kuya e-amygdala kuvela.

Ingabe ukuqonda kwe-anatomical kungasekela izingxoxo zemisebenzi yokuqala? Iminyaka eminingana edlule ibone ukuqhuma kwesithakazelo ngendlela izindawo ezahlukahlukene zobuchopho ezixhumana ngayo. Le ntshisekelo ivele ngokwengxenye ngenxa yokuvela kwamasu wokuhlola ukuxhumana okusebenzayo ne-fMRI, ukuhlinzeka okokuqala amandla okuhlola ukusebenzisana phakathi kwezifunda zobuchopho kubantu abaphilile. Kodwa-ke, izingxoxo zale datha, kanye namamodeli avela kule mininingwane, bekungakaze kucindezelwe yi-anatomy. Njengoba lezi zinhlobo ziye zaba nethonya ngokwengeziwe, sikholwa ukuthi kungasiza ukuhlola ukuthi zihambisana kahle kanjani neuroanatomy echazwe ngenhla. Sikholwa ukuthi amamodeli anjengalawa adinga ukuhambisana nawo womabili amaphethini wokuxhumana ahlanganayo ahlanganisa izifunda ezihlukile zecortical kanye ne-subcortical kanye nesimo sokuphendula / sokuphakelayo kwalezi zindlela. Lapho amamodeli engahambisani nalezi zingqinamba, ahluleka ukubonakala, noma okungenani adinga incazelo yokuthi angasebenza kanjani ngokunikezwa ukungahambisani kwawo nokuxhumana kwengqondo okwaziwayo.

Izincwadi ezikhulayo ngokwengqondo zizama ukuqonda indlela izinqubo “ezinengqondo” ezisebenzisana ngayo nezinqubo “zemizwa”. Ngenkathi kukhona imikhawulo eqondile yokuhlukaniswa okwenziwe phakathi kwezinqubo zokuqonda nezomzwelo (I-Pessoa, 2008), umehluko ubonakale ulusizo ekuboniseni uhla lwezimilo ezinjengokulawulwa kwemizwelo, ugqozi, izinqumo zomnotho kanye nokuqondiswa kwezindlela zokunaka. Ezigabeni ezilandelayo, sichaza idatha kanye namamodeli avelayo wokulawulwa kwesimo, inkumbulo yokusebenza nokusebenzelana kwangaphambi kwekhaya, kugxilwe ekuvumelaneni kwabo nedatha ye-anatomical. Sigxila kakhulu ezincwadini zokulawula imizwa, njengoba lezincwadi ziya ngokuya zanda ezingxoxweni ze-psychopathology kanye nokwelashwa kwengqondo.

Izifundo ze-Correlational of deactivation ze-amygdala

Ukuze uqonde kabanzi ukuthi umthethonqubo we-top-phansi uxhumana kanjani ne-amygdala, isethi yezifundo zokulawulwa kwemizwelo idlulele ngaphezulu kokuhlukahluka komsebenzi wokuphenya ukuphenya okuqondile kokuncipha komsebenzi we-amygdala (Bona Ithebula 2). Lokho ukusho ukuthi, esikhundleni sokubuza ukuthi yiziphi izindawo ezenza umsebenzi owaziwa njengokulawula phansi ukusebenza kwe-amygdala, bavivinya ngokusobala ukuxhumana noma ukusebenza / ukuxhumana okusebenzayo phakathi kwe-amygdala nalo lonke ubuchopho ngesikhathi sokusebenza komthetho wokuzwela. Ngenye indlela, ezinye izifundo ezihlanganisiwe ze-amygdala ziyancipha kanye nezifunda ezisetshenzisiwe ezisetshenzisiwe kusukela ekuqhathaniseni komthethonqubo omkhulu. Lolu cwaningo lukhombisa ukuthi ukuncipha kwe-amygdalar kuhlobene kabi nezindawo eziningi zomsebenzi we-PFC. Okuqaphelekile ikakhulukazi ukusebenza kwe-VMPFC, kufaka phakathi i-BA 11m / 14r (5, 37, −12; −6, 46, −20: Urry et al., 2006, U-Ochsner et al., 2002 ngokulandelana). Ngaphezu kwalokho, izifunda ezingaphili kanye nezangaphambilini zokwe-cingate zaqapheleka ukuthi zihambisana kabi nomsebenzi we-amygdala ngesikhathi somthethonqubo. Ngokwesibonelo, U-Urry nozakwabo (2006) ubike isifunda se-BA 32 / 10 (esiphezulu ku-−23, 43, −10) enwetshiwe ngokwedlule nangokwangaphakathi. UDelgado et al. (2008b) futhi ubike ukuhlangana okungafani phakathi kwe-BA 32 (0, 35, −8) kanye ne-amygdala iyancipha. Izindawo ze-Posterior (BA 13) ze-OFC nazo zihlobaniswe kabi nge-amygdala deactivation (−24, 28, −14; 26, 24, −22: Banks et al, 2007: −30, 22, −16; I-34: U-Ochsner, uRay et al., 2004). Izindawo ezingaphansi kwe-PFC ezise-BA 47 (34, 54, 12) kanye ne-BA46 (−54, 12, 12: Urry et al., 2006; I-−54, 42, 12: U-Ochsner et al., 2002), futhi kwavela kulezi zifundo. Izifundo ezimbili zixhumanise izifunda ezithile ze-DLPFC ezifundeni ezithile, ezihambelana nokuncipha kokuphendula kwe-amygdala. Esifundweni sika Urry et al. (2006), ukuhlaziywa kokulamula kubonise ukuxhumana phakathi kwe-amygdala, BA 10 (3, 63, 18) kanye nesifunda se-DLPFC (−50, 23, 19). UDelgado et al. (2008b) kungenjalo basebenzise i-medial BA 32 esifundeni njengembewu yokuhlaziywa kwe-PPI yabo eyabe ikhomba isifunda sase-amygdala nesifunda se-DLPFC. Ngokusemqoka, lolu cwaningo lukhomba izifunda ezihambelana nama-amygdala okwehla okuye kwaqalwa nakho ngaphezulu njengokuqagela kuma-amygdala anjenge-dorsal anterior cingulate, cingrate engaphandle kanye ne-posterior orbitof Pambal cortex.

  

Ithebula 2

Izifundo ezibika ukuhlangana phakathi kokuncipha komsebenzi we-amygdala nesifunda sangaphambi kokukhuphuka kuyanda phakathi kwemisebenzi yokulawula imizwa.

