Neurosci Biobehav Rev. I-2012 kaJanuwari; 36(1): 479-501. Ipapashwe kwi-intanethi ye-2011 Agasti 25. ikhonkco: I-10.1016 / j.neubiorev.2011.08.005
URebecca uRay1 kwaye UDavid H. Zald2
Inguqulelo yokugqibela yomhleli yeli nqaku iyafumaneka apha Neurosci Biobehav Rev
Bona amanye amanqaku ku-PMC Wisdom nqaku epapashwe.
Abstract
Uphando ngengqondo luthi lubonise ukuba iinkqubo zeemvakalelo zinxibelelana nezinye izinto zokuqonda. Izifundo zibonakalise amandla okuphembelela ngokweemvakalelo ukuphembelela uluhlu olubanzi lwemisebenzi yokuqonda, kunye nokubanakho kwabantu ukusebenzisa iindlela zokulawula ukuqonda ezikwinqanaba eliphakathi ukulawula ulawulo lweemvakalelo. Amacandelo e-cortex yangaphambili abonakala edlala indima ebalulekileyo kolu lonxibelelwano. Nangona kunjalo, indlela la kusetyenziswana ngayo iphumelelayo ihleli kuphela icacisiwe. Kuphononongo lwangoku sichaza unxibelelwano lwe-anatomical phakathi kwe-ventral kunye nendawo yangaphambili dorsal kunye nokunxibelelana kwabo nemimandla ye-limbic. Kuphela kukusetyenzwa kweendawo eziphambili zangaphambili ezinokuthi zibe nefuthe ngqo ekusetyenzisweni kwe-amygdalar, kwaye njengemodeli zolawulo lwangaphambili lweemvakalelo kunye neemodeli zokulawulwa kweemvakalelo kufuneka zinyanzeliswe kwiindlela ezibonakalayo zempembelelo. Sikwajongisisa kwakhona indlela isakhelo esithile sokudibana kunye nokunxibelelana kwengxelo phakathi kwale mimandla okunokuthi kulandele imeko yokuhamba kolwazi phakathi kweendawo ezingena ngaphakathi kunye nezase dorsal. Ezi patheni zonxibelelwano azingqinelani kunye neembono ezixhaphakileyo eziveziweyo malunga nobume zonxibelelwano phakathi kwemvakalelo kunye nokwazi.
1. intshayelelo
Uphando oluzenzekelayo luvavanya ukuba inxibelelana njani nemvakalelo kunye neminye imiba yokuqonda. Umsebenzi onjalo uye wacacisa iindlela ezinokuthi ulwazi olungqineke ngokweemvakalelo lungaqondisa okanye luthande ingqalelo (U-Ahman et al. 2001; Uninzi et al. 2005; I-Mathews kunye ne-Wells, ngo-1999), kunye neenkqubo zokwenza izigqibo (UKnututson et al. 2008). Kwangelo xesha, uncwadi olukhulayo lubonisa ukuba iinkqubo zokuqonda ezinjengokuphinda uvavanye kunokulawula ukuphendula ngokweemvakalelo (UJackson et al. 2000; UKim noHamann, 2007; I-Ochsner et al. 2002; I-Ochsner, uRay, et al. 2004; URay, uWilhelm kunye noGross, 2008). Ewe kona, unxibelelwano phakathi kwemisebenzi echazwe ngokwesini ngokungqongqo ngokweemvakalelo okanye ngokungqongqo ngokwengqiqo kakhulu ngokwaneleyo ukubangela ukuba kuthandabuzwe ukwahlulwa okwenziwe rhoqo phakathi kwezi ndawo (bona umzekelo I-Pessoa, 2008). Nangona kunjalo, ukwahlula kubonelela ngexabiso elibonakalayo ekubeni ukuqhutywa kweemvakalelo kuneempawu ezithile zomsebenzi ezinokuthi zahlulwe kwezinye iziza zengqondo ngendlela efanayo iinkqubo zokunikela ingqalelo okanye iinkumbulo zineempawu ezahlukileyo kwaye zixhonywa ngokwahlukeneyo (nangona kunjalo ngamanye amaxesha kuyadibana) uthungelwano kwimimandla yobuchopho.
Indlela apho imvakalelo kunye neminye imimandla yokuqonda inxibelelana iye yaba yinto ephambili kwiimodeli zengqondo. Umzekelo, ukucengulwa kweziphazamiso zoxinzelelo zihlala zigxila kukukhathalelwa kwengqondo okunyanzelisiweyo ukuya kwisoyikiso (Bishop, 2007; ICisler kunye neKoster, ngo-2010; I-Ouimet, iGawronski kunye neDozois, ngo-2009; Williams et al. 1996). Ngokufanayo, ukusilela ukusebenzisa ulawulo olukumgangatho ophantsi kwiimvakalelo zijongwa njengokubaluleka kokuphazamiseka kwengqondo okubangelwa kukudakumba okukhulu (Intsomi et al. 2008; UJohnstone et al. 2007; UAlmeida et al. 2009; UTaylor Tavares et al. 2008), ukuya kumda wobuntu obuphazamisekileyo (Entsha et al. 2008).
Imimandla ephambili ibala ngokubonakalayo kwimodeli ye-neurobiological yonxibelelwano phakathi kwemvakalelo kunye nezinye izinto zokwazi. Nangona kunjalo, iimpawu anatomical zemimandla eyahlukeneyo yangaphambi kokuhlala zihlala zinikwa kuphela ulwazi lokujonga ngokujonga ubunyani beemodeli ezinjalo. Ukuya kufikelela kwinqanaba lokuthathelwa ingqalelo, kuhlala kuxoxwa kuphela ngokubanzi malunga nokuba indawo inonxibelelwano ngqo okanye udibaniso olusebenzayo kunye nemimandla ene-limbic, njenge-amygdala okanye i-hypothalamus. Nangona kunjalo, iinkcukacha zolu nxibelelwano zibalulekile ekuqondeni unxibelelwano lwengingqi. Umzekelo, imodeli ecela ukuba i-dorsolateral pre kwangaphambilial cortex (DLPFC) ngqo umsebenzi we-amygdalar unokuba nesandi kuphela ukuba ubonakalisiwe ukuba i-DLPFC ithumela uqikelelo oluthe ngqo kwi-amygdala. Ukuba uqikelelo olunjalo luthobekile okanye alukho, ezinye iimodeli ezizezinye zixhomekeke kwimimandla esembindini ziyakufuneka ukuba zichaze iimpembelelo ze-DLPFC ezineempendulo kwiimpendulo ze-amygdalar.
Amanqanaba obume bemimandla eyahlukeneyo yangaphambili kunye nephethini ye-laminar yoqhagamshelo lwayo inokunikezela ngokubonakalayo okukhulu kwentsebenzo phakathi kwemvakalelo kunye neenkqubo zengqondo eziphakathi zecortex (PFC). Ngokukodwa, imisebenzi ye-cytoarological yemimandla eyahlukeneyo ye-cortical ichaza indlela abaqhuba ngayo ulwazi kunye nokunxibelelana neminye imimandla. Eli nqanaba lesibini lokuhlalutya ngokubanzi alingenanga kwiingxoxo ze-neural substrates zentsebenzo-yokuqonda imvakalelo, nangona ineempembelelo ezinkulu zokuqonda ezi nkqubo.
Kwiphepha langoku, sizama ukucacisa amanqaku aliqela onxibelelwano olwahlukeneyo phakathi kweendawo ezahlukeneyo ze-PFC, kunye nokunxibelelana kwabo ne-amygdala. Sijolise ikakhulu kuthengiso phakathi kwe-orbital kunye ne-dorsolateral PFC ngenxa yokudibana kwexesha elide kwe-orbitofrontal cortex (OFC) kwiinkqubo zeemvakalelo (UZald kunye noKim, ngo-1996kunye nolwalamano lwexesha elide lwe-DLPFC kwimicimbi yolawuloI-Fuster, 1989; ISuss kunye neBenson, ngo-1986). Sikwachaza nendima yendawo yokuhlala yangaphandle (ACC) / izakhiwo zangaphambili zentsebenzo kolu nxibelelaniso, njengoko ulwazi olongezayo libonisa ukuba ezi zinto zibonelela ngonxibelelwano olubalulekileyo phakathi kwemvakalelo kunye neminye imiba yokuqonda.
2. Itoyografu kunye neempawu ze-cytoaruralural PFF
Iteyitogragi
I-PFC yahlulwahlulwahlulwa yangommandla obanzi we-6, dorsolateral, ventrolateral (VLPFC), frontopolar (FP), OFC, ventromedial (VMPFC), kunye dorsomedial (DMPFC) (jonga Umzobo 1). Imida yendawo ngqo yale mimandla isetyenziswa ngendlela eyahlukileyo ngabaphandi, kodwa uluhlu olubanzi luye lwangqineka luluncedo njengesikhokelo sokucwangciswa ngokubanzi sokuqonda ubuchule kunye nomsebenzi we-PFC.
IPhylogeny kunye neCytoarchitecture
I-PFC iqulethe iindlela ezimbini ezinokwahlulwa-hlukeneyo, ezineempawu eziqinisekileyo zephylogenetic (IBarbas, 1988; IiSanides, 1969; I-Yeterian kunye nePandya, ngo-1991). Imeko ye-basoventral isuka kwi-ophodoory (yolwabiwo) isisiseko se-OFC kwaye isasazeka ngaphandle kwendawo yangaphambili yecala, kwaye kamva kwi-VLPFC (ephela kwingingqi yeBrodmann (BA) V46). Ngokwahlukileyo, imeko ye-mediodorsal iqala ecaleni kwe-corpus collosum, iqhubekeka ngodonga lwe-lobe yangaphambili kwaye isongele umda ophezulu we-lobe ungene kwi-DLPFC (iphelela kwi-BA D46). Uncedo ngalunye kwezi lubonisa ipatheni yamanqanaba alandelelanayo yoyilo lwe-cortical oluboniswe kuphuhliso kunye nobubanzi boluhlu lwe-granular IV. Elona candelo lindala lendaleko lelokwendalo lihlala kwindalo, ngelixa iindawo ezincinci ezinokuvela zinomda omdaka kunye ochazwe kakuhle. Kwindlela ye-basoventral, oku kuqhubela phambili komzimba kuqalisa emva kwe-OFC (indlela ye-agranular isebenzisa isigama sikaCarmichael kunye nexabiso (UCarmichael kunye nexabiso, ngo-1994)) elandelwa yi-dysgranular (buthathaka granular) cortex kwimimandla esembindini ye-OFC, ukuya eulamini izendlalelo njengoko zihamba ziye kwicala elingaphambi kunye nakwindawo ezingena zithuthi (IBarbas kunye nePandya, ngo-1989; UCarmichael kunye nexabiso, ngo-1994; IPetrides kunye neMackey, ngo-2006; Ixabiso, 2006a). Imeko ye-Mediodorsal ibonakalisa ukuqhubela phambili kokuhamba komzimba okufanayo. Ukuqala nge-periallocortex cortex ecaleni kwe-rostral corpus collosum, imeko iba dysgranular kwi-cingate (kubandakanywa imimandla engaphandle, yangaphambili, kunye nemimandla engaphezulu), eulaminate I njengokuba umntu ehamba ngaphandle ecaleni kodonga lwendlu okanye ngaphezulu kweyona ndawo iphambili ye-gyrus. II kwimimandla ye-dorsolateral (BA 8 kunye ne-46).
Ukwenzela ukunqanda ukudideka, siphawula ukuba ukusetyenziswa kwendlela yeemododoral akufuneki kudityaniswe nomda we-DMPFC ochazwe kwi Umzobo 1. Inkqubo ye-Mediodorsal ibandakanya i-DMPFC, kodwa ikwabandakanya neendawo ze-VMPFC i-25 kunye ne-32, kunye neenxalenye ze-BA 10 eludongeni lwe-medial (indawo ye-10m kwindawo yokuma kwe U-Ongur et al. (2003); Umzobo 2).
Ipateni yophuhliso lwe-cytoaracological njengoko enye ihamba kwi-agranular ukuya kwi-eulaminate II cortex ihamba kunye nokunyuka kwenani elipheleleyo le-neurons (ubungakanani beeseli), ubungakanani beeseli zepramramidal kwizaleya II no-V, kunye nenqanaba le-myelination (IBarbas kunye nePandya, ngo-1989; Dombrowski et al. 2001; Umzobo 3), ezikhokelela kunye kwiimpawu ezahlukeneyo zokulungisa ulwazi kwimimandla eyahlukeneyo. Olunye umohluko omkhulu phakathi kwemimandla yangaphambili ivela ngokwembonakalo yokubekwa kwembali, ihlala ibonisa iimpawu ezahlukileyo zobumbano. UCarmichael kunye nexabiso (UCarmichael kunye nexabiso, ngo-1994) Yahlula i-Macaque OFC kunye ne-medial PFC kwiicandelwana ezininzi ngokuxhomekeke kwizinto ezinjalo (jonga Umfanekiso we4), kwaye uninzi lwezi mpawu zinokuchongwa ebantwini (I-Ongur et al., 2003). Iimpawu ezihlangeneyo zangaphakathi ezibonakala ngakwimimandlana engaphambi kwempembelelo zinefuthe leempawu ezithile zokulungiswa kolwazi olufezekiswe kukuhanjiswa kwangaphambili (Wang et al., 2004; I-Zald, 2007), kodwa ngaphaya kobungakanani beli phepha. Ngokucacileyo, izahlulo ezichaziweyo ngokwendlela efanelekileyo ye-PFC ineepatheni ezahlukeneyo zonxibelelaniso ngaphakathi kwi-PFC kunye neminye imimandla yobuchopho bokungafani.