Ezifundeni ezibikiwe kusuka kulokhu kuhlaziya okuhlanganisiwe noma ukuhlaziya okuningi kokucwaninga, inani elilinganiselwe lazo linokuxhumana okuqondile okuqondile kwi-amygdala. Izifunda ezivame kakhulu ezihlobene ngokungaphendulwanga nempendulo ye-amygdala yizifunda ezingemuva kwe-OFC kanye ne-cingulate engaphansi kwe-VLPFC (Umfanekiso we-12). Ezifundeni ezilandelwayo zangaphambi kokuphela kwengxenye ye-BA 47 / 12 yamuva kuphela enengqungquthela eqinile ye-amygdala. Izindawo ze-BA 32 zangaphandle nazo zikhonjwa ekuhlaziyweni kokuhlobanisa, okungakhombisa ukuqagela ku-testal and basal lateral nucleus ye-amygdala (UCeba et al, 2001).

  

Umfanekiso we-12

Abaxhumanisi bakhonjwe ngaphakathi Ithebula 2 njengoba kuhlobene nokusebenza okwenziwe ku-deictivations ku-amygdala ngesikhathi sokulawulwa kwemizwelo okuhlelwe ngaphezulu kwengqondo yethempulethi (phezulu kwesokunxele nakwesokudla) futhi kunikezwe kwingqondo yengilazi (ukubuka okuphansi nokubuka kwesobunxele). Amamaki e-cyan ...

Amamodeli okulawulwa kwemizwelo

Kuze kube manje, imodeli edonsa kakhulu yedatha yokulawulwa kwemizwelo ivela ocwaningweni lokuphinda kuqalwe kabusha ngu I-Wager nozakwabo (2008). Umphumela wokuhlukahluka wenzalo ukuguqulwa kokuthinteka okungekuhle okubikwe ngokwakho. Indlela esetshenziswayo yokulinganisa isetshenziswe kudathabheyisi ye-neuroimaging evela kupharadesi yokuphinda ifane nale esetshenziswa U-Ochsner et al. (2002; 2004). I-VLPFC eyiyo yakhethwa njengesiqalo sokuhlaziya, ngokuxhumanisa okugxilwe endaweni okubandakanya kuyo ingxenye yengxenye yendawo ye-47 / 12 ngokuqagelwa kwe-amygdala. Ababhali baqale basebenzisa indlela ye-ROI ukuvivinya indima yama-amygdala nama-nucleus accumbens njengabalamuli phakathi kwe-VLPFC yangakwesokudla futhi banciphisa umthelela ongemuhle owakhonjwa njengethimba eliphambili lokuphumelela kabusha. Kulolu hlaziyo lwe-ROI zombili izinhlaka zaboniswa ukulamula ubudlelwano phakathi kwe-VLPFC yangakwesokudla kanye nokwehla kokuzibika kokuthinteka okungalungile (bona Umfanekiso we-13).

  

Umfanekiso we-13

Umdwebo wokuhlaziywa kokulamula uvivinya ubudlelwano phakathi kwe-VLPFC yangakwesokudla futhi wehlise kokubi kuthinta okuthethwe kusebenze ku-amygdala kanye ne-nucleus accumbens. Umdwebo uguqulwe nemvume evela UWager, uDavidson, uHughes, uLindquist, ...

Ababhali babe sebesebenzisa ukucubungula kwengqondo yonke kanye nokuqonda okungatheni uhlangothi ukuthola izindlela ezimbili zokuxhumana phakathi kwe-VLPFC kanye nezinguquko ekuthini zithinteke ngendlela engeyiyo (bona Umfanekiso we-14). Inethiwekhi eyodwa inokukhetha okungaqondile okuqondile kokukhulisa ushintsho kokuthinta okungalungile. Le nethiwekhi ifaka izifunda zama-nucleus accumbens, subgenual cingulate (BA 25), i-pre-SMA, i-precuneus, i-DMPFC (MNI: 24, 41, 40), ne-front front gyrus (24, 21, 58). Phakathi kwalezi zifunda, ama-nucleus accumbens kanye ne-cingrate engaphansi kwe-infinual ukuxhumana kakhulu ne-amygdala. Inethiwekhi yesibili ekhonjwayo inokukhetha okungaqondile okungaqondile ekunciphiseni ushintsho kumthelela omubi nasekunciphiseni ukuphumelela kabusha. Le nethiwekhi ifaka i-rostral dorsal ACC, amygdala (amazwe amabili) ne-posterior-lateral OFC (48, 24, −18). Umsebenzi wekusasa kuzodingeka ucacise ukuthi izingxenye zamanethiwekhi zisebenzisana kanjani nokuthi amanethiwekhi ahlobene naluphi uhlobo lwecebo lokuqondisa imizwa.

  

Umfanekiso we-14

Imodeli yendlela yenethiwekhi ebandlulula ngokuphelele kwinethiwekhi ephuzi futhi engakhethi kahle ngokuluhlaza okuluhlaza phakathi kobudlelwano phakathi kwe-VLPFC kanye nokwehla kokuthinteka okungekuhle okubikiwe. Umdwebo uguqulwe nemvume evela UWager, uDavidson, uHughes, uLindquist, ...

Abaphenyi abaningana bakhiphe amamodeli wethiyori mayelana nezindlela zemvelo ezisetshenziswayo zokulawula imizwelo. Elula lala amamodeli liphakamisa ukuthi inani elilinganiselwe lezindawo linomthelela oqondile kuma-amygdala. UDelgado et al. (2008b), UHansel noVan Kanel (2008) futhi I-Quirk ne-Beer (2006) isiphakamiso ngasinye sokuthi i-podC ye-ventromedial phansi ilawula izifunda ze-amygdala. Lezi amamodeli kubalulekile ukuzama ukuqonda kwethu izisekelo ze-neuroanatomical zokulawulwa kwesimo somzwelo wabantu ezincwadini zezilwane ezinwebekayo ekuqothulweni kanye nokuxhumeka kwe-PFC okuxhumeka kubantu abaningi abahlanganisiwe e-basolateral amygdala (UMorgan, uRomanki noLeDoux, ngo-1993; Likhtik et al., 2005; I-Quirk et al., 2000). I-Quirk ne-Beer (2006) yakhela phambi kobukhona bokuthi kube nemiphumela ethokozisayo nethikamezayo yokuqokwa kwe-PFC ye-"ventral" kuma-amygdala kubantu kanye namagundane. Isifunda esingaphandle kwesisekelo se-cingrate, i-BA 25, kuthiwa singavimbi kakhulu ngenkathi i-dorsal ne-anterior BA 32 kuhlongozwayo ukuthi kube nokuxhumana okujabulisayo ne-amygdala. Bobabili i-BA 25 ne-32 inokuxhumana ne-amygdala. I-BA 32, noma kunjalo, inokuxhumana okukhawulelwe okuningi.