Ubunjineli beCytoar kuMntu
Nangona kukho i-homology ebalulekileyo kulungelelwaniso lwangaphambili kunye nokwenziwa komntu ngaphakathi kwe-lobes yangaphambili, kunye neendlela eziqhelekileyo ze-phylogenetic zabiwe kuzo zonke iintlobo zezilwanyana, ubunzima obahlukeneyo buvela ekuhambeni phakathi kwedatha yabantu neyesilwanyana. Okokuqala, izifundo ze-neuroimaging zabantu zihlala zibhekisa kwiindawo zaseBrodmann (Brodmann, 1914), kodwa angabonisi uphuculo ekubonakalisweni kweendawo ezakhiwe ngokwaselwandle kunye nemida yendawo eyenzekileyo ukusukela oko umsebenzi ka-Brodmann enguvulindlela malunga nenkulungwane eyadlulayo. Okwesibini, ukusetyenziswa kwezilebhile zendawo kuhlala kusekwe kwiTalairach atlas (ITalairach kunye neTournoux, ngo-1988), kodwa le atlas yeyona ndlela isondele, kuba uhlalutyo lobunjineli bojikelezo lobugcisa ayenzelwanga kwingqondo eyila isiseko se-atlas. Okwesithathu, kukho ukungangqinelani phakathi kweelebheli zezilwanyana kunye neelebheli zabantu kwisebe yangaphambili yangaphambili, kuba idatha yezilwanyana zisebenzisa ezahlukileyo kwilebheli eyenziwe ngu-Walker (Walker, 1940), ezongezwe ngababhali ngoku ebantwini (IPetrides kunye neMackey, ngo-2006; I-Ongur et al., 2003), ngelixa uninzi lwabaphandi abanomdla besebenzisa iilebro zeBrodmann. Ngelishwa, ngamanye amaxesha akucaci ukuba yeyiphi iileyibheli zenkqubo yophando ezinxibelelanisa xa zisenza ingxelo. Oku kuvelisa ukuguquguquka okukhethekileyo kwi-lateral ye-OFC / VLPFC, apho abaphandi abangabantu bahlala bebhekisa kwi-BA 47, kodwa uncwadi lwezilwanyana lubhekisa kwingingqi ye-12. Ileyibhile ye-47 / 12 ngoku yamkelwe ngabathile be-neuroanatom ukuchaza le ndawo ebantwini, nangona umda we-medial wale ndawo uhlala uphikiswa ngabakhokeli be-neuroanatomists (IPetrides kunye neMackey, ngo-2006; I-Ongur et al., 2003). Ngokufanayo, iindawo i-13 kunye ne-14 zichongwe ngokucacileyo kwiinkawu, kwaye iindawo zasekhaya zijongwa ebantwini, kodwa akubanjwa i-Brodmann okanye Talairach, owafaka ilebhile yendawo eyi-11 kuwo omabini la macandelo angasemva nakwingaphandle kwele-OFC. Ukuchaza idatha ye-neuroimaging yabantu, ngokubanzi sibhekisa kwinkqubo ebanzi yelebheli echazwe I-Petrides neMackey (2006), endaweni yokufumana i-Talairach atlas ukuze uthathe ithuba ledatha kwizifundo zangaphambi kobuntu.
3. Unxibelelwano
Uninzi lwedatha esele ikhona kuqhagamshelo olungaphambili lufumaneka kwizifundo zezilwanyana. Nangona kunjalo, xa sinikwa isifundo se-cytoaruralural Homology (IPetrides kunye neMackey, ngo-2006; I-Ongur et al., 2003), ngokubanzi kucingelwa ukuba ukuqhagamshelwa kwezi ndawo zigcinwe ubukhulu becala kuhlobo lweentlobo. Kananjalo, kusengqiqweni ukusebenzisa uncwadi olungelulo olwangumntu ekubumbeni njengesiseko sokuvavanya uxhulumaniso ebantwini. Sijolise kwiindidi ezimbini zonxibelelwano apha: unxibelelwano lwe-amygdala-PFC, kunye nonxibelelwano phakathi kwemimandla eyahlukeneyo ye-PFC.
Igalelo lama-Amygdalar kwi-PFC
I-OFC kunye ne-PFC yelanga ifumana igalelo elikhulu kwi-amygdala (Amaral et al., 1992; UCarmichael kunye nexabiso, ngo-1995; IBarbas kunye neZikopoulos, ngo-2006). Oku kuma ngokwahluke mpela kwi-DLPFC, efumana uqikelelo oluncinci ngokuthe ngqo kwi-amygdala. Uphengululo loncwadi lubonisa ukuba ezinye zezinto eziza kuqikelelwa kwi-ventral nakwi-medic ye-PFC ziyahluka ngokuxhomekeka kwinowuthi okanye kwi-subnikhaliAmaral kunye nexabiso, ngo-1984; IBarbas & De, 1990; Amaral et al., 1992; UCarmichael kunye nexabiso, ngo-1995). Nangona kunjalo, ezi nkcukacha zingaphaya kobubanzi beli phepha, kwaye umfanekiso oqinileyo wokuxhuma uphuma kuyo yonke i-nuclei eyahlukeneyo ukwazisa ngengxoxo ngokubanzi yeepatheni zokunxibelelana. Umzobo 5 bonisa ubume ngokubanzi boqikelelo lwe-amygdalar (evela kwi-amygdalar nuclei) kwingqondo ye-medial kunye ne-ventral kwengqondo ye-macaque usebenzisa i-Carmichael kunye ne-Intenclature yexabiso. Eli nani libonakalisa ukuba umphezulu we-orbital awufani ngokudibeneyo kwe-amygdala. Eyona nto iqaphelekileyo kukungabikho kokufakelwa okukhulu kweendawo i-13m, 13l, 12m, 11l, kunye ne-10o kumhlaba orbital. Udonga lwe-medial lukwafumana igalelo le-amygdalar enkulu, kodwa kwakhona ayifani, njengoko kungekho ndawo 10m okanye ndawo 9 ifumana igalelo elibalulekileyo le-amygdalar.
Izigqibo ezibini zinokuthathelwa kule ndlela yokufaka. Okokuqala, igalelo le-amygalar kwi-PFC licwangcisiwe kuyilo kwaye lijolise kwimimandla encinci eyakhiwe ngokwendawo. Oku kubonisa ukuba kuya kuba yimpazamo ukuphatha ngokwendalo yonke i-OFC okanye i-PFC yodidi ngokungathi inxulumene kakhulu ne-amygdala. Endaweni yoko, ukujonga indawo ngaphakathi kwe-OFC kunye ne-medial PFC kucetyiswa ngaphambi kokuzoba ukuthotyelwa malunga nokuxhunyaniswa kwe-amygdalar. Okwesibini, i-DLPFC kunye ne-FP bafumana igalelo elibuthathaka ngokuthe ngqo kwigalelo le-amygdalar (inene kuphela iindlela ezibuthathaka kakhulu zibonisa ubungqina bokufakwa kwe-amygdalar). Ngenxa yoko, iimpembelelo ze-amygdalar kwi-DLPFC kunye ne-FP kusetyenzwa zinokungangqinelani, nokuba zidluliselwa ngakwicala le-cingulate okanye lendawo yangasemva ye-OFC (okanye ngokusebenzisa ezinye iindlela ezibanzi, ezinje ngokuguqulwa kweenkqubo ze-neurotransmitter).
Iziphumo eziphambili zangaphambili kwi-amygdala
Iziphumo ze-PFC eziya kwi-amygdala zikwindawo ethile yommandla (Ixabiso, 2006b; UGhashghaei et al., 2007; UStefanacci kunye neAmaral, 2002; UStefanacci kunye neAmaral, 2000). Ngokubanzi, iindawo eziphambili zangaphambili ezifumana uqikelelo kwi-amygdala zithumela uqikelelo ku-amygdala, ngelixa iindawo ezingafumani galelo likhulu le-amygdalar (njenge-DLPFC kunye ne-FP) zinobungqina obuthathaka kakhulu kwi-amygdala. Ubungqingqwa boqikelelo lubonisa ubukhulu be-cytoarchitectonics, kunye nobuthathaka obuthathaka boxinaniso lweengqikelelo njengoko usuka kwiindawo ezigudileyo ukuya kuphuculo olungendawo lwe-eulaminate isocortex. Le patheni ibonisa ukuba iindawo zecocortical (DLPFC kunye ne-FP) azinakho ukubonelela ngefuthe eliqilima ngqo kwi-amygdala, kwaye ukuya kuthi ga apho banefuthe kwi-amygdala, impembelelo inokubakho ngokungathanga ngqo. Oku akuthethi ukuba akukho luqikelelo luthe ngqo olusuka kwi-DLPFC ukuya kwi-amygdala, njengoko izifundo ezininzi ziye zabona ulwaziso ngqo ukusuka kwindawo 9 kunye 46 (UStefanacci kunye neAmaral, 2002; I-Aggleton et al., 1980; Amaral kunye no-Insausti, 1992). Endaweni yoko, uqikelelo ngokubanzi ukukhanya kakhulu kunikezela ngomgaqo obanzi wokusebenza kwe-amygdalar.
Nangona i-cytoarchitectonics ngokubanzi ibonelela ngenqophiso eyomeleleyo yenqokelelo ngokubhekiselele kuqhagamshelo lwe-amygdala-pre-prealal, ukusasazwa kwengingqi kunye nokungenayo akuphumanga ngokulinganayo (UGhashghaei et al., 2007). Ngokukodwa, eyona galelo liphezulu le-amygdalar kwi-PFC likummandla we-agranular insular ecaleni kwendawo engasemva ye-OFC, ngelixa, indawo enkulu ekuphumayo kwi-amygdala ivela kwindawo yangasemva ye-cingate cortate (BA 25) kunye nezahlulo ze-dorsal anterior cingulate (BA 24 ). Ngokubanzi, iindawo zodonga lwe medial zinesiphumo esiphezulu kunegalelo le-amygdala, ngelixa iindawo ezingasemva kwe-OFC zinegalelo elingaphezulu kunokukhupha. Okubangel 'umdla kukuba, ngokuthe ngcembe ngakumbi okunxibelelana ngakumbi kwi-PFC, imimandla ye-DLPFC (i-BA 8, i-9 kunye ne-dorsal 46) inegalelo elikhulu kunokungeniswa kwi-amygdala, ngelixa le patheni iphinda ibuyiselwe ngaphakathi kwi-VLPFC. Inqaku malunga noku, lo mmandla ungasemva wengingqi ye12l ngaphakathi kweVLPFC ubonelela ngoqikelelo olusemgangathweni kwi-amygdala, eyenza ibe yeyoqobo le-PFC yengingqi enegalelo elibalulekileyo kwi-amygdala. Ewe, oluqikelelo luluqine kunoko kubonwa kwimimandla ye-anterior orbital, eyabelana ngeendawo ze-12l ubukhulu becala lokuphuma kunokokufaka, kodwa bonisa ngokubanzi amanqanaba obuqhetseba kunendawo ye-12l.
Inani le-subnuclei ezahlukeneyo ze-amygdala zifumana igalelo le-PFC. Isiseko kunye nesiseko sezinto ezingasisiseko kunye ne-medial nuclei zifumana iingqikelelo ezibalaseleyo, kunye nokwamkela uqikelelo kubalo oluphezulu lwemimandla ye-PFC, ngelixa i-neneal, embindini kunye ne-cortical kini ifumana uqikelelo lwe-PFC, kodwa kwinqanaba elincinci nelibanzi (UStefanacci kunye neAmaral, 2002). I-BA 25 iyaphawuleka kuba ayithumeli nje kuphela iingqikelelo ezixineneyo kwi-amygdala, ikwathumela noqikelelo kuluhlu olubanzi lwee-nuclei, njengoko zonke i-amygdala subnucleus ephawulwe ngentla ifumana igalelo kwi-BA 25. Nangona ivela njengokukhanya kwikholamu B ye Umzobo 6, kubalulekile ukuba uqaphele ukuba i-BA 32 inikezela ngengqikelelo echazwe kakuhle kwi-amygdala. Ngeendlela ezininzi i-BA32 ibonakala i-homologous kwi-prelimbic cortex kwiintonga (Ixabiso, 2006a). Kwiintonga, iiprojekthi zecortex ye-prelimbic ukuya kumacandelo easolateral kunye nentloko ye-amygdala ephakathi (I-Vertes, 2004). Kwimimandla engengowomntu, uqikelelo lubekiwe ukusuka kwi-BA 32 ukuya kwinxalenye ye-discal basal nucleus (I-Chiba et al., 2001). Ke, nangona zincinci kwaye zinabile kunokuba kuqikelelwa kwi-BA 25, i-BA 32 ibonakala ikwindawo yokusebenzisana neenkqubo ze-amygdalar ezikhethiweyo.
IBarbas kunye neZikopoulos (2006) Ukuphikisana ukuba iziphumo zangaphambi medialal kunye ne-OFC kwimveliso ye-amygdala zinokuba neempembelelo ezahlukileyo ekusebenzeni kwe-amygdalar. I-BA25 kumphezulu wentambo inikezela ngokukhethekileyo iziphumo ezivuthayo zeenxalenye zesiseko se-amygdala, yona ekuboneleleni ngoqikelelo lwethamsanqa kwi-hypothalamus. Ngokuchaseneyo, i-post yangaphambili ye-agranular ye-OFC ibabeka ngaphakathi ngaphakathi ubukhulu bezinto ezihlanganisiweyo ezijikeleze i-basal nucleus (jonga Ikhiwane. 7). Imasisi ezibandakanyekileyo zinika igalelo le-inhibitory kwi-nucleus esembindini. Xa ivuselelwe, iingxangxasi ezihlanganisiweyo zimisa umendo wendlela we-tonic inhibitory ukusuka kwi-nucleus esembindini ukuya kwi-hypothalamus, ngaloo ndlela kubangela ukuphazamiseka kwe-hypothalamus. Ukuqikelela okuchulumancisayo okulula kufikelela kwi-nucleus esembindini ngqo ukusuka ngasemva kwe-OFC, ivumela i-OFC ukuba inyuse okanye inciphise i-nucleus yokudubula ephakathi.
Izinto eziphambili eziphambili eziqikelelweyo kwi-hypothalamus nakwindlela yokusebenza kwengqondo
Iindawo ze-OFC kunye ne-PFC ye-medial eneengqikelelo kwi-amygdala ikwayi projekti yendawo ye-hypothalamus kunye ne-autonomic brainstem / periaqueductal grey (I-et al., 1998; UBarbas et al., 2003; Ixabiso, 2006b; Rempel-Clower kunye neBarbas, 1998), ukubonelela ngokubanakho ngqo kwempembelelo kwimimandla yeefandisi zesiphumo ezihambelana nokuphuma kwemvakalelo (jonga Umzobo 8). Oluqikelelo lubonakala lomelele ngakumbi kulwakhiwo lodonga lwedial, kodwa luvela nakwindawo efana nendawo kumhlaba orbital apho igalelo le-amygdalar likhulu. Njengokungafikeleli kwayo ngokuthe ngqo kwi-amygdala, i-DLPFC kunye ne-FP ubukhulu becala azinalo ucwangciso ngqo kwezi ndawo. Ukongeza, izahlulo ezingaphandle ze-OFC zibonisa kancinane ukuphuma ngokuthe ngqo kula maziko azimeleyo.
Unxibelelwano ngaphakathi kwe-lobe yangaphambili
Njengoko kuphawuliwe apha ngasentla i-PFC inokwahlulwa ibe ziindlela ezimbini eziphambili ze-phylogenetic. Olona nxibelelwano luphakamileyo lommandla ngamnye lineendawo ezifanayo, ngakumbi iindawo ezingabamelwane ezingabikho kwinqanaba elinye lophuhliso kude nendawo ekuthethwa ngayo. Yiyo loo nto, umzekelo, imimandla ye-agranular insular yendawo engasemva ye-OFC inokunxibelelana okubonakalayo nakweminye imimandla ye-agranular kunye ne-dysgranular orbital, kodwa ngokubanzi ayinalo unxibelelwano kwimimandla ye-isocortical efana ne-46 yendawo yokuma kwendlela yayo. Apho kuvela khona uqhagamshelo lwendawo yokuhamba phakathi, abahlali bagxume ngaphezulu kwenqanaba elinye kophuhliso loyilo. Umzekelo, indawo ye-isocortical ventral 46 idibene ngamandla ne-isocortical dorsal ndawo 46 kwindlela ye-Mediodorsal, kodwa ayixhumeki kwiindawo eziphuhliswe kakubi zedonga medial ezifana ne-cgenate ye-subgenual (BA 46). Iindawo ezingakumbi zangasemva kunye nezingasemva ze-OFC zinonxibelelwano olugcweleyo lwendawo eyi-25 yendawo engummelwane ne-46, kodwa uqhagamshelo oluxhuma isalphic eyintloko kwindawo yendawo eyi-45 ayinqabile.
Nangona kunjalo, iindawo ezininzi ze-OFC zibonakala zinonxibelelwano ngqo ne-DLPFC. Iindawo ngokukodwa i11m, 12o, 13a kunye 14r nganye inento enxibelelanayo ne-DLPFC. I-gyrus rectus (equka indawo ye-14r) inokujongwa njengenxalenye yendlela yokuthetheleleka okanye njengendawo yenguqu phakathi kweendlela, ngenxa yoko uqhagamshelo lwayo alumelani nokutsiba kwendlela yokuhamba. Nangona kunjalo, iindawo i-11m, i-12o, kunye ne-13a zidityaniswe njengenxalenye ye-basoventral trend, ngenxa yoko amakhonkco abo kunye ne-DLPFC amele uxhulumaniso phakathi kwezinto. Ukuqonda isikhundla esikhulu senethiwekhi kwezi ndawo, kuluncedo ukuqwalasela enye indlela yokuhlela imimandla ye-orbital kunye neyelaphaya. Endaweni yokuseka iimodeli ezisekwe kwi-phylogeny, UCarmichael kunye nexabiso (1996) Yahlulahlula i-OFC kunye nodonga lodidi lube yindawo ye-orbital kunye nenethiwekhi yokubonisa ngokudibeneyo ngokuqinileyo kumandla onxibelelwano phakathi kwemimandla (bona Ikhiwane. 9). Olu hlobo lwenkqubo yokuhlelwa lubonisa ukugqagqana okukhulu kunye nokwahlula kwe-phyolgenetic phakathi kwe-basoventral kunye neendlela zokuziphatha eziphakathi, akumangalisi ukuba umbutho uqhagamshelo sele uxubushile. Nangona kunjalo, ezi nkqubo zimbini zokuhlelwa azifani. Into enomdla kukuba, zonke iindawo ze-orbital ezinxibelelana ne-DLPFC ziyinxalenye yomnatha we-medial kaCarmichael, okanye uthathwa njengomlamli phakathi kothungelwano. Umzekelo, indawo ye-11m ithathwa njengenxalenye yenethiwekhi yoku medial, kuba inonxibelelwano olukhulu nemimandla yedonga loku medial kune-OFC iphela. UCarmichael kunye neXabiso bahlukanisa iindawo i-12o kunye ne-13a njengemimandla yojongano kuba ziqukethe uqhagamshelo olunzima kuzo zombini iindawo zokomoya kunye ne-orbital. Ezi patheni zahlukileyo zoqhakamshelwano zenza kucace ukuba kuya kubakho ukuphawulwa kwengingqi, okanye nokuba kukwahlula-hlulwa, umahluko kubukho be-OFC ukunxibelelana nezinye iindawo eziphambili. Ngokukodwa, i-gyrus rectus, kunye ne-11m, i-12o, kunye ne-13a zikwindawo yokunxibelelana nazo zombini iindawo zodonga lwendondo (umz. I-cingate) kunye neendawo ze-DLPFC, lo gama ezinye iindawo ezizungelayo zingenalo ulwalamano ngqo.
Uqhagamshelo lwenethiwekhi ephambili ilawula iindlela kwi-amygdala
Kwiindawo ezingenakho ukuvela ngokuthe ngqo kwi-amygdala, amandla okuphembelela ukusebenza kwe-amygdalar kufuneka axhomekeke kwindlela ezingathanga ngqo, kwaye ezi ndlela ziya kuthi zichazwe ikakhulu zizikhundla zazo kuthungelwano oluphambili. Ngenxa yamandla e-infgenual cingulate's (BA 25) ukuqikelela kwi-amygdala, inokubonelela ngodluliselo olubaluleke kakhulu apho imimandla eyahlukeneyo ye-PFC inefuthe kwi-amygdala. Njengoko kunokubonwa Umzobo 9I-BA 25 ifumana uqikelelo oluninzi oluvela kwiindawo zenethiwekhi zedial kunye nakwiindawo zomzimba ezihambelana nothungelwano lwe-medial. Ngokwahlukileyo, ulwaziso ngakumbi dorsolateral lunqabile. UVogt noPandya (1987) Qaphela ukuba i-BA 25 ifumana uqikelelo kwi-DLPFC, kwaye ichaze ngokuthe gca i-9 yendawo kwindawo ephezulu ye-DLPFC. Nangona kunjalo, amandla oqhagamshelo abonakala engenamandla, kwaye engakhange abonwe ngokucacileyo kwezinye izifundo (IBarbas kunye nePandya, ngo-1989). Nangona kunjalo, i-BA 9 idibene kakuhle ne-BA 32 kudonga lwe-medial, yona ke idityaniswe kakhulu ne-BA 25 (UCarmichael kunye nexabiso, ngo-1996; IBarbas kunye nePandya, ngo-1989), kwaye ke ngoko kubonelela ngendlela enokwenzeka yokungaqhubekiyo yokwenza i-DLPFC ukuphembelela i-BA 25. Ngokufanayo, i-dorsal BA 46 ayinakukwazi ukunxibelelana kakhulu ne-BA 25, kwaye kuya kufuneka ukuba ibandakanye i-BA 32, okanye mhlawumbi ezinye izahlulo ze-cingex cortex, ukuze inxibelelane ne-BA 25.
I-dorsal ACC (BA 24) ikwabonelela ngommandla wokukhupha iziphumo kwi-amygdala. Lo mmandla unepateni efumileyo yamagalelo evela kwi-PFC (UCarmichael kunye nexabiso, ngo-1996; IVogt kunye nePandya, 1987; IBarbas kunye nePandya, ngo-1989). Oku kubandakanya igalelo elibalulekileyo elivela kwi-BA 9, kunye nokuya kwi-BA 46 kwi-DLPFC, izahlulo ze-BA 32, kunye ne-BA 10 eludongeni lwe-medial, kunye nemimandla eliqela ye-OFC (ngakumbi imimandla yendlela yangaphakathi / yangaphakathi ye-13a kunye ne-Iai, kunye ne-12o ). Yiyo loo nto i-dorsal ACC ibonakala ikwindawo eqinileyo yokuhlanganisa iinkalo zokusebenza kwe-PFC kwimimandla emininzi.
Nangona imimandla engaphandle ye-OFC kunye ne-VLPFC ingabonakala inomlinganiselo omkhulu wokukhutshwa kwe-amygdala kunokokufaka okuvela kwi-amygdala, kuba ezi zinto ziqikelelwa zithobekile, le mimandla ingaphandle ingasebenzisa iindlela ezingathanga ngqo ukubandakanyeka kwi-amygdala. Kwimimandla engaphandle ye-OFC, oku kungabhekiswa kakhulu kwimimandla yangasemva ye-OFC. Ngokwahlukileyo, kwimimandla yokuhambahamba ngenqanawa, indawo engasemva ye12l inokubonelela ngendlela ethe ngqo yokubandakanya i-amygdala, inikezwe isikhundla sayo esingafaniyo naphakathi kwenethiwekhi yangaphambili kunye ne-amygdala.
4. Imodeli yolwakhiwo
Iimpawu ezijikeleze ujikelezo lommandla we-cortical zone ziphembelela kakhulu indlela le ngingqi inxibelelana ngayo neminye imimandla yobuchopho. Ngokukodwa, inqanaba lobuhlakani kunye nophuhliso lwe-laminar linefuthe kwinqanaba lokuphononongwa kunye nengxelo yengqikelelo (IBarbas kunye neRempel-Clower, ngo-1997; IBarbas 2000). Ngaphakathi kwimodeli evezwe yiBarbas, iingqikelelo zokuphinda zichazwe ngendlela eyiyo njengokuvela kwizinto ezingaphezulu kunye nokubhengezwa kwezinto ezinzulu zecortex. Kwinkqubo yeemvakalelo, amabakala okuqala omjelo wokuqhubela phambili anikezela ngolwazi kumanqanaba alandelayo ngoluhlobo lokuqikelela okuphindiweyo (I-Rockland kunye nePandya, ngo-1979; IPandya, i-1995). Ngaphakathi kweenkqubo ezinendawo yokusebenza ecacileyo yokuhamba kolwazi, ezinje ngeesistim zeemvakalelo, iindlela zokuphinda zondlalwe nazo zinokuchazwa njengokuba zisenyuka ukusuka kwindawo yokuqala ukuya kumanqanaba aphezulu kumjelo wokuqhubekeka (umzekelo i-V1 ukuya kwi-V2). Ngokumalunga neenkqubo zokuqonda, ukudluliselwa kolwazi kuhambelana ngqo neeleyibhile zesiko lokwazisa ngokwenkcubeko njengenkqubo yokuqhubela ezantsi (IKastner kunye neUgerleider, 2000).
Ngokwahlukileyo, uqikelelo lwempendulo luqala kumaleko ezinzulu ze-cortex kunye neprojekthi ukuya kumaleko angaphezulu kakhulu e-cortex (bona Umzobo 10). Kwiinkqubo zeemvakalelo ezinesakhiwo esicacileyo sendlela yokusebenza, ezi zinto zinokuchazwa zinokuchazwa njengezisehla, njengoko zihamba ukusuka kwinqanaba elizayo ukuya kwinqanaba langaphambili lomjikelo wokusebenza wezinzwa (umzekelo, iV2 ukuya kwiV1). Uqikelelo lwempendulo isenzo sokuguqula okanye ukukhumbula ubalo olwenziwe kumanqanaba okuqhubela phambili (IRaizada kunye neGrossberg, 2003). Umzekelo, ezi zinto ziqikelelweyo zokwenza ingxelo ukunceda ukunceda ukonyusa iimpendulo zeeseli zokufaka iikhowudi kwizinto okanye kwiindawo, ngelixa unika okanye ucinezela iimpendulo kwizinto ezinganwanga (Mehta et al., 2000; Saalmann et al., 2007). Ezo ngxelo zoncedo kwiinkqubo ezisisiseko ezibonakalayo njengokucalulwa komhlaba (IDomijan kunye neSeti, ngo-2008; URoland et al., 2006), kunye nokuvumela ulawulo oluphezulu lwezinto ezenziwa kumjelo wolwazi (I-Grossberg, 2007). Kwimibono yokuqonda, olu lawulo luphezulu lusebenzisa uguquko lwento esekwe kulindelo, kwiinjongo zangoku nasekuqwalaselweni kwengqondo (UGlibert noSigman 2007).