Phillips et al (2008) bakhiqize imodeli yesekethe ezizama ukuchaza ukuqonda okungaphansi kwe-neural kwezinhlobo eziningi zokulawulwa kwemizwelo (bheka Umfanekiso we-15). Imodeli iqukethe izingxenye zezigaba ze-DLPFC, OFC, VLPFC, DMPFC ne-ACC. Okuthakazelisa ngokukhethekile, ababhali bahlukanisa phakathi kwezindawo ezibandakanyeka emthethweni othomathikhi wokulawula imizwa (ku-Agengenation ne-rostral ACC) kanye nezifunda eziqoqelwe ukulawulwa kokuzithandela (DLPFC kanye ne-VLPFC). Babiza lezi zifunda zokugcina njengezintsha ngokwedlule nangokunikeza impendulo ezinqubweni zakudala zokuzala zemizwa. I-OFC, i-DMPFC kanye ne-ACC, ngakolunye uhlangothi izifunda ezindala ezinama-phylogenetically ezichazwa njengezisebenza ngezinqubo ezilandelwayo zokudlulisa imininingwane yezwe yangaphakathi kwi-DLPFC ne-VLPFC. Ababhali babeka i-DMPFC njengomugqa lapho i-OFC idlulisela khona ulwazi oluyimfihlo phambili ezifundeni ezinobuchopho izinqubo zezinqumo.

  

Umfanekiso we-15

Phillips et al. (2008) imodeli yokusebenzelana kokuqala kwe-amygdala a) I-OFC, i-ACG engaphansi (ACC), kanye ne-rostral ACG (ACC) imininingwane ehambelana ngqo ne-MdPFC kanye nasezifundeni ze-PFC ezilandelwayo ukuthatha isinqumo nezinyathelo. B) Izinqubo zempendulo ezivela ku- ...

Isici esisodwa esiyingqayizivele kulo modeli ukucaciswa okucacile kwezinqubo ze-feedxp kanye nempendulo. Imodeli iyathandeka futhi inamqondo ophelele ngemibono yendabuko nge-DLPFC esebenzisa ukulawula okungaphezulu kwezifunda "ezinemizwa" engaphezulu. Kodwa-ke, kunzima ukuvumelanisa le conceptualization nemodeli yesakhiwo, uma kunikezwe ukusatshalaliswa okuxhunyiwe kwe-PFC ukuxhumana (IBarbas & Rempel-Clower, ngo-1997; IBarbas, i-2000). Impela, imodeli yokwakheka isikisela ukuthi imininingwane yokugeleza phakathi kwe-DLPFC ne-OFC empeleni isephambili nenqubo evela ku-OFC nokuya e-DLPFC kubonakaliswe njengempendulo, kanye nalabo abasuka eDLPFC futhi abaya e-OFC ngokuyinhloko kubhalwe njengo-feedirect.

Phillips et al. imodeli nayo iyaphawuleka ekubekweni kwayo kwezifunda ezibizwa nge- “othomathikhi mithetho” njenge-cgenrate engaphansi kanye ne-OFC njengomgwaqo wokuqala lapho izifunda ezintsha kakhulu zephylogenetically zithinta izindawo ezinamandla njenge-amygdala. Lokhu kuyavumelana kakhulu (ikakhulukazi nesifunda sangokunye kokuhlangana) ngamalungiselelo wenethiwekhi achazwe ngenhla. Kungacatshangwa, noma kunjalo, ukuthi kungahle kube nendlela engaphezu kweyodwa lapho izindawo zokulawula imizwa yokuzithandela zingathinta khona ukucubungula kwe-amygdala. Ikakhulu, i-VLPFC yangemuva ingakwazi ukuthonya ngqo izinqubo ze-amygdala ngaphandle kokudinga ukubandakanyeka kwesinye sezifundazwe ze- "othomathikhi okuzenzakalelayo", inikezwe okufakwayo okuqondile ku-amygdala nuclei.

Ngokufingqa, ingcebo yemininingwane ikhombisa ukubandakanyeka kwezifunda ze-PFC ngesikhathi semisebenzi yokulawula imizwa, ngomsebenzi eqenjini elikhethiwe lezindawo (i-BA 47 / 12, BA25 ne-BA 32) ekhombisa izinhlangano ezinekhono lokulawula umsebenzi we-amygdala. Kuvuselelwa amamodeli asezingeni eliphakeme ukuchaza le mininingwane. Ukuvela kwalezi zinhlobo kuyathandeka, njengoba nokukhathazeka okukhonjiswe ngababhali babo ngokwenzelela kwamanethiwekhi ahlangene ahlongozwayo. Siyazi, noma kunjalo, ukuthi azikho amamodeli kuze kube manje ezivumile ngokucacile iphethini yokuxhuma okuphakathi kwezifunda ze-PFC ezahlukene. Ngokwesibonelo, U-Wager et al (2008) inikeza imodeli eyinkimbinkimbi kakhulu yesu elithile lokulawula imizwa, kepha alisho uhlobo lokugeleza kolwazi phakathi kwezigaba zezingxenye. Phillips et al. faka ngokucacile umqondo wolwazi lwe-feedxp kanye nerekhodi, kepha ungahambisani nale mibono ngephethini ebanjiwe yokuphendula kanye nokuqagelwa okuphakanyisiwe ezifundeni okukhulunywa ngazo. Sikholwa ukuthi ukuhlanganisa lezi zinkinga kunikeza enye yezinselelo ezibalulekile kubacwaningi abazama ukuqonda imigatshana ye-neural sub Rule.

Ukucindezelwa kokuthinteka ngokomzwelo ngesikhathi somsebenzi wokuqonda

Izifundo eziningi zisebenzise ubuhlakani lapho abahlanganyeli kufanele baphendule esivumelwaneni esisebenzayo esingahlobene nesikhuthazo (njengombala) futhi bangaphazanyiswa okuqukethwe yimizwa (ie, amagama emizwelo), noma ukubambelela ekuvuseleleni okungekho emthethweni (ie a indlu) ngenkathi unganaki ukuvuselela okungokomzwelo (ubuso obesabekayo). Isibonelo, izifunda ze-rostral (dorsomedial, pregenual and dorsal ACC) nazo zombili izifunda ze-DLPFC kanye ne-VLPFC konke kubhekwe kuzimpawu zesimo somoya lapho izifundo kufanele zigweme ukuphazamiseka ngokuqukethwe yimizwa yamagama (I-Whalen et al., 1998; ICompton et al., 2003; UHerrington et al., 2005; UMohanty et al., 2007). Ukubuyekezwa okuningiliziwe kokuthi ukuphathwa kokunakwa kanye nokulawulwa kwemizwelo kungabandakanya kanjani abafundi bezimpawu ezingezansi ze-neurocognitive okukhulunywa ngazo UBlair noMitchell (2009) futhi I-Mitchell (2011).