Ukucacisa, kuluncedo ukwahlula phakathi kwesigama sokunyuka / ukuhla, ukuphawulwa / ingxelo, kunye nokuhla / phezulu-ezantsi, njengoko zithetha izinto ezahlukileyo (bona UPenny et al. 2004 yengxoxo). Ukunyuka nokunyuka kweqikelelo kubhekisa kumanqaku athile obuchwephesha, kwaye isigama siluncedo ikakhulu kwimeko yemijelo ecociweyo echaziweyo. Sisebenzisa eli gama lisezantsi kunye phezulu-ezantsi ukubhekisa ngqo kwiinkqubo zokuqonda, ukuya emazantsi ebhekisa kwiinkqubo ezizenzekelayo, ezinje ngeempendulo eziqhutywa kukuqonda kwesikhuthazo, kunye nokubhekisa ezantsi kwiindlela ezivumayo yemodyuli eguqulwayo yokuqhubela phambili ukuhlangana kunye neenjongo zangoku kunye nolindelo. Amagama okuphakanyiswa kunye nengxelo kulo mxholo aneentsingiselo ezimbini, njengoko zichazwa ziimpawu ezithile ze-laminar zoqikelelo, kodwa zikwathetha neempawu zokulungisa ulwazi.
Kungenxa yokuba ingxelo kunye nengqikelelo ye-feedirect ichazwa ngamanqaku e-laminar, inkxalabo inokuphakanyiswa malunga neendlela ezithile ze-laminar ezisetyenziswa yiBarbas kunye noogxa bokuphawula uqikelelo njenge-feedxp okanye impendulo ngaphandle kwemijelo yokusombulula imizwa. Kwimodeli yenkqubo yokubonakalayo, ukuqikelela okuphindaphindiweyo kuchazwa ngokunxibelelene nokupheliswa kwe-laminar IV, ngokunyuka okuqikelelweyo kokuphakanyiswa kweendonga ezingaphezulu kunye nokupheliswa komaleko IV (ngokuchasene nobubanzi benkqubo; UFelleman kunye noVan Essen, ngo-1991). Ngokwahlukileyo, iBarbas isebenzisa inkcazo ebanzi engafaniyo nokwahlula phakathi kongqameko lwe-IV kunye nolwakhiwo lwe-infragranular. Olu lwandiso lusemgangathweni ofanelekileyo ngenxa yobukho bendawo yangaphambi kokuba inesiseko esomeleleyo seplanular, kunye neendlela zokuphelisa amatye okuphelisa amatshe eziqwalaselwe kule mimandla. Nangona kunjalo, iimpembelelo ezisebenzayo zolwandiso zihlala zichaziwe.
Kuvela umba owahlukileyo kwinkcazo yoqikelelo lwempendulo. Inkcazo kaBarbas yengxelo ijolise ngokukodwa kuqikelelo olusuka kubunzulu (infragranular) layita kunye nokupheliswa kweengqimba eziphezulu, ngokungqinelana nomsebenzi wakuqala I-Rockland nePandya (1979). Nangona kunjalo, UFelleman noVan Essen (1991) Ukuphika ukuba uludwe lweempendulo ezongeziweyo zinokuba nemvelaphi ye-bilaminar ngokudibeneyo kwemvelaphi ye-infra- kunye ne-supra-granular. Ukusukela, uBarbas ugcina inkcazo egciniweyo ngakumbi, ukwamkelwa kwendlela yakhe yokugweba ngokuthe ngqo kuyacaca. Nangona kunjalo, kukhokelela kwinto yokuba umlinganiso woqikelelo ubonakaliswe njengempendulo kwi-PFC unokuba phezulu usebenzisa inkcazo yenkululeko.
Eyona nto iphambili kwimodeli eyakhiweyo kukuba inqanaba lokuphendula kunye nolindelo oluqinisekileyo phakathi kwemimandla luqingqwe ubukhulu becala ngumgangatho wokuphuhliswa kobunjineli bemimandla. Ukuqikelelwa ukusuka kwi-cortex eyahluliweyo ethe kratya (okt, eyahlulahlulweyo ngakumbi, kunye ne-denser granular layer) ukuya kumgangatho ophantsi wobunjineli bobuchwephesha bokulandela isakhelo sephepha, ngelixa ezo zisuka kuphuhliseko olungephi oluphuhliweyo ukuya kwicortex ephuhliswe ngokwendlela ilandela uhlobo lwempendulo. Le patheni ihambelana noko kubonakala kwiinkqubo zeemvakalelo, kodwa le pateni ibonakala inokufana kwiinkqubo ezininzi. Kwi-PFC, imodeli yolwakhiwo icingela ulungelelwaniso lwe-feedxpo kunye nengqikelelo yengxelo malunga ne80% yexesha, kunye nokulingana kwengxelo kunye nokunxibelelana okuphindaphindiweyo kuya kusiba mkhulu ngakumbi umahluko kuphuhliso lobugcisa be-cytoarological phakathi kwale mimandla mibini ikulo mbuzo (IBarbas kunye neRempel-Clower, ngo-1997).
Eyona nto isetyenziswayo yemodeli yolwakhiwo ukuya kwisihloko esikhoyo kukuba ikhokelela kwisimo sezulu esiqikelelweyo malunga nobume bonxibelelwano phakathi kwemimandla yobuchopho nangona kungekho datha esebenzayo. Ewe kunjalo, ekugqibeleni, i-electrophysiological okanye ezinye iindlela ezinobuchule zokuvavanya ukuhamba kolwazi lwe-laminar ziya kuba yimfuneko ukuqinisekisa ukuba iipatheni zentlelo ye-laminar kwi-PFC isebenza ngokufana noko kubonwa kwii-sensor cortices (okt, ingxelo echaziweyo yokwenziwa kwengxelo kunye neprojekti yokutya iipropathi ezifanayo zolwazi zihamba ngaphandle kwenkqubo exeliweyo). Izifundo ezinje nge-electrophysiological zinokuthi ekugqibeleni zincedise ukuphucula imilinganiselo yokuchaza ngendlela eyi-feedfoward kunye nengqikelelo yengxelo. Okwangoku, imodeli yolwakhiwo ibonelela ngesiseko esomeleleyo esikhoyo ngoku sokuxela uhlobo lokuhamba kolwazi kwi-PFC. Ukuba imodeli yolwakhiwo ichanekile ekubonakalisweni kokungena kolwazi ngaphakathi kwi-PFC, ineempembelelo ezibonakalayo kwiimodeli zokunxibelelana nemvakalelo.
Iipateni zeLamari kunye nokunxibelelana kwangaphakathi
Ngokuhambelana nemodeli yolwakhiwo, uhlalutyo lweepatheni zamalin yeendlela eziqikelelweyo zibonisa ukuba idreyini ye-OFC ibonakaliswa luphawu lweengxelo ezinamandla kunxibelelwano lwayo kunye nemimandla engaphezulu yophuhliso lwe-PFC (IBarbas, 2000). Ngothelekiso kwiinkqubo zeemvakalelo, oku kuya kuthetha ukuba uqikelelo lwe-OFC lwenzelwe ukukhetha okanye ukulungisa ukubala. Ngokwahlukileyo, i-eulaminate DLPFC inamanqanaba aphezulu okuqikelelwa okuphindaphindiweyo, okukuvumela ukondla iziphumo okanye iziphumo zokuqokelelwa kwayo kwimimandla yengqondo elandelayo. Le patheni iyonke yokuqikelela okuphindaphindiweyo kunye nengxelo yengxelo ikwaphawuleka unxibelelwano oluthile phakathi kwe-OFC ne-DLPFC. Ukudityaniswa kwangaphambili kwe-OFC kuqala ikakhulu kulwakhiwo oluphezulu lwe-cortical (2-3) kwaye ii-axons zazo ziphela zisilele ikakhulu kwizendlalelo ezinzulu (i-4-6), ehambelana nephethini yokuhambisa (IBarbas kunye neRempel-Clower, ngo-1997). Ngokuchaseneyo, i-OFC eqikelelwe kwi-PFC ye-latal isuselwa ngokuyintloko kwizendlalelo ezinzulu (i-5-6) nge-axons yazo iphelisa ubukhulu becala kuluhlu oluphezulu (i-1-3), isimbo esibonisa ingxelo. Le patheni ibonakala isebenza phantse kwi-70-80% yoqikelelo. Ke ngoko, ukuhamba kolwazi ukusuka kwi-OFC ukuya kwi-PFC ebubumbeneyo kubandakanya impendulo, ngelixa ulwazi luhamba kwelinye icala luhambelana nephethini ehambelana noko.
Imodeli yolwakhiwo iyacaphukisa kuba iphakamise ukuba imeko yonxibelelwano lwommandla ingangeniswa ngokusekwe kunxibelelwano lwe laminar. Ukuba imodeli yolwakhiwo ichanekile, iyasinyanzela ukuba sibenako ukondliwa kunye nesimo sengxelo nonxibelelwano lwommandla. Iimodeli ezenza ukuba i-PFC yecala isebenze ngokuyintloko okanye ngokukhethekileyo ngokusetyenziswa kweendlela ezisezantsi zinzima ukuvisisanisa neempawu zayo ezibalaseleyo zokutya. Ngokukwanjalo, iimodeli ze-OFC ezijonga njengokudlulisela iziphumo zekhomputer (umzekelo wexabiso lomvuzo) kwi-PFC yecala, ziyasilela ukubamba amandla alo mmandla kukhuphiswano oluzenzekelayo olwenziwa kwi-PFC yecala. Kodwa, njengoko kuchaziwe kamva kweli nqaku, imifuziselo ekhoyo yothethathethwano phakathi kwemimandla ye-PFC, naphakathi kweendawo ezibandakanyeka kukwiimvakalelo ngokweemvakalelo. Ewe, iimodeli ezikhoyo zihlala zifuna imisebenzi ye-PFC yecala, ngakumbi imisebenzi ye-DLPFC, ngolawulo olusezantsi, kwaye kunqabile ukuba kuthathelwe ingqalelo indawo ezinokuphuculwa ngokwendawo ezinjenge-OFC zinokubonelela ngefuthe eliphantsi kwimimandla ye-PFC.
Iipateni zeLaminar zonxibelelwano lwangaphambili-amygdalar
Uqikelelo lwe-amygdala olusemva kwe-OFC lugcina yonke imiqolo ye-cortex, kwaye ngenxa yoko ayinakuba nemida engqongqo yokuqikelela okanye uhlobo lwengqikelelo (UGhashghaei et al., 2007). Nangona kunjalo, kuyacaca ukuba kukho icandelo eliqinileyo lokutya kwezi ziqikelelwe ngokusekwe ekuphelisweni kwe-laminar. Ngokwahlukileyo, iingqikelelo ze-OFC eziya kwi-amygdala ikakhulu zivela kumaleko we-5, zibonisa ubume bazo njengoluvo lwengxelo (kucebisa ukuba basebenze ekuqhubeni inkqubo ye-amygdalar kunokuba kudluliselwe ulwazi oluthile njengeempawu zoluvo lwesizathu). Okubangel 'umdla kukuba, iingqikelelo zokutya ezisuka kwi-CCC yecala zibhekiswa kububanzi be-5 ye-OFC, olungumaleko wokuqala ophuma apho kuqikelelwa i-OFC ukuya e-amygdala.
Ngaba ulwazi lwe-anatomical lunokwazisa malunga neengxoxo zemisebenzi ephambili? Iminyaka eliqela edluleyo ibone ukuqhushumba komdla kwindlela ezihamba ngayo iingqondo ezahlukeneyo. Lo mdla uye kwinxalenye ubangelwe njengesiphumo sokuvela kweendlela zokuhlola unxibelelwano olusebenzayo kunye ne-fMRI, ukubonelela okokuqala amandla okuvavanya ukusebenzisana phakathi kwemimandla yobuchopho kubantu abaphilayo. Nangona kunjalo, iingxoxo ngezi datha, kunye neemodeli ezivela kule datha, azange zihlale zithintelwa yi-anatomy. Njengoko ezi ntlobo ziye zaba nefuthe, siyakholelwa ukuba ziluncedo ekuvavanyeni ukuba zilungelelana kangakanani na neuroanatomy echazwe apha ngasentla. Siyakholelwa ukuba ezo ntlobo zemodeli kufuneka zihambelane kunye neepateni ezidibeneyo zonxibelelwano ezidibanisa imimandla eyahlukeneyo yecortical kunye ne-subcortical kunye nengxelo / uhlobo lwendlela yezi ndlela. Xa iimodeli zingangqinelani nale miqobo, azinakho ukubakho, okanye ubuncinci zifuna inkcazo yokuba zinokwenzeka njani ngokunikwa ukungahambelani kwazo nokudibana kwengqondo eyaziwayo.
Uncwadi olukhulayo lwengqondo luzama ukuqonda indlela iinkqubo “ezibonakalayo” ezinxibelelana ngayo neenkqubo “zvakalelo”. Ngelixa kukho umda oqinisekileyo wokwahlulahlula phakathi kweenkqubo zokuqonda kunye neemvakalelo (I-Pessoa, 2008), umahluko uye wangqineka uluncedo ekuboniseni uluhlu lwezinto zokuziphatha ezinje ngommiselo wemvakalelo, inkuthazo, ukwenza izigqibo kwezoqoqosho kunye nolwalathiso lweendlela zoqwalaselo. Kumacandelo alandelayo, sichaza idatha evelayo kunye neemodeli zokulawulwa kweemvakalelo, imemori yokusebenza kunye nokusebenzisana kwangaphambi kweshishini, kugxilwe ekungqineni kwabo nedatha ye-anatomical. Sijolise ikakhulu kuncwadi lwemigaqo yolawulo lweemvakalelo, njengoko olu loncwadi lukhula ngamanani kwiingxoxo zengqondo kunye nonyango lweengqondo.
5. Ummiselo woLawulo
Ummiselo wommiselo uchazwe njengazo iinkqubo ezichaphazelekayo ekutshintsheni ukuqala, ixesha, ubungakanani okanye umxholo wempendulo yemvakalelo (Ubungakanani, 1998; Ubungakanani, 2008). Inkqubo yolawulo lweemvakalelo isusela kwizenzo ezenziwa kwakudala phambi kokuba kuvuke imvakalelo, njengokuchongwa kwemeko, kwezo nkqubo zibandakanyeke ngaphambi okanye kwangaphambi kokuba imvakalelo iqalile ukuvela, njengokuhambisa ingqalelo okanye ukuphinda uqapheleke kwakhona.Ubungakanani, 1998). Kukulezi ntlobo zokugqibela zobuchule apho uphando khona malunga nobudlelwane phakathi kwemimandla enxulumene nolawulo lwengqondo lwemvakalelo kunye nezo zinxulumene nokuphendula ngokweemvakalelo ziba nomdla omkhulu. Olu phando lubeka ngokungqalileyo okanye ngokucacileyo ichaze immiselo yemozulu njengothunyelo lwe-top-down, 'ukubanda' kwengingqi zolawulo lwe-PFC ukuya kulawula ezantsi, iinkqubo ezisebenzayo ezibandakanya imimandla engaphantsi ye-limby njenge-amygdala. Ukusilela ekufakweni ngempumelelo kwenkqubo ye-PFC yokulawula ubucukubhede okanye iinkqubo eziphezulu ze-amygdala eziphakanyisiweyo kucetyisiwe ukuba zibenegalelo kwiindlela ezininzi ze-psychopathology (IRottenberg kunye neGross, 2003; URottenberg noJohnson, ngo-2007).
Isicwangciso sokulawula iimvakalelo esithe safumana ingqalelo enkulu kuncwadi lwe-neuroimaging kukuphinda uqaphele kwakhona. Esi sicwangciso sokulawula siquka ukuqaphela ngenkcazelo yolwazi ngokweemvakalelo ukuze sitshintshe impendulo kwiimvakalelo (Ubungakanani, 1998). Ukuphononongwa kwakhona kufakwa kwinqanaba elibanzi leenkqubo ezinxulumene noko. Umzekelo, ukuphononongwa kwakhona kungagxila ekuguquleleni kwakhona intsingiselo yobuqu kwinto yeemvakalelo ukuyenza ngaphezulu okanye ngaphantsi kokuzithemba. Ngenye indlela, ukuphononongwa kwakhona kungagxila ekuchazeni kwakhona unobangela, isiphumo, okanye ubunyani bokubetheka ngokweemvakalelo ngaphandle kokutshintsha ubudlelwane bomntu kwisikhuthazo. Umzekelo, umntu unokuphinda afumane ingozi yemoto kwicala lendlela mhlawumbi ophela ngawo onke amaqela ahamba isigameko ngokukhuselekileyo. Inani lezifundo ezisebenzayo ze-neuroimaging ngoku zenziwe ngexesha lomsebenzi wokuphononongwa kwakhona, kwaye zidweliswe apha 1 Table, kunye nendawo yokusebenza kwe-PFC kuboniswe kuyo Umzobo 11. Sebenzisa amagama aphambili mgaqo wokumiswa, ukuphazamiseka kunye nokuphinda usebenze, amanqaku axhasayo ommiselo wokuzithandela wokulawulwa kweemvakalelo afakiwe. Ezi zifundo ze-fMRI bezibandakanya ukuphononongwa kwakhona kwengqondo okuyalelweyo, ukunyanzeliswa kweemvakalelo kunye nezifundo eziphazamisayo kwiindawo ezingezizo ezeklinikhi. Olu luhlu lwezifundo zokulawula iimvakalelo alupheleli; Umzekelo, ayibandakanyi iikhonsepthi ezinxulumene noko njengemodem regulation. Siphawula ukuba kuzo zonke iiTafile sigcine umbhalo obhalisiweyo (iilebhile zeBrodmann, okanye inkcazo engokusetyenziswa / yengingqi) esetyenziswe ngababhali bamaphepha obuqobo. Kukho iimeko ezithile apho imibuzo inokuphakanyiswa malunga nokusetyenziswa kweelebheli, kodwa sinqongophele “inkqubo yegolide” epapashiweyo yokulungelelanisa uninzi lweendawo ezingaphambi, asizitshintshanga iileyibhile, ngaphandle kokuba kwisicatshulwa siphawula ngokukhethekileyo ukusebenza kweVLPFC ezihambelana nenxalenye yangasemva ye-BA 47 / 12. Ukuphulukana nomda ocacileyo wenxalenye ye-BA 47 / 12 enokudityaniswa okubalulekileyo kwe-amygdala ebantwini, sijonga isahlulo somhlaba esingemva kwe-y = 32 njengemele ngokubanzi posterior BA 47 / 12. Sibonisa kwisicatshulwa xa i-OFC egxilwe iyahambelana nendawo ye-BA 13 (nokuba ithini).
Eyona paradigm ixhaphakileyo yokufunda ukuphinda ufundise kwakhona icela abathathi-nxaxheba ukuba bajonge ngokungathandabuzekiyo ukungqinisisa, ukuvusa umdla, imifanekiso eshukumayo (umz., Ukuguqula, ukubetha, ukubola, kunye nokubeka esweni) kunye nokuthelekisa ukusebenza kwe-neural ngexesha lokuvavanywa ukuphononongwa kwakhona kwengqondo ngovavanyo oluqondwe ngokujonga nje (Eippert et al., 2006; UKim et al., 2007; I-Ochsner et al., 2002; I-Ochsner et al., 2004; Phan et al., 2005; Urry et al., 2006; UVan Reekum et al., 2007). Ngelixa kukho umahluko kwiinkcukacha zemiyalelo yokuphinda ubeke esweni ukuya kufundo, bahlala becinga ukuba abathathi-nxaxheba benze utoliko olutsha lwentsingiselo, unobangela, iziphumo okanye ukubaluleka komfanekiso xa uvavanywa kwakhona. Ukuqwalaselwa ngokutsha kuthelekiswa nokujonga okungalawulwanga kwemifanekiso emibi kuthatha iindawo ezibanzi ze-PFC, kubandakanya ne-DLPFC yamazwe amabini kunye ne-VLPFC (ehlala ishiywe kakhulu kwicala), kunye nemimandla ye-ACC ye-dorsal kunye / okanye ye-PFC yokuxhasa njengokuxhasa iinkalo zolawulo lwengqondo. Umzobo 11 bonisa indawo yokusebenza kwakhona okunxulumene nokuvuselelwa (izikhombisi ze-cyan zokunciphisa isikhuthazo esingasasebenziyo, kunye nephuzi lokunciphisa isikhuthazo esihle) kwizifundo ezibonisiweyo.
Iparadigm ehambelana nayo isebenzisa imifanekiso eshukumayo ye-movie endaweni yemifanekiso emileyo. Olu phononongo lukwabonakalisa ukuqeshwa kwakhona kwe-DLPFC yamazwe amabini ngexesha lokuphinda uqashelwe kwakhona kodwa iyahluka ngokubhekisele kwimimandla ye-ACC kunye ne-PFC yasemva kokuba iphinde yaqeshwa ukuze inciphise usizi, intiyo okanye inkanuko yesini (I-Beauregard et al., 2001; UGoldin et al., 2008; I-Levesque et al., 2003, 2004).
Kwizifundo ezininzi zokuphononongwa kwakhona kusetyenziswa imifanekiso emileyo okanye eguquguqukayo, ukwehla kwe-amygdala kwasetyenziswa njengommeli otshintshileyo kwimpikiswano engendawo nokuvuselela kunye nokuncipha kokugaya abantu abafundisayo kwezinye izifundo (UGoldin et al., 2008; I-Levesque et al., 2003; I-Ochsner et al., 2002; 2004; Phan et al., 2005). Siphawula ukuba ukulinganisa okulula komsebenzi we-amygdalar kunye nefuthe elibi kuyingxaki, unikwe ukuba i-1) i-amygdala iba nakho ukusebenza kwiimeko ezingezizo ezimbi, kunye ne-2) amava axhasayo abandakanyekayo abandakanya amacandelo e-cortical kunye ne-subcortical eyandisa ngaphaya kwe-amygdala. Nangona kunjalo, unikwe umdla wethu kunxibelelwano lwengingqi yengingqi, ukumiselwa okuphantsi kwemisebenzi ye-amygdalar kubonelela ngesalathiso esifanelekileyo sokulinganisa ukuhlangana kwangaphambi kwembali ye-limbic nokuba kungakanani na umsebenzi wayo uhambelana nefuthe elibi. Uninzi lwezifundo zifumana ukwehla kwi-amygdala yasekhohlo, kwaye rhoqo kwi-amygdalae ephakathi, xa usebenzisa ukuphinda usebenze kwakhona ukulawula ulawulo olubi. Bambalwa kuphela abafundi abaphonononge ukuphinda bakhuthazwe kwakhona. Xa ubuzwa ukuba uphinde uqalise ukusebenza phantsi okanye ulawule isikhuthazo esifanelekileyo okanye ngokwesondo, inqanaba lokuvula le-amygdala lasekunene liye lancipha (I-Beauregard et al., 2001; UKim noHamann, 2007). Oku kungonyusa ingqikelelo malunga nokunyanzeliswa kwemigaqo yeemvakalelo, kodwa ngokubanzi, izifundo kuvavanyo lokunxibelelana ngokusesikweni kunye ne-amygdala kamva alikho.
Elinye icebo lokulawula iimvakalelo libandakanya ukuzisa engqondweni imifanekiso emihle okanye ethobisayo nokuba yeyendalo okanye yangaphambili okanye yokutshintsha okanye yokuchasana nefuthe elibi. Uvavanyo lokuziphatha kakuhle lubonisa ukuba ukukhumbula iinkumbulo ezingezonyani okanye imifanekiso enciphisa iyachaphazeleka kakubi (U-Erber kunye no-Erber, ngo-1994; UJoormann, uSeimer kunye neGotlib, ngo-2007; IParrot kunye neSabini, ngo-1990; Ukurusa kunye neDeHart, 2000). Izifundo ezibini ezinomtsalane ziye zathelekisa ukulawula imeko yomntu ngokubiza umfanekiso okhumbuzayo okanye inkumbulo kulindelo olungalawulwanga. I-Kalisch kunye noogxa (2005) uvavanyo olucatshulwe ngeethoni ezibonisa ukuba ngaba kunokwenzeka ukuba wothuke kwezo zilingo okanye hayi. Kwizilingo zommiselo, abathathi-nxaxheba bayalelwa ukuba bahlukane neemvakalelo zabo zoxinzelelo kwaye bacinge ngendawo ekhethekileyo echongwe ngaphambili. Kwizilingo ezingekho semthethweni, abathathi-nxaxheba bayalelwa ukuba bazibandakanye neempendulo zabo kwiimvakalelo. Uhlalutyo lwe-ROI lubonise ukuba le ndlela yommiselo yaqesha kwakhona indawo yengingqi ye-anterolateral yangaphambili (MNI: 42, 48, 18) kunye nommiselo wokubakho koxinzelelo lokufumana imimandla kwi-medic PFC kunye ne-rostral ACC (−4, 46, 28). Kwisifundo esifanayo, UDelgado kunye noogxa (2008b) Usebenzise iibhloko ezinemibala ukuyila uvavanyo apho kunokwenzeka ukuba wothuke, kwaye ucele abathathi-nxaxheba ukuba balawule uxinzelelo lwabo ngokuthi bakhumbule enye yeendawo ezimbini ezichongiweyo kwindalo. Uhlalutyo lwabo lwe-ROI lubonisa ukuba ukukhumbuza imifanekiso yendalo xa ulindele ukothuka iphinda ibambe i-front front gyrus (Talairach: −43, 28, 30). Indawo yokuma eyayinxulumene nempumelelo yomgaqo. Ummiselo kwakhona wakhokelela ekusebenzeni kudonga lwe-ventral medial kunye ne-cgenate ye-subralual (BA 32; −3 36, −8 kunye ne-BA 25; 0, 14, −11), abathi ababhali bayo bayayanyaniswa nokutshatyalaliswa (Iiphelps et al., 2004) kwaye iyancipha kwimisebenzi yasekhohlo ye-amygdalar. Ngelixa zombini ezi zifundo zisebenzisa iiparadigms ezifanayo, iindlela zazo zokuhlalutya kubandakanya ukukhethwa kwee-ROIs kunye nokumodareyithwa kweethonic ezichasene nethamsanqa zinokuba noxanduva lokumnye umehluko kwimimandla ekunikwe ingxelo ngayo ngokuzoba imifanekiso emihle okanye epholileyo yokulwa nexhala elinxulumene nokulinda okunokwenzeka wothuka.