Ukukhawulelwa kokuhumusha kweziningi zalezi zinhlaka kuphakama kodwa akuhlali kucace ukuthi lezi zifunda zenziwa yini ngoba zisebenzisa ukulawula ingqondo, ukuqapha ukungqubuzana, kubandakanyeka ngenxa yengxabano / ukuphazamiseka okukhulu ngaphandle kokulawula ukungqubuzana / ukuphazamiseka, noma bamane baphendula esimweni esingokomzwelo sesisusa. Ngokwesibonelo, UMohanty nozakwabo (2007) bonisa ngokubonisa ukuthi isifunda sangaphambi kokuhlangana (cishe i-BA 24 / 32) sikhombisa ukwanda kokusebenza ngesikhathi somsebenzi we-Stroop ngamagama atholakalayo, nokuthi lokhu kuyahambelana nesikhathi esengeziwe sokuphendula emsebenzini. Lokhu kungahunyushelwa ngokuya nge-RACC ebandakanyeka ukuze kunikezwe amandla wokucabanga ngokuqondisa abaphazamisi abanemizwa. Noma kunjalo, kunikezwe ukuthi ukusebenza kwale ndawo kuvumelana nesikhathi sokuphendula esikhulu, izinga lokwenza kusebenze akubonakali lihlobene nokuvinjwa okuphumelelayo kwabaphazamisi. Ngaphezu kwalokho, kukhombise ukuhlangana okusebenzayo kanye ne-amygdala, okusobala ukuthi akuhambisani ne-hypothesis yokuthi i-RACC yayiqhuba umthetho-jikelele we-amygdala. Impela kuyaphawuleka ukuthi ababhali baphakamise ukuthi esikhundleni sokubonisa ukumiswa kwe-RACC kwe-amygdala, ukuxhumana okuphezulu ngesikhathi sokuchayeka kweziphazamiso ezingokomzwelo kungakhombisa ukulawulwa kwe-amygdalar noma okufakwayo ku-RACC, kunokuba kunenye indlela eyindilinga.

Phakathi kwezingcezu ezibucayi kakhulu zobufakazi zokulawulwa kokuqonda kwengaphambili kokulungiswa kwemizwelo e-amygdala kuvela ocwaningweni U-Etkin et al. (2006), lapho ababambiqhaza benze khona umsebenzi ofana nse Stroop lapho ukubukeka kobuso obungokomzwelo kungahle kube kokuhlangene noma kungasebenzi ngamagama aqukethe umzwelo. Ukwakheka kwalolu cwaningo bekuyinkimbinkimbi njengoba ababhali bebengagxili ekuqhathaniseni okulula kovivinyo olungathathi hlangothi noma izivivinyo ezingezinhle ezilinganayo, kepha bahlola imiphumela ngesikhathi sokulingwa okungalungile okulandela ngqo uvivinyo oluhle noma olungalungile. Kuyamangaza ukuthi i-DLPFC, isifunda se-DMPFC endaweni ephakeme yangaphambili, kanye ne-rostral (pregenual) i-ACC ikhombise ukwenziwa kwezinto ngesikhathi sokuvivinywa okungalungile okuncike ekutheni ucwaningo lwakudala lwalunenhlonipho yini noma cha. I-DLPFC (kanye ne-DMPFC) baphendule kakhulu kwizivivinyo ezingalungile ezalandela ukuqulwa kwecala eliphelele, kuyilapho i-RCCral ACC isabele kakhudlwana kwizivivinyo ezilandela olunye uvivinyo olungafanele. Lolu cwaningo ngesinye sezifundo ezimbalwa ezincwadini zokulawulwa kwengqondo ezihlolisisa ubudlelwane bendawo zangaphambi kwecortical cortical nomsebenzi we-amygdala (kusetshenziswa ukuhlaziywa kwengqondo yokusebenzisana kwengqondo, UFriston et al. I-1997). Ngokumangazayo, umsebenzi omkhulu ku-rostral ACC wayehlotshaniswa ngobuhlakani nomsebenzi we-amygdala. Ngokusekelwe kuphethini yezimpendulo ze-amygdala, ababhali bathi umsebenzi we-amygdala uhambisana nezinga lokungqubuzana kwecala elinikeziwe, nangokucindezela umsebenzi we-amygdala i-rostral ACC ihlinzeka ngokulawula lokhu kungqubuzana. Ukusekelwa kwalo mbono kuqhamuka kwidatha yokuziphatha kokuthi labo abakhombise ukuxhumana okuphezulu kokusebenza okulinganayo kwizivivinyo ezingenangqondo bakhombise ukulungiswa okukhulu kwengxabano njengoba kukalwa ngezikhathi zokuphendula emsebenzini. Esifundweni esilandelwayo U-Etkin et al. (2010) waphawula ukuthi lokhu kucindezelwa kwemisebenzi ye-amygdala kubonakala kubuthakathaka ezigulini ezinenkinga yokuphazamiseka kokukhathazeka okuhlobene nokulawulwa okunempilo, okunikezela ngokulungiswa okungaphakathi kwe-neural kobunzima bokulawula ukuphazamiseka okungokomzwelo noma izingxabano kulaba bantu abagulayo.

I-bakoat ebalulekile ifanelekile maqondana nale ncwadi. Okokuqala, ucwaningo olwenziwe yiqembu lika-Etkin alukhuthazi ubukhona bokuvinjelwa komhlaba wonke we-amygdala yizifunda ze-PFC ngesikhathi semininingwane engqubuzanayo, noma ukubandakanyeka okuqhubekayo "kwezifunda zokulawula okuqondayo", kepha kunalokho umsebenzi othize ovinjelwe kuncike kuzinga ukungqubuzana phakathi kokukhuthaza ngokushesha. Uma kunembile, amandla okubheka ukuzihlanganisa okungafani phakathi kwe-cingate yangaphambili (noma ezinye izifunda ze-PFC) ne-amygdala kungaba ngumsebenzi wokuhlaziya kakhulu.

Eminye imigqa yobufakazi iphinde iphakamise ukuthi ezinye izindawo zangaphambili, ikakhulukazi i-dorsal ACC, zingakwazi ukulawula ukubuswa kwe-amygdala. Isibonelo, ocwaningweni usebenzisa i-paradigm efanayo ne-Etkin et al. (UChechko et al., 2009), iziguli ezinenkinga yokwethuka zikhombise ukwehla okukhulu kunokulawulwa okunempilo ngesikhathi sokuvivinywa okungahambi kahle ngokomzwelo, kanye ne-amygdala ephakeme, kepha umsebenzi ophansi we-dorsal ACC / DMPFC, okuholela kusikisela sokuthi ukwethuka komoya kubonakaliswa ukulawulwa okungalungile kwe-DMPFC / dorsal ACC. Ngokufanayo, UHariri et al., (2003) wabona ukuhlangana okungahambi kahle phakathi kwe-amygdala ne-dorsal ACC (ne-VLPFC) lapho izifundo kwadingeka ukuthi zifake ilebula vs. ukufanisa izithombe ezingokomzwelo (ngomsebenzi we-amygdala owanda ngesimo somdlalo, kanye ne-VLPFC kanye nomsebenzi we-dorsal ACC okwandayo ngesikhathi sesimo sokulebula). Kuphakanyisiwe nokuthi i-dACC ingahle ibe namandla okulawula ama-amygdala noma ngabe kungekho ukungqubuzana okuthile noma ukuphazamiseka kwemizwelo komsebenzi. Pezawas et al. (2005) kubone ukuzihlanganisa okuphambili phakathi kwe-dACC nomsebenzi we-amygdala ngesikhathi somsebenzi wokusondeza ubuso besongo. Kungaphawulwa futhi ukuthi i-ACC engaphansi ocwaningweni lwePezawas yayihlotshaniswa kahle nomsebenzi we-amygdala, iphakamisa ukuhlangana okuhlukile phakathi kwezindawo ezihlukile ze-cingate kanye ne-amygdala, nokuphakamisa okunye, njengasekuqaleni UMonhaty et al. (2007) iphepha, ukuthi i-racC, okungenani kwezinye izimo iyakhanya, kunokuba imbi, ihlanganiswe ne-amygdala.

Ukusebenza Kwememori

Okunye ukuhlolwa kokulawulwa kokuqonda okugxile emandleni okucindezela ukuphazamiseka okungokomzwelo ngesikhathi sokusebenza kwememori yokusebenza. Ngoba inani lolwazi olungabanjwa futhi lusetshenziswe ku-inthanethi lilinganiselwe (I-Cowan, i-2010), kubalulekile ukuthi abantu babeke phambili ngendlela efanele ukuthi yiluphi ulwazi olungena kulesi sitolo se-inthanethi. Ngokufanelekile, kufanele sigcine imininingwane efanele umgomo efanelekile ephathelene nemininingwane engabalulekile kangako, kodwa futhi sikwazi ukulahla okuqukethwe kwimemori yokusebenza lapho ulwazi olubaluleke kakhulu lufaka imigomo yangaphambilini. Kanjalo, inkumbulo yokusebenza ihlinzeka ngesizinda esingahle siwusizo ekuhlolweni kokusebenzelana kokuqonda, ngokunikezwa indima ebalulekile ye-DLPFC ne-VLPFC kwinqubo yememori yokusebenza (I-Badre et al., 2005; Blumenfeld et al., 2010; UCurtis & D'Esposito, 2004; UJonides et al., 2005; I-Levy & Goldman-Rakic, 2000, UNee noJonides, 2010; Iposi, i-2006; UThompson-Schill et al., 2002).

Imibiko emibili eyenziwe ngabakwaDolcos nozakwabo bayaphawuleka njengoba bexhumanisa ukusebenza kwengqondo ngokusebenza okuphumelelayo noma badonsa izingqinamba zokuxhumana okusebenzayo (UDolcos noMcArthy 2006; I-Dolcos et al., 2006). Yomibili le mibiko ihlaziye idatha kusuka kwento elula yokuphendula ukubambezeleka yokuphendula lapho izithombe ezethusayo noma ezingathathi hlangothi zivezwa phakathi nesikhathi sokulibaziseka (ukulungiswa) komsebenzi. Ocwaningweni lokuqala, bakhombisile ukuthi i-ventrolateral cortex (BA 45 / 47) ivulwe ngaphakathi ngesikhathi sokuthinteka ngokomzwelo nabaphazamisi abangathathi hlangothi. Ababambiqhaza abakhombise umsebenzi omkhulu wokuhamba kwamaphaphu lapho kukhona abaphazamisi abanemizwelo balinganise labo baphazamisayo ngokungaphazamisi okuncane. Ocwaningweni olwalandelayo bakhombisa ukuthi umsebenzi we-BA 45 oshiywe (kodwa akulungile i-BA 45) wehlukaniswe phakathi kovivinyo lapho abantu baphumelele ngempumelelo.wayengakunaki ukusiphazamisa (njengoba kuboniswe ukusebenza ngokulungile noma okungalungile kokulibaziseka kokusebenza). UDolcos et al. (2006) futhi babike nge-VLPFC- amygdala uxhumano olusebenzayo, futhi zombili lezi zindawo zanda phakathi kokulingana okungokomzwelo nezilingo zokuphazamisa ukungathathi hlangothi. Ngokubalulekile lokhu kuxhumana kusendleleni enhle futhi akunakuhunyushwa njengokubonisa ukucindezela kokudubula kwe-amygdalar.