Kuyafana nesicwangciso esiliqili sokumisela imvakalelo, ukuphazamiseka kubandakanya ukubamba ulwazi olungathathi hlangothi kunye nolungafanelekanga kwimemori yokusebenza komuntu. Uphando lwezinto oziphetheyo lubonisa ukuba ukwenza njalo kunciphisa ifuthe elibi kubo bobabini abantu abanesifo sokudakumba kunye nondysphoric (UFennell, Teasdale, uJones, noDamle, ngo-1987; ILyubomirsky, iCaldwell, kunye neNolen-Hoeksema, 1998; I-Teasdale kunye neRezin, ngo-1978). Ngokuthatha ubungakanani bememori yokusebenza ngokusebenza okunganyanisekanga kwengqondo, iingcinga zokomoya ezihambisanayo zithintelwe ekufikeleleni kwizibonelelo zononophelo (I-Minaer, i-2005). Izifundo ze-Neuroimaging zokuphazamiseka zisebenzisa ii-paradigms ezimbini ezahlukeneyo. Owokuqala, oqashwe ngu UKalisch et al. (2006), basebenzise ulindelo lwe-paradigm yothusayo, ngaphandle kwendawo yokuba abathathi-nxaxheba bakhumbule imemori emnandi okanye ekhuselekileyo, kukho umyalelo ophazamisayo ovulekileyo apho uthathi-nxaxheba wakhuthazwayo ukuba acinge ngayo nantoni na eyenye ngaphandle kokothuka. Le paradigm ichonge ummandla we-PFC yasekhohlo (MNI: −56, 30, 22) eyayisebenza ngakumbi kwizilingo apho abathathi-nxaxheba bayalelwa ukuba baziphazamise kunakwizilingo eziphazamisayo. Inkqubo yesibini yokuphazamisa ukubandakanyeka ibandakanyeka kumsebenzi wokuphazamisa owabelweyo (umsebenzi wokukhumbula umsebenzi waseSternberg) apho abathathi-nxaxheba babambe khona uluhlu lweeleta kwimemori yokusebenza ngelixa bejonga imifanekiso emibi okanye engathathi cala kunye nokulandela umfanekiso ophelileyo kufuneka uphendule ekubeni ileta enye ibikuyo iseti ababeyibambe engqondweni. McRae et al. (2009) Xela ukuba ukuzibandakanya kwimemori yokusebenza ngelixa ujonga izilayidi ezingalunganga xa kuthelekiswa nokujonga nje okunyuselweyo kuyandisa impendulo ye-BOLD ngakwesobunxele nangaphezulu gyri (MNI: BA6; −6, 10, 62 kunye −56, −4 , I-48 kunye ne-48, i-42, i-32; i-BA 9; −42, i-22, i-30 kunye ne-42, i-30, i-34; i-BA 10; I-36p; 62, 12, −38).
Uninzi lweengxelo ezilandelanayo zomgaqo wolawulo-mvakalelo ngokucacileyo ezibonisa imimandla ye-DLPFC njengokuzibandakanya kolunye uhlobo lolawulo lokuqonda kwaye ilumkisa ngokubonisa ukuncipha okufanayo kwimpendulo ye-amygdala kunxibelelwano oluthe ngqo kunye ne-amygdala. Kwimeko yokuphinda uqwalaselwe kwaye uphazanyiswe, oku kulumkiswa kufanelekile ingakumbi kuba ezi nkqubo zivelisa izinto ezijolise kuyo yonke iPFC (Umzobo 11). Njengoko bekutshiwo ngaphambili, iphethini yeendlela zokwenza izinto ezivela kwi-cortex zicebisa ukuba iindlela ezi ngqo ezivela kwimimandla ye-DLPFC akunakwenzeka ukuba zikhuphe ulawulo oluluqilima lokwenza i-amygdala. Iindawo ze-PFC ezinamaqashiso amdaka ngendlela ethile kwiFCC ifumaneka kuphela kwinxalenye encinci yeVLPFC, ngakumbi kwimimandla engasemva ye-BA 47 / 12. Ngelishwa, njengoko bekutshiwo ngaphambili, igama elichaziweyo elisetyenzisiweyo ukuxela ukusebenza kulo mmandla kuninzi lwezifundo kudala ubumbano xa kufikwa kwimibuzo yokunxibelelana ne-amygdala. Izifundo zokuphononongwa ngokutsha, inkumbulo eyakhayo okanye ukubandakanyeka komfanekiso kunye nokuphazamiseka kuxhaphake ingxelo yokusebenza kwimimandla ngokubanzi ye-VLPFC kunye ne-medC ye-OFC (U-Eippert et al., 2007; UGoldin et al., 2008; UKim noHamann, 2007; U-Lieberman et al., 2007; McRae et al., 2009; I-Ochsner, uRay et al., 2004). Ngokukodwa, uninzi lwezifundo eziphindiweyo ziphinda zenze ukuba kusebenze i-BA 47 / 12 xa kuncipha imvakalelo emibi okanye eyakhayo. Njengoko kuphawuliwe apha ngasentla, i-BA 47 / 12 yindawo enkulu enezinto ezinzulu kwaye ingumda we-BA 47 / 12 kuphela zindawo ezinokubakho kwe-amygdalar. Ke ngoko, iingxelo ezomeleleyo malunga nefuthe lokuqonda ngokuthe ngqo kwi-amygdala ziba nokucaca ngakumbi kwezo zifundo zenziwe zisebenze kweli candelo le-BA 47 / 12.
Imimandla yemedical ye-PFC ihlala ithathwa njengofumana ilungelo lokufikelela kwiingingqi ezincinci njenge-amygdala. Nangona kunjalo, ngokuhambelana nokunxibelelanisa ngokuthe ngqo kwimephu ye-medial kwi-amygdala, kuphela loo mimandla ye-cgenate cgenate (BA 25) kunye ne-dorsal ACC (BA 24) inexinene ngqo ne-amygdala. Izifundo kuphela UDelgado noogxa bakhe (2008a, 2008b) Xela ukugxila kumphezulu wezendondo okwimimandla ebekelwe ukuba nefuthe ngokubanzi kwi-amygdala. Ukunikezelwa kwedatha ye-anatomical, kunokubonakala kumangalisa ukuba ukusebenza kwe-BA25 akuveli rhoqo kwezi zifundo. Nangona kunjalo, kucacile ukuba ukuphuma kwesibonakaliso kwi-VMPFC yangaphambili kuthintele izifundo ekuboniseni usebenze ngakumbi kulo mmandla. Rhoqo, izifundo zokuthintelwa / zokucinezelwa, ukuphazamiseka kunye nokuphinda kuqwalaselwe ingxelo kuphela kwi-BA 32, enokubonisa ukuguqulwa okungaphezulu kwe-amygdala, xa kunikwe uhlobo lwe-BA 32 yegalelo kwi-amygdala.
Izifundo zeCorrelational deactivation ze-amygdala
Ukuqonda ngakumbi iinkcukacha malunga nendlela umda weemvakalelo ezisezantsi ezinxibelelana ngayo ne-amygdala, iseti yezifundo zemigqaliselo yeemvakalelo ihambele ngaphezulu komsebenzi ngokuchaseneyo nolawulo lokuphanda ulungelelwaniso oluthile lokwehla komsebenzi we-amygdala (jonga 2 Table). Oko kukuthi, endaweni yokubuza ukuba zeziphi iindawo ezenza umsebenzi owaziwayo ukubeka phantsi imisebenzi ye-amygdala, bavavanya ngokucacileyo unxibelelwano okanye ukusebenza / ukunxibelelana okusebenzayo phakathi kwe-amygdala kunye nengqondo yonke ngexesha lokusebenza kommiselo weemvakalelo. Ngenye indlela, ezinye izifundo ezihambelana ne-amygdala ziyancipha kunye nemimandla esele ichongiwe ngaphambili kusukelo oluphambili lommiselo. Olu phononongo lubonisa ukuba ukuncipha kwe-amygdalar kudityaniswa ngokungalunganga kwiindawo ezininzi zomsebenzi we-PFC. Okona kuqatshelweyo kukusebenza kweVMPFC, kubandakanya i-BA 11m / 14r (5, 37, −12; −6, 46, −20: Urry et al., 2006, I-Ochsner et al., 2002 ngokulandelanayo). Ukongeza, imimandla yangaphambi kokuzalwa kunye neyangaphambi kokuzalwa kwayo yaqwalaselwa ukuba inxulumene kakubi nomsebenzi we-amygdala ngexesha lommiselo. Njengokuba, Urry kunye noogxa (2006) uxele ummandla we-BA 32 / 10 (ubuninzi kwi-−23, 43, −10) eyandiswa ngokuqhubekayo nangokuphakathi. UDelgado et al. (2008b) Xela ulungelelwaniso olungafaniyo phakathi kwe-BA 32 (0, 35, −8) kunye ne-amygdala iyancipha. Iindawo zasePosterior (BA 13) ze-OFC nazo zidityaniswa kakubi kukusebenza kwe-amygdala deactivation (−24, 28, −14; 26, 24, −22: Banks et al, 2007: −30, 22, −16, 34, 24, 16, XNUMX, XNUMX, XNUMX I-XNUMX: I-Ochsner, uRay et al., 2004). Iindawo ezincinci zangaphakathi ze-PFC kwi-BA 47 (34, 54, 12) kunye ne-BA46 (−54, 12, 12: Urry et al., 2006; I-−54, 42, 12: I-Ochsner et al., 2002), nayo ivukile kwezi zifundo. Izifundo ezibini zidibanise imimandla ethile ye-DLPFC kwimimandla ye-medical, eyayihambelana nokuncipha kwempendulo ye-amygdala. Kwisifundo sika Urry et al. (2006)Uhlalutyo lokulamla lubonise unxibelelwano phakathi kwe-amygdala, BA 10 (3, 63, 18) kunye nommandla we-DLPFC (−50, 23, 19). UDelgado et al. (2008b) ngenye indlela basebenzise i-med X BAUMN yeXMUMX njengembewu kuhlalutyo lwabo lwe-PP emva koko ichonge indawo yase-amygdala kunye nomda we-DLPFC. Ngokubalulekileyo, ezi zifundo zichonga imimandla ehambelana ne-amygdala yokwehla eye yaqatshelwa apha ngasentla njengomba we-amygdala njenge-dorsal anterior cingate, i-cgenergen yangaphakathi kunye ne-posterior orbitofrontal cortex.
Kwimimandla ekunikwe ingxelo ngayo kolu hlalutyo lophononongo okanye uhlalutyo oluninzi lokuphonononga, inani labo linomda ngqo kwi-amygdala. Ezona ndawo ziqhelekileyo zidityaniswe ngokungalunganga nempendulo ye-amygdala yimimandla yangasemva ye-OFC kunye ne-CLLFC ye-FC eseleyo.Umzobo 12). Kwimimandla esembindini yangaphambili kuphela inxalenye engemva ye-BA 47 / 12 ineengqikelelo ezomeleleyo kwi-amygdala. Imimandla ye-engaphandle ye-BA 32 ikwachongwa kuhlalutyo lobumbano, olunokubonisa uqikelelo kuvavanyo kunye nakwi-basal lateral nucleus ye-amygdala (U-Cheba et al, 2001).
Iimodeli zokulawulwa kweemvakalelo
Ukuza kuthi ga ngoku, eyona datha iphambili iqhutywa yimodeli yokuqhutywa kwemvakalelo ivela kwisifundo sophononongo kwakhona Wager kunye noogxa (2008). Isiphumo sokuhluka komdla lutshintsho kuchaphazeleko oluzimeleyo oluchazayo. Indlela yokusebenza yokulinganisa yasetyenziswa kwidathabheyisi yokuhamba kwezinto ukusuka kwiparraisal paradigm efanayo naleyo yasetyenziswa I-Ochsner et al. (2002; 2004). I-VLPFC elungileyo yakhethwa njengesiqalo sohlalutyo, kunye nemithombo yolungelelwaniso egxilwe kwindawo ebandakanya ngokucacileyo isahlulo sangasemva sendawo ye-47 / 12 kunye nolwaziso lwe-amygdala. Ababhali baqale basebenzisa indlela ye-ROI yokuvavanya indima ye-amygdala kunye ne-nucleus accumbens njengabalamleli phakathi kweVLPFC elungileyo kwaye banciphisa nefuthe elibi elachongwa njengeyona metric eyintloko yokuphumelela kwakhona. Kolu hlalutyo lwe-ROI zozibini ezi zakhiwo zaboniswa ukulungelelanisa ubudlelwane phakathi kwe-VLPFC elungileyo kunye nokuhla kwengxelo kuchaphazeleka kakubi (bona Umzobo 13).
Ababhali ke basebenzise uhlalutyo lobuchopho olupheleleyo kunye nokuthobela okungathathi cala kuchonga iinethiwekhi ezimbini njengabalamli phakathi kobudlelwane phakathi kweVLPFC kunye notshintsho ekuchazeni ingxelo eyichaphazelayo (jonga Umzobo 14). Inethiwekhi enye inokhetho oluchanekileyo olungathanga ngqo ekunyuseni utshintsho kwimpembelelo ezimbi. Le nethiwekhi ibandakanya imimandla ye-nucleus accumbens, i-subgenual cingulate (BA 25), i-pre-SMA, i-precuneus, i-DMPFC (MNI: 24, 41, 40), kunye ne-front front gyrus (24, 21, 58). Phakathi kwale mimandla, i-nucleus accumbens kunye ne-subingual cingulate ineyona nto inxibelelanayo kakhulu ne-amygdala. Inethiwekhi yesibini echongiweyo inethagethi engathanga ngqo ekunciphiseni utshintsho lwempembelelo ezimbi kunye nokunciphisa impumelelo kwakhona. Le nethiwekhi ibandakanya i-rostral dorsal ACC, i-amygdala (kabini) kunye ne-poster-lateral ye-OFC (48, 24, −18). Umsebenzi wexesha elizayo kuya kufuneka ucacisa ukuba iinxalenye zenethiwekhi zinxibelelana njani nokuba ezi ntetho zikhethekileyo kolu hlobo oluthile lwesicwangciso sokumiselwa kwemvakalelo.
Iqela eliphandayo likhuphe imodeli yethiyori malunga neural neendlela zokusebenza emva kommiselo wemvakalelo. Eyona ilula kwezi modeli iphakamisa ukuba inani eliqingqiweyo leendawo linefuthe ngqo kwi-amygdala. UDelgado et al. (2008b), UHansel noVan Kanel (2008) kwaye IQuirk kunye neBhiya (2006) Isiphakamiso ngasinye sokuba i-podC ye-ventromedial ilawula imimandla ye-amygdala. Ezi modeli zibaluleke ngokuzama ukumisela ukuqonda kwethu iziseko ze-neuroanatomical zomgaqo wolawulo lweemvakalelo zoluntu kwimibhalo ebanzi yezilwanyana ekuphelisweni kunye nokunxibelelana kwe-PFC yokunxibelelana nabantu abahlanganayo kwi-basolateral amygdala (UMorgan, uRomanki kunye noLeDoux, ngo-1993; Likhtik et al., 2005; I-Quirk et al., 2000). IQuirk kunye neBhiya (2006) wakhela ubukho bazo zombini ezivuselelayo kunye nefuthe lesithintelo 'se-ventral' ye-PFC eqikelelweyo kwi-amygdala ebantwini nasegumbini. Ummandla wokuphinda uzalwe, i-BA 25, kuthiwa ayinabunkunkqele ngelixa i-dorsal engaphezulu kunye ne-anterior BA 32 icetywayo ukuba inonxibelelwano olonwabisayo kunye ne-amygdala. Zombini i-BA 25 kunye ne-32 zidibene ne-amygdala. I-BA 32, nangona kunjalo, inokunxibelelana okungaphezulu kakhulu.
Phillips et al (2008) Uphuhlise imodeli yesekethe ezizama ukucacisa ukungaphantsi kwe-neural yeentlobo ezininzi zemmiselo yomgaqo (jonga Umzobo 15). Imodeli inemimandla yecandelo le-DLPFC, OFC, VLPFC, DMPFC kunye ne-ACC. Eyona nto ibangela umdla, ababhali bahlula phakathi kweendawo ezichaphazelekayo kulawulo lweemvakalelo ezizenzekelayo (kwi-Agengenic ne-rostral ACC) kunye nemimandla eqeshwayo kulawulo lwemvakalelo ngokuzithandela (i-DLPFC kunye neVLPFC). Baphawula le mimandla yokugqibela njengeyona phylogenetically entsha kunye nokubonelela ngengxelo kwiinkqubo zokudala iimvakalelo ezindala. I-OFC, i-DMPFC kunye ne-ACC, kwelinye icala imimandla emidala ethe phylogenetically ichazwa njengokusebenza kweenkqubo zokuhambisa ingxelo yolwazi lwangaphakathi kwi-DLPFC kunye neVLPFC. Ababhali babeka i-DMPFC njengekhonkco apho i-OFC itya ulwazi lwexabiso ngaphambili kwimimandla yobuchopho kwiinkqubo zesigqibo.
Inxalenye eyodwa kule modeli kukucacisa ngokucacileyo kweenkqubo ze-feedxp kunye nengxelo. Imodeli inomdla inomdla kwaye inomdla ngokufanelekileyo kwimibono yesiko malunga ne-DLPFC isebenzisa ulawulo oluphezulu lwendawo engaphezulu kwemvakalelo. Nangona kunjalo, kunzima ukuvisisanisa le conceptualization kunye nemodeli yolwakhiwo, unikezwe usasazo olunamandla lokuqhagamshelwa kwe-PFC (IBarbas kunye neRempel-Clower, ngo-1997; IBarbas, 2000). Ewe, imodeli yolwakhiwo ibonisa ukuba ulwazi oluhamba phakathi kwe-DLPFC kunye ne-OFC ngokwenene luhamba kwicala elingafaniyo neenkqubo ezisuka kwi-OFC kwaye ziye kwi-DLPFC zibonakaliswe ikakhulu njengempendulo, kunye nezo ziphakamayo kwi-DLPFC kwaye ziye kwi-OFC ubukhulu becala iphawulwe njenge-feedirect.
I Phillips et al. Imodeli ikwabalulekile ekubekweni kwayo kwemimandla ebizwa ngokuba "ngummiselo wokuzenzekelayo" njengemimandla yokuzalwa kunye ne-OFC njengeyona ndlela iphambili apho imimandla emitsha ye-phylogenetically intsha ichaphazela iindawo ezinamaqhina njenge-amygdala. Oku kuyahambelana kakhulu (ngakumbi ingingqi engaphantsi yecingate) namalungiselelo othungelwano achazwe apha ngasentla. Kungacingelwa, nangona kunjalo, kunokuba kukho indlela engaphezu kweyodwa apho ulawulo lweemvakalelo ngokuzithandela lunokuchaphazela ukusebenza kwe-amygdala. Ngokukodwa, i-VLPFC yangemva inganakho ukuba nefuthe ngqo kwiinkqubo ze-amygdala ngaphandle kokufuna ukubandakanyeka kwenye yeendawo “ezizezomthetho ozenzekelayo” zommandla, inikezwe igalelo layo ngqo kwi-amygdala nuclei.
Isishwankathelo, ubutyebi bedatha bubonisa ukubandakanyeka kwemimandla ye-PFC ngexesha lomsebenzi wokumiselwa kwemvakalelo, kunye nomsebenzi kwiqela elikhethiweyo leendawo (BA 47 / 12, BA25 kunye ne-BA 32) bonisa imibutho enokwazi ukulawula imisebenzi ye-amygdala. Ukunyuka kweemodeli ezinobunkunkqele kucacisiwe ukuchaza le datha. Ukuvela kwezi modeli kuyathandeka, njengoko nenkxalabo iboniswe ngababhali babo ngobukho bothungelwano olucetyiweyo. Siqaphela, nangona kunjalo, ukuba akukho mizekelo ukuza kuthi ga ngoku evume ngokucacileyo iphethini ye-laminar yonxibelelwano phakathi kwemimandla eyahlukeneyo ye-PFC. Njengokuba, I-Wager et al (2008) inikezela ngemodeli eyinkimbinkimbi yesicwangciso esithile sokumiselwa kwemvakalelo, kodwa ayisijongi imeko yokuhamba kolwazi phakathi kwemimandla yecandelo. Phillips et al. bandakanya ngakumbi umbono wenkcazelo yokudluliselwa kwengcaciso kunye nengxelo, kodwa ungangqinelani nezi ngcinga kunye nephethini yengxelo egciniweyo yengxelo kunye nothelekelelo lwendlela kwimimandla ekubhekiswa kuyo. Sikholelwa ukuba ukulungisa le micimbi kubonelela ngomnye wemiceli mngeni ophambili kubaphandi bezama ukuqonda i-neural substrates of regression memo.
6. Ukulawulwa kokuqonda kokuphazamiseka ngokweemvakalelo
Ngelixa uninzi lohlalutyo lwethu lujolise kwizifundo zokulawulwa kwemvakalelo, imiba emininzi efanayo iyavela xa kuthathwa uncwadi kulawulo lwengqondo. Ngokubanzi, ulawulo lwengqondo lubhekisa kwiinkqubo zolawulo ezikumgangatho ophezulu ezikhuthaza ukusebenza ngokuchanekileyo, ngelixa kuthintela ukwenziwa kwezinto okungafanelekanga. Eli gama lisetyenziswa ngokukhethekileyo ekusebenzeni kwimisebenzi efuna ingqalelo ekhethiweyo yolwazi lweemvakalelo olungenayo kunye nokuthintelwa kolwazi lweemvakalelo ezingafanelekanga, kunye nokukhethwa okuhamba kunye kweempendulo zokukhuthaza injongo kunye nokucinezela iimpendulo zokhuphiswano olungafanelekanga. Inkqubo enjalo yokhetho ihlala icaciswa gca ngokwemeko yokuguqulela okuphezulu kunye nokucalulwa kweendlela zokwenza. Izifundo zokulawulwa kwemvakalelo zinokujongwa njengecandelwana elithile lolawulo lwengqondo olujolise ekumodareyelweni kwempendulo ehambelana nayo. Ngokwahlukileyo koko, uninzi lolunye uhlobo lwezifundo zolawulo lokuqonda olubandakanya ukugxininisa kwimvakalelo kwisakhono sokunqoba ukuphazamiseka okubangelwa yimvakalelo. Ngenxa yeempawu zabo zendalo (ezihlala zizenzekelayo) zokufumana ingqalelo (Uninzi et al., 2005; 2007; I-Pessoa, 2008), iimvakalelo ezishukumisayo zihlala zivusa isidingo esomeleleyo solawulo lokuqonda ukuze kugcinwe ulwazi olululo lweenjongo ezifanelekileyo. Le mfuno yokuphepha ukuphazanyiswa kukuvuselela ngokweemvakalelo ngokukodwa kwenzeka kwizifundo apho imvakalelo yokuqhubela phambili isenzeka ngaxeshanye kunye nesinye isikhuthazo, ukungahambelani nezinye iimfuno zomsebenzi, okanye ngexesha lokusebenza kwememori yokusebenza, apho ukuphazamiseka kungaphazamisana nolondolozo lolwazi olusemgceni. Sizijongisisa ngokufutshane ezi zifundo ngenjongo yokuqaqambisa ukuhlangana kwazo ngoncwadi olunokumisela iimvakalelo. Ukuphononongwa ngokucokisekileyo okucacileyo kolu fundo ngabafundi ekubhekiswa kubo Banich et al. (2009).
Uxinzelelo lweemvakalelo ezishukumisayo ngexesha lemisebenzi yokuqonda
Izifundo ezininzi zisebenzise iiparadigms apho abathathi-nxaxheba kufuneka baphendule kwindawo engeyiyo eyombonakalo yomfanekiso weshukumisi (onjengombala) kwaye bangaphazanyiswa ngumxholo weemvakalelo (okt, amagama axhasayo), okanye ukubambelela kunyuselo olungelulo (okt. indlu) ngelixa ungayinaki into evuselela iimvakalelo (ubuso oboyikisayo). Umzekelo, imimandla ye-rostral (dorsomedial, pregenual and dorsal ACC) kwaye zombini imimandla ye-DLPFC kunye ne-VLPFC zonke ziye zaqwalaselwa kwimibono ye-Stroop paradigms apho izifundo kufuneka zikuphephe ukuphazanyiswa ngumxholo wemvakalelo wamagama (UWhalen et al., 1998; ICompton et al., 2003; UHerrington et al., 2005; UMohanty et al., 2007). Ukuphononongwa ngokucokisekileyo ngakumbi kokulawulwa kwengqwalaselo kunye nolawulo lweemvakalelo olunokuthi lubandakanye abafundi abasebenza ngokumajukujuku UBlair noMitchell (2009) kwaye I-Mitchell (2011).
Umda wokutolika uninzi lwezi paradigms ziphakamayo nangona kungasoloko kucacile ukuba le mimandla iyabandakanyeka na ngenxa yokuba isebenzisa ulawulo lwengqondo, ukubeka iliso kwimpikiswano, kubandakanyeka kungquzulwano olukhulu okanye ukuphazamiseka ngaphandle kokulawula ungquzulwano / ukuphazamiseka, okanye bamane baphendula kubume beemvakalelo zesimo esishukumisayo. Umzekelo, UMohanty kunye noogxa (2007) bonisa ngokubonakalayo ukuba ingingqi yentshukumo yangaphambili (malunga ne-BA 24 / 32) ibonisa ukunyuka kokusebenza ngexesha lomsebenzi we-Stroop ngamagama ovakalelo, kwaye oku kuyangqinelana nokwanda kwexesha lokuphendula emsebenzini. Oku kunokutolikwa ngokwemigaqo ye-RACC yokuzibandakanya ukuze kusebenze ukulawula kwengqondo kubaphazamisi bemvakalelo. Nangona kunjalo, inikwe into yokuba ukusebenza kwale ndawo kuyahambelana nexesha elininzi lokuphendula, inqanaba lokuvula kwalo akubonakali ngathi linxulumene nempumelelo yabaphazamisi. Ngaphezu koko, ibonise ukunyuka kokusebenzisana kunye ne-amygdala, nto leyo ngokucacileyo ingahambelani ne-hypothesis yokuba i-RACC yayiqhuba umthetho olawula i-amygdala. Ewe kona kuyaphawuleka ukuba ababhali bacebise ukuba endaweni yokubonakalisa ukulawulwa kwe-RACC ye-amygdala, ukuphakama koqhakamshelwano ngexesha lokuvezwa kwezinto eziphazamisayo ezinokubonakalisa ukulawulwa kwe-amygdalar okanye ukufakwa kwi-RACC, kunokuba kungenye indlela ejikelezayo.