Izifundo ezenziwe ngabakwaDolcos nozakwabo nazo zinikeza ubufakazi bamaphethini angahlukeki wokusebenza nokwenza usebenze kuzo zonke izifunda ezingaphambili. Ngokukhethekile, izindawo ezithutha umoya zanda ngeziphazamiso ezingokomzwelo, ngenkathi i-DLPFC (BA 9 / 46) incipha, iphakamisa ubudlelwano bokubuyelana phakathi kwalezi zifunda. Lobu budlelwano bokuphindisela bufana nephethini yangaphakathi ehlukanisiwe yangaphakathi Perlstein et al. (2002) ababenezifundo ezenze umsebenzi wokukhumbula okusebenza kuzo lapho izithombe eziqinisekiswe ngokomzwelo zivela khona njengezifanele kanye nezinqubo ezifanele [ngokuthokozisayo, ubudlelwane bokubuyisa buhlobene kakhulu nobumbano ne-DLPFC ekhuphuka ngezindawo ezivuthisayo nezishubile (BA 10 / 11) ekhombisa umsebenzi owandisiwe kwesikhuthazo esingesihle]. Iphethini ehlukile phakathi kwezifunda ze-dorsal kanye ne-ventral PFC nayo ibonwe kwezinye izinhlaka zokusebenza kwememori yokusebenza, nge-DLPFC enkulu ehlobene nomsebenzi wangaphandle ohambisana nomthwalo wememori omkhulu osebenzayo (I-Rypma et al., 2002; UWoodward et al., 2006), noma izifunda ezithile ze-ventral PFC ezibandakanyekile kulezi zifundo ziyehluka. Iphethini ebonakalayo eguquguqukayo yezifunda ezingaphakathi kanye neyamadolobha iveza ukungqubuzana okuphikisanayo phakathi kwalezi zifunda, kepha ungakhombisi isimo sobuhlobo. I-Ranganath (2006) iphakamisa ukwakheka kwe-hierarchical izinqubo zememori zokusebenza lapho izifunda ze-caudal / ventral PFC zinikezela ngokulawulwa okuphansi kwezinhlelo zangemuva, ngenkathi i-dorsal / rostral PFC inikezela ukulawula kwezifunda zangaphambili ze-caudal. Ngaphakathi kwalombono, iRanganath ithi izinqubo zokukhetha ezisetshenziswe yi-rostral / dorsal PFC zibandakanya ukuguquguquka komsebenzi ku-caudal / ventral PFC. Kodwa-ke, njengoba kuchaziwe ngezansi, ukuguquguquka kolunye uhlangothi nakho kucatshangelwa.

Imininingwane ye-Emotion Regulation

Ngokusekelwe esimweni sokukhetha semigwaqo ye-neuroanatomical phakathi kwe-PFC ne-amygdala, amamodeli asobala wokushintshwa kwe-PFC ngesidingo kumele abandakanye ukushintshanisa, noma ukudlulisa, ngokusebenzisa i-dorsal anterior cingulate, isifunda esingaphandle esisuka kufree rectus, noma engxenyeni engemuva yendawo 47 / 12. Ngalesi sigaba somkhakha, izitatimende ezilula zokuthi i-PFC ibandakanyeka emithethweni yemizwelo inikezela ngemininingwane enganele ukuze ibe wusizo, futhi ezimeni eziningi kungenzeka idukise, njengoba iningi lezifunda ze-PFC lingenakho ukubikezela okuqinile kuma-amygdala. Ukuvela kwamamodeli endlela agxila kuma-node aqakathekile afinyelela kuma-amygdala, njengamamodeli ahlongozwayo futhi ahlolwe nguWager et al. noPhillips et al. intuthuko ekhuthazayo kulokhu. Sisola ukuthi ukuze kube nenqubekelaphambili ethe xaxa yokuqonda ukubandakanyeka kwe-PFC emithethweni yokulawula imizwa, izindima ezilinganayo ze-dorsal anterior cingulate, i-postterior 47 / 12 kanye nesifunda esingaphansi kokulawula i-amygdala kuzodingeka kunqunywe.

Umbuzo osemqoka ubuye futhi ukuthi maqondana nokwanda kokusebenza kwe-PFC okuvela kakhulu ngesikhathi sokulawulwa kwemizwelo kuhlobana kanjani nalezi zindawo ezisemqoka, ngoba yizifundo ezimbalwa kuphela ezihlole ngokuqondile ukusebenza kwe-intra-PFC. Ngokungazi, lezi zindawo ze-PFC azixhunyiwe ngokulinganayo ku-dorsal anterior cingulate, i-posterior 47 / 12 noma isifunda esingaphansi komhlaba, futhi ngenxa yalokho kungenzeka zihlotshaniswe ngokukhethiwe nezindlela ezihlukile eziya e-amygdala. Sisola ukuthi ukukuqonda ngokuphelele ukubandakanyeka kwe-PFC emithethweni yokulawula imizwa kuzodinga ukucaciswa ukuthi zingaki kulezi zindawo ze-PFC ezingenalo izisulu eziqondile zokuhlangana ngokuhlangana nezinye izifunda ze-PFC ezinokuqagela okwanele kokulungisa imilenze.

Imininingwane ekuqondisweni kwamathonya

Siphikise ukuthi ukusebenzisana okuphezulu kwe-intra PFC, kanye nokusebenzisana kwe-PFC-amygdala okucacisa ukulawulwa kokuqonda kwengqondo okuqinile okungagxiliwe kwezinqubo zemizwa akuhambelani nezimpawu zokuxhuma okuphakathi kwezifunda. Ukuphikisana kwethu nalezi zinhlobo zendabuko eziphezulu ze-PFC-amygdala nokusebenzisana kwe-PFC kuncike kakhulu kwimodeli yesakhiwo echazwe nguBarbas nozakwabo, lapho iphethini le-laminar yemiklamo ibeka ukuthi ngabe ukuqagela kufanekisela ukukhetha okufana nokuphendula, noma Ukudluliswa okuphindwe kabili kolwazi. Uma kunembile, izindawo ezihlobene nomzwelo zibonakala zinikeza okuphezulu kakhulu kokulawulwa kokuphendula okungaphezulu kokwethula okuphakelayo kolwazi ukwedlula izindawo zokuqonda ze-PFC ngokwesiko.

Sikholwa ukuthi isigatshana somthethonqubo ongaphezulu kwesisekelo siholele ekucabangeni okuqondakalayo ekuqondeni ubudlelwano phakathi kwezifunda zobuchopho kanye nezinqubo zengqondo. Lokhu kukhetha kufanelana nombono wefilosofi wezindima zezinqubo “zokuqonda” kanye “nezemizwelo” ezibeka ukuqonda ngaphezu kwemizwa yobuzwe. Kepha lokhu kuthambekela kungaphazamisa amandla ethu okuthola ukuqonda okugcwele ngendlela ubuchopho obucwaninga ngayo ulwazi. Uma izinqubo zemizwelo zilawula ukusebenza kwe-“cognitive” futhi kuthande ezinye izindlela ukuzungeza, kakhulu noma ngaphezulu kunenye indlela ejikelezayo, amagama asezingeni eliphakeme naphansi angahle angalungisi lapho kucatshangelwa ukusebenzelana kokuqonda.

Ukulinganiselwa ekufiphiseni umsebenzi kusuka kusakhiwo

Ubuhle bemodeli yokuhleleka ukuthi kuholela ekubikezelweni okunamandla mayelana nohlobo lokuxhumana kwesifunda. Kodwa-ke, ukugxekwa okuningi kungaphakanyiswa ngokushesha ekudwebeni iziphetho ezisebenzayo ngokusekelwe kuzimpawu ze-anatomical. Okokuqala, yize imodeli yokwakheka isekelwa ngokuqinile ngokulandela ukubikezela kwayo kwamaphethini wokuxhuma we-laminar asuselwa ekwakhiweni kwe-cytoarureure, ukuthathwa kwemininingwane ephathelene nemiphumela yokusebenza kwalezi zindlela zokuxhuma ze-laminar akutholanga ukuhlolwa okuhlelekile kumasekethe ngaphandle kwemisinga yokucubungula imizwa. Yize kubukeka kunengqondo ukucabanga ukuthi izici ezifanayo ezisebenzayo zibonisa amakhethini we-laminar wokuqagela kulo lonke ubuchopho, lokhu akunjalo empeleni. Njengoba kunje, ukucekelwa phansi okuphathelene nezakhiwo ezisebenzayo zokuxhuma ku-PFC kusebenza kuphela uma izici ezisebenzayo zokuphakanyiswa kwesakhiwo kanye nokuqagelwa kwempendulo sekufakazelwe ukubamba kuzo zonke izinhlangano zososeshini.

Sithumele ukuxhumaniseka okuqinile phakathi kokuphendula okusebenzayo kanye nomthethonqubo we-top-phansi, nokuxhumana okufanayo okuqinile phakathi kwenqubo yokuphakelayo nokuya phansi. Izimpendulo zemibandela ne-feedSO zivela kumqondo wokulawula, ozama ukuchaza ukusebenza kwezinhlelo ezinamandla. Ukwemukelwa kwalawa magama ngabasebenza ngama-neuroscientist kanye nezazi zokusebenza kwengqondo akumangazi njengoba umqondo wezindlela zokunikeza ngezindlela zokulawula kanye nezinqubo ezilandelanayo ezinikeza ukudluliselwa kwemininingwane ezindaweni eziphakeme ekusakazweni kokuphawuleka kuyinto enembile. Kodwa-ke, i-equation elula yokulawulwa okungaphezulu kwephasi ngempendulo nangaphansi-up nge-feed Express kuyinkinga kuze kufike lapho izinkomba ezingeziwe zikhonjiswa nge-conceptualizations ephezulu nezansi. Izici ezinjalo ezingeziwe azivezwa ngokucacile, kepha zingafakazela ubucayi ekuqineni izindlela zokucubungula imininingwane. Sisola ukuthi abanye abafundisi bezenkolo basebenzisa amagama aphezulu-phansi naphansi ezansi ngezindlela ezingahambisani nempendulo kanye nezinqubo ezichazayo njengokuchazwa komqondo wokulawula, kepha lokho kungahambisani akuvamisile ukuthi kuchazwe ezincwadini.

Ekubonakaliseni impendulo kanye nokuqagelwa kwe-PFC, siphawula ukuthi asisho ukuthi konke ukuqagela kohlobo olufanayo. Izindawo zinenhlanganisela yokuxhuma, yokuphakelayo, nokuxhumeka okwalandelwa, kepha ukulingana kwalokhu kuxhumana kuhluke kakhulu ezindaweni zonke. Ngakho-ke sibonisa amaphethini amakhulu wokuxhuma, kepha lokhu akusho ukuthi ukuxhumana okusele akubalulekile. Isibonelo, izifunda ze-PFC ezi-eulaminate ngokuqinisekile zinemiklamo eyanele yokuphendula yokusiza ukuphatha izingxenye zezifunda ezingenamkhawulo, noma ngabe lokhu akuyona indlela ebanzi yokuxhumana phakathi kwezindawo.

Ngaphezu kwalokho, ukuxhumeka kokuqagela kohlobo lokuphakelayo kungenzeka kwezinye izimo kuguqule ukucubungula ezindaweni eziqondiwe kunokuba kumane kuthathwe imininingwane. Mhlawumbe isibonelo esihle salolu hlobo lokuguqulwa kwe-feedirect sivela kumamodeli wokuncintisana ahlanganisiwe (I-Desimone ne-Duncan 1995; UDuncan et al. I-1997) lapho ukuzuza kwesethulo esisodwa kuphumela ekucindezelweni komunye. Kumamodeli anjalo, ukudluliselwa kwangaphambi kwesethulo esinikeziwe kungaholela ekwandisweni kokusebenza kwalelo khuthazo, kanye nokucindezelwa komunye nomunye umfutho (I-Desimone ne-Duncan 1995). Ngale ndlela, okudluliselwa phambili kungathatha isinyathelo sokushintsha ukucubungula ezindaweni eziqondiwe. Ngokwesimo sokusebenza kwe-PFC, isiginali ye-DLPFC ingaguqula umncintiswano phakathi kwezethulo ezingaba khona ku-OFC ngalolu hlobo lokuqagela okuphindaphindwe. Lolu hlobo lwendlela yokuncintisana iyamangaza ngoba izosho izici ezithile zekhompiyutha (UWalther noKoch, 2006), okungakaze kufakwe kumamodeli wokulawulwa okungokomzwelo.

Uma ucubungula imodeli yokuhleleka, kubalulekile ukuphinda ugcizelele ukuthi imigomo esetshenziswa nguBarbas nozakwabo ukuchaza okuphakanyisiwe nokuxhumeka kwempendulo akuhambelani ngokuphelele nezindlela ezisetshenziswe abanye abaphenyi abahlola ukulungiswa kwe-laminarical proion. Ngokukhethekile, izincazelo zempendulo nokuxhuma ngaphambili zihlala zichazwa ngokubhekiswa kusendlalelo IV, njengokuthi ukuqagela (okwenyukayo) kokuqagelwa kuchazwa ngokunqanyulwa kwabo kungqimba IV (noma ubukhulu becala kungqimba IV), ngenkathi impendulo (yehlela) inqamuka ngaphandle kongqimba IV. Ngenkathi ukunamathela ngokuqinile kusendlalelo se-IV kungafuneki kululekwe kabi, njengoba kungafani nalawa maphethini (1)I-Pandya ne-Rockland, i-1979; UFelleman noVan Essen, 1991), umthelela wezinqubo zokunwebeka wokuvumela ukuqagela kuqeda ezingxenyeni ezingamapulani V no-VI kuphathwe njengokuqagelwa okuphambili akuqondakali ngokuphelele. Ngokucatshangwayo, izifundo zesikhathi sokudubula ezingxenyeni ezihlukile ze-PFC zingawuphendula lo mbuzo, kepha imininingwane kulolu daba iyantuleka.

Umbuzo wezimpawu ubangela ukumiswa isikhashana ngaphambi kokucabanga ukuthi i-OFC-DLPFC ngempela inephethini lapho i-OFC kufanele iphathwe njengezinga eliphakeme kune-DLPFC, futhi akuyona inhloso yethu ukuphikisana nalokhu. Noma kunjalo, kungashiwo ngokusobala ukuthi amaphethini wokuqagela ngokuqinisekile awahambisani nenhlangano yabaphathi lapho i-DLPFC isesimweni sobukhosi ngaphezulu kwe-OFC, ngendlela efana nezindawo zemizwa ezisezingeni eliphakeme ezihlala ngaphezu kwezindawo zemisindo ephakeme. Kanjalo, amamodeli enhlangano ye-PFC angaba ukuhlakanipha ukugwema ukubekeka okuphezulu kwe-DLPFC njengokuhlala phezulu kwesikhundla esiphakeme sezifunda ze-PFC.

Ukufaniswa kokuxhuma kwe-feedirect kanye nempendulo

Ekuhloleni amamodeli akhona wokusebenzelana kokuqonda imizwelo kuyaphawuleka ukuthi izifundo ezimbalwa ezishicilelwe kuze kube manje zifake izivivinyo ezithile zokuthi ukuqagela kubonisa impendulo, ukuqagela okuphambili noma okwakamuva (ngaphandle kokuqashelwa Seminowicz et al. I-2004). Ucwaningo oluningi lwe-neuroimaging lweqiniso alunikezi imininingwane ethile ye-laminar engalungisa le nkinga. Kodwa-ke, intuthuko yakamuva kumasu wokubonisa ukuxhumana okusebenzayo ihlinzeka ngamathuluzi angasetshenziswa ukubonisa uhlobo nokuqondisa kokuxhumana phakathi kwezifunda. Isibonelo, umfuziselo oshukumisayo we-causal (DCM) usebenzisa ukulinganiswa kwezinga lomndeni ne-Bayesian imodeli yokulinganisa ingasetjenziswa ukuhlola ama-hypotheses maqondana nokuqondiswa nobunjani bokuhamba kolwazi kanye nokushintshana kokushintshana kwezifunda zobuchopho ezahlukene (UFriston et al. I-2003; Chen et al. I-2009; UDaunizeau et al. I-2009; UFriston noDolan 2010; UPenny et al. I-2010). I-DCM futhi ingahlola amamodeli wokuncintisana anjengokunikeza ikhanda ukuqhathanisa ukuthi i-DLPFC phansi ilawula i-amygdala ngqo noma ngesakhiwo esithile sommeli. Kuze kube manje, kucwaningwe kuphela izifundo ezimbalwa ze-DCM ezihlobene nokusebenza ngokomzwelo (Ethofer et al. I-2006; USmith et al. I-2006; URowe et al. I-2008; U-Almeida et al. I-2009), futhi olwazini lwethu azikho izifundo ezishicilelwe ngqo eziphathelene nokulawulwa kwezimo. Kodwa-ke, ukusetshenziswa kwalezo zinqubo kungenzeka kuthuthukise kakhulu ukuqonda kwethu ukusebenzisana kwezimo zengqondo eminyakeni ezayo.

Ukuhlolwa okuqondile kwethonya

Mhlawumbe indlela engcono kakhulu yokwakha ubudlelwano obusebenzayo phakathi kwezifunda zobuchopho ngokuhlolwa kwesifunda esisodwa ngesikhathi sokukhetha komzimba waphezulu noma phansi kwendawo ethile. Isibonelo, uma i-DLPFC isebenza ngempela ukuqeda ukucubungulwa kwe-OFC, umuntu angalindela izimpendulo ezixakile ku-OFC lapho i-DLPFC ingeniswa ungaxhunyiwe ku-inthanethi. Lokhu kungenzeka kubhekwe ngokuhlola imisebenzi ye-OFC nge-fMRI ezigulini ezinezilonda ze-DLPFC. Ngenye indlela, i-transcranial magnetic stimulation (TMS) ingasetjenziswa ngaphezulu kwe-DLPFC ukuguqula isikhashana ithonya le-DLPFC kwimisebenzi ye-OFC. Impela, UKnoch et al. (2006) usanda kubika ukuthi i-TMS engakwesokudla i-DLPFC ikhiqize izinguquko zomsebenzi we-OFC wamuva ngendlela encike kumvamisa. Ngokufanayo, kungaba nentshisekelo ukwazi ukuthi izilonda engxenyeni eyodwa ye-cortex yangaphambili zithinta kanjani ukucubungula kwezinye izingxenye zenethiwekhi. Isibonelo, uma i-OFC ibalulekile ukubala inani lomvuzo elihlanzekile, kwenzekani ezindaweni ezingaphezulu lapho isuswa i-OFC? Saddoris et al. (2005) basebenzise lolu hlobo lwendlela yokuhlola ukuthi ukudubula kwe-OFC kuguqula kanjani ukudubula kwama-amygdalar ezindongeni zamagundane, kepha ezinye izifundo ezisebenzisa le ndlela azivamile ukwenzeka. Izincwadi ezikhulayo zokuxhumeka okusebenzayo kungenzeka ngokufanayo ukuthi zikhulise ukuqonda kwethu ukuthi lezi zindawo ezibucayi zobuchopho zixhumana kanjani. Kodwa-ke, ukuqonda okugcwele kwalokhu kuhlangana kuzofinyelelwa kuphela ngokucophelela ezicini ezithile ze-neuroanatomical zalezi zisekethe.

Lo msebenzi ubusekelwa izibonelelo T32MH018931-21, T32MH018921-20, & 5R01MH074567-04 ezivela kwiNational Institute of Mental Health. Sibonga uTawny Richardson ngosizo lokulungiselela lo mbhalo wesandla.

Ukuzikhulula komshicileli: Leli fayili le-PDF yesandla esingenakubalwa esamukelwe ukuze sishicilelwe. Njengenkonzo kumakhasimende ethu sinikeza le nguqulo yokuqala yombhalo wesandla. Umbhalo wesandla uzothola ukukopisha, ukufaka izinhlobo, nokubukeza ubufakazi obulandelayo ngaphambi kokuba ushicilelwe efomini layo lokugcina. Uyacelwa ukuthi uqaphele ukuthi ngesikhathi sezinqubo zokukhiqiza kungenzeka ukuthi zitholakale ezingahle zithinte okuqukethwe, nazo zonke izinqamulajuqu ezisemthethweni ezisebenza kulo magazini.

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