Phakathi kwezinye iziqwenga ezibubungqina obuhle kakhulu bokulawulwa kwangaphambili kwengqondo kulawulo lweemvakalelo kwi-amygdala ivela kwisifundo ngu Etkin et al. (2006), apho abathathi-nxaxheba benze umsebenzi we-Stroop-apho iimvakalelo zobuso zinokuba zihle okanye zingachanekanga ngamagama abiza imvakalelo. Uyilo lophononongo lwalunzima ngokungaqondakaliyo njengoko ababhali bengagxili ekuthelekiseni okulula kovavanyo lweemeko okanye ukungathathi cala kwezilingo, kodwa bavavanya iziphumo ngexesha lolingo olungelulo olwalandela ngqo uvavanyo oluhle okanye olungelulo. Into ebangela umdla kukuba i-DLPFC, ingingqi ye-DMPFC kwindawo ephambili ye-gyrus, kunye ne-rostral (eyenzelwe yona) i-ACC ibonise ukwenziwa kwezinto ngexesha lokulingwa okungalunganga obekuxhomekeke ekubeni ulingo lwangaphambili lwalulungile okanye akunjalo. I-DLPFC (kunye ne-DMPFC) iphendule enkulu kwizilingo ezingezizo ezalandela emva kovavanyo olufanelekileyo, ngelixa i-ACC ye-rostral yaphendula kakhulu kwizilingo ezilandela olunye uvavanyo olungelulo. Olu phononongo lolunye lophononongo ezimbalwa kuncwadi lolawulo lwengqondo ezivavanya ngokuthe ngqo ubudlelwane bemimandla yecortal yecortical nomsebenzi we-amygdala (kusetyenziswa uhlalutyo lonxibelelwano lwengqondo. UFriston et al. 1997). Ngokumangalisayo, umsebenzi omkhulu kwi-rostral ACC yayihambelana kakhulu nomsebenzi we-amygdala. Ngokusekwe kwipatheni yeempendulo ze-amygdala, ababhali baxoxa ukuba umsebenzi we-amygdala ulungelelaniswe nenqanaba lokungqubana kwityala elinikiweyo, kwaye ngokunyanzela umsebenzi we-amygdala i-rostral ACC ibonelela ngolawulo kule ngxabano. Inkxaso yolu luvo ivela kwidatha yokuziphatha ekubeni abo babonisa umsebenzi omkhulu onxibelelanayo kwizilingo ezingalunganga babonisa ukusonjululwa kwempixwano enkulu njengoko kulinganiswa ngamaxesha okusabela emsebenzini. Kwisifundo esilandelayo Etkin et al. (2010) Uqwalasele ukuba le uxinzelelo lomsebenzi we-amygdala ubonakala buthathaka kwizigulana ezinesifo sokudakumba ngokubanzi esihambelana nolawulo olusempilweni, ukubonelela ngokulungelelanisa okunokubakho kwe-neural kobunzima bokulawula ukuphazamiseka ngokweemvakalelo okanye ungquzulwano kuluntu lwezigulana.
Ipakiat ibalulekile ekufanele ukuba yenziwe ngokubhekisele kolu loncwadi. Okokuqala, uphononongo olwenziwe liqela lika-Etkin aluchazi ubukho bothintelo lwethoni ye-amygdala yimimandla ye-PFC ngexesha lokungqubana kolwazi lweemvakalelo, okanye ukubandakanyeka rhoqo "kwemimandla yolawulo lokuqonda", kodwa kunokuba ngumsebenzi othile othintele kuxhomekeke kwinqanaba. ungquzulwano phakathi kwangoko kwangoko. Ukuba kuchanekile, ukubanakho kokubona ukunxibelelana okungafaniyo phakathi kwendalo yangaphambili (okanye eminye imimandla ye-PFC) kunye ne-amygdala kunokuba ngumsebenzi omkhulu nohlahlelo oluthile.
Eminye imigca yobungqina ikwaphakamisa ukubakho kwezinye iindawo eziphambili, ngakumbi i-ACC, inokubangela ulawulo lwe-inbitory kwi-amygdala. Umzekelo, kwisifundo usebenzisa iparadigm efanayo ne-Etkin et al. (UChechko et al., 2009), abaguli abanesifo sokungakhathali babonakalise ukucothisa okukhulu kunolawulo olusempilweni ngexesha lovavanyo olungenangqondo ngokweemvakalelo, kunye ne-amygdala ephezulu, kodwa umsebenzi we-dorsal ACC / DMPFC, okhokelela kwisimvo sokuba ukuphazamiseka kwengqondo kuphawulwa ngokungalunganga kwe-DMPFC / dorsal ACC control. Ngokufanayo, IHariri et al., (2003) Kuqwalaselwe ukunxibelelana okungalunganga phakathi kwe-amygdala kunye ne-dorsal ACC (kunye ne-VLPFC) xa izifundo kwakufuneka zifakele iilebhile ngokuchaseneyo nemifanekiso yeemvakalelo (kunye nomsebenzi we-amygdala wonyuka ngemeko yomdlalo, kunye ne-VLPFC kunye ne-dorsal ACC eyandisayo yomsebenzi ngexesha lelebheli). Kucetyisiwe ukuba i-dACC inokukhupha ulawulo lwe-amygdala nangona kungekho kungqubana okuthile okanye ukuphazamiseka ngokweemvakalelo komsebenzi. UPezawas et al. (2005) Kuqwalaselwe ukuhlangana okungathath'inxaxheni phakathi kwe-DACC kunye nomsebenzi we-amygdala ngexesha lomsebenzi osongelayo wobuso. Kananjalo kuya kuqatshelwa ukuba i-ACC yangaphantsi kwisifundo sePezawas yayihambelana kakuhle nomsebenzi we-amygdala, iphakamisa ukuhlangana okungafaniyo phakathi kweendawo ezahlukeneyo ze-cingate kunye ne-amygdala, kunye nokuphakamisa ngakumbi, njengakwindawo UMonhaty et al. (2007) iphepha, ukuba i-RACC, ubuncinci kwiimeko ezithile ngokuqinisekileyo, kunokuba kakubi, idityaniswe ne-amygdala.
Inkumbulo yokusebenza
Olunye ugcino lweenzame zolawulo lokuqonda lujolise kwisakhono sokucinezela ukuphazamiseka ngokweemvakalelo ngexesha lomsebenzi wememori yokusebenza. Kuba ubungakanani bolwazi olunokubanjwa luze lusebenze kumgca we-intanethi lilinganiselwe (I-Cowan, i-2010), kubalulekile ukuba abantu babeke phambili ngokufanelekileyo ukuba loluphi ulwazi olungena kule venkile. Ngokufanelekileyo, kufuneka sigcine injongo efanelekileyo yolwazi enokuhambelana nolwazi olungabalulekanga kangako, kodwa sikwazi nokulahla imixholo yememori yokusebenza xa ulwazi olubaluleke ngakumbi luhambisa ngaphezulu kweenjongo zangaphambili. Ngenxa yoko, inkumbulo yokusebenza inikezela ngendawo enokuba luncedo ekuvavanyeni ukusebenzisana kwemvakalelo, ngakumbi inikwe indima ebalulekileyo ye-DLPFC kunye ne-VLPFC kwiinkqubo zememori yokusebenza (I-Badre et al., 2005; I-Blumenfeld et al., 2010; UCurtis & D'Esposito, ngo-2004; UJonides et al., 2005; Umrhumo kunye neGoldman-Rakic, 2000, Nee & Jonides, 2010; Iposti, 2006; UThompson-Schill et al., 2002).
Iingxelo ezimbini ezenziwe ngabakwaDolcos kunye noogxa babo inqaku njengoko banxibelelanisa ngokukhethekileyo ukusebenza kobuchopho kwintsebenzo enempumelelo okanye ukucofa kwimicimbi yokudityaniswa kokusebenza (IDolcos kunye neMcArthy 2006; I-Dolcos et al., 2006). Zombini ezi ngxelo zahlalutya idatha ukusuka kulungelelaniso olulula lokuphendula olucothayo lomsebenzi apho imifanekiso yovakalelo okanye yokungathathi cala iboniswe ngexesha lokulibaziseka (ukugcinwa) kwexesha lomsebenzi. Kwisifundo sokuqala, babonisa ukuba i-ventrolateral cortex (BA 45 / 47) yasebenza kabini ngexesha lokubonana ngokweemvakalelo nabaphazamisi abangathathi hlangothi. Abathathi-nxaxheba ababonise umsebenzi omkhulu wokuhambahamba ngeenyawo phambi kwabaphazamisi ngokweemvakalelo bakala ezo zinto zingabaphazamisi kangako. Kuphononongo olwenziweyo lubonisa ukuba umsebenzi we-BA 45 washiyekileyo (kodwa awulunganga umsebenzi we-BA 45) umahluko phakathi kwezilingo apho abantu baphumeleleyo ukulandela.okungawunanzi umsebenzi ophazamisayo (njengoko kubonisiwe ngokuchanekileyo okanye ngendlela engalunganga yokusebenza kwempendulo). UDolcos et al. (2006) Nika ingxelo nge-VLPFC- amygdala unxibelelwano olusebenzayo, kunye neendawo zombini ezanda ngexesha lokubonana ngokweemvakalelo kuvavanyo lokuphazamisa ukungathathi hlangothi. Ngokubalulekileyo olu nxibelelaniso lukwindlela elungileyo kwaye alunakho ukutolikwa libonisa uxinzelelo lokudubula i-amygdalar.
Izifundo ezenziwe yiDolcos kunye nabalingane bakwabonelela ngobungqina beepatheni ezingasebenziyo zokwenza isebenze kunye nokusebenza ngokuzimeleyo kwimimandla engaphambili. Ngokukodwa, iindawo zokwenza i-ventrolatal ziye zanda ngeziphazamiso ngokweemvakalelo, ngelixa i-DLPFC (BA 9 / 46) iyancipha, iphakamisa ubudlelwane bokubuyelana phakathi kwale mimandla. Obu budlelwane buphinda bufana ne-dorsal vs. patral patheni egcinwe yi U-Perlstein et al. (2002) owayenezifundo zenza umsebenzi wokukhumbula owakhutshwa kwimifanekiso engqinelweyo ngokweemvakalelo njengemisebenzi efanelekileyo kunye nezinto ezibonakala njengezinto ezinomdla kunye nomdla [onomdla, ubudlelwane bokuphindwayo babunxulunyaniswa ngokusondeleyo kunye ne-DLPFC yokunyuka kunye nemimandla enomdla kunye ne-ventral (BA 10 / 11) ibonisa umsebenzi owandisiweyo kwisikhuthazo esingalunganga]. Iphethini eguqukayo phakathi kwemimandla engakumbi ye-dorsal kunye ne-ventral ye-PFC nayo iye yaqwalaselwa kwezinye iindlela zokusebenza kwememori, kunye ne-DLPFC enkulu enxulumene nomsebenzi wangaphandle we-ventral onxulunyaniswa nomthwalo omkhulu wenkumbulo yokusebenza (URypma et al., 2002; UWoodward et al., 2006), nangona imimandla ye-PFC yecandelo elithile ebandakanyeka kwezi zifundo iyahluka. Ipateni ebonakalayo yokwahluka kwengingqi kunye ne-dorsal yendawo iphakamisa ukungavisisani okukhoyo phakathi kwale mimandla, kodwa ungabonisi imeko yolo lwalamano. IRanganath (2006) iphakamisa ubume bendawo yokusebenza kwiinkqubo zememori yokusebenza apho imimandla ye-caudal / ye-ventral ye-PFC ibonelela ngolawulo olusezantsi lweenkqubo zangasemva, ngelixa i-dorsal / rostral PFC ibonelela ngolawulo lwemimandla engaphambili ye-caudal yangaphakathi. Ngokwolu luvo, iRanganath ithi iinkqubo zokhetho eziphunyezwe yi-rostral / dorsal PFC zibandakanya ukuguqulwa kwemisebenzi kwi-caudal / ventral PFC. Nangona kunjalo, njengoko kuchaziwe ngezantsi, iimodyuli ezikhokelela kwicala elahlukileyo nazo ziya kuthathelwa ingqalelo.
7. Ummiselo olungileyo weendawo zokuqonda
Ngenxa yemodeli yolwakhiwo echazwe kumacandelo angaphambili, uqikelelo lwe-OFC kwi-CCC yecandelo, kubandakanya ne-DLPFC inokwahlulwa ngokokubonelela ngengxelo. Ngenxa yoko, oluqikelelo lunokunikezela ngomkhethe kunye nokulawulwa kwemimandla ephuculweyo yoyilo. Nangona kubonakala ngathi kuchasene nembono yentanda-bulumko ebeka uburharha ngaphezulu kwemvakalelo, uluvo lokuba indawo ebandakanyeka kulungiso lobambiso inokubonelela ngohlobo lokuphendula ngokubandakanyeka kwiindawo ezibandakanyeka kolunye unxibelelwano lwengqondo luhambelana ngokulula nezimvo zanamhlanje zemvakalelo egxininisa amandla emvakalelo wokubeka phambili kunye nokukhetha Ukulungiswa kolwazi ukuze kuququzelelwe iinjongo zebhayoloji kunye nezentlalo. Olu luvo lweemvakalelo luchazwa ngokukhethekileyo nguGrey kunye noogxa (Grey, 2001, Grey, uBraver, uRaichle, i-2002), ngubani oyeka le ndlela kunye nokurhoxa ilizwe ngokuphembelela ngamandla ekusebenzeni kwemisebenzi ethile yengqondo, kokubini kuphucula kunye nokuphazamisa imisebenzi eyahlukeneyo yokuqonda ukuze kuhlangatyezwane neemfuno zomsebenzi. Ubungqina bokumodareyitha ngokweemvakalelo kukwamkelwa kakuhle ekwenzeni izigqibo (Delgado et al., 2003; IGrabenhorst kunye neeRoll, 2009; IHardin, iPine kunye ne-Ernst, 2009; I-Piech et al., 2010), kodwa nayo inokuqwalaselwa kweminye imisebenzi yangaphambili yokulamla njengememori yokusebenza. Umzekelo, ukusebenza kwendawo yokusebenza kwememori yokusebenza ngaphakathi kuguqulwa ngendlela eguqulweyo ngokuqalwa kweemeko ezichaseneyo ezimbi, ngememori yokusebenza yendawo iyaphuculwa yimeko yokuphelisa imeko kunye nokuphazamiseka yimeko yendlela, kunye nememori yokusebenza yomlomo ebonisa isiphumo esichaseneyo (Grey, 2001). Ukongeza, ulwazi lweemvakalelo ezilungileyo nezingalunganga zinciphisa ukuphazamiseka kwememori yokusebenza ngokuthelekiswa nolwazi olungathathi hlangothi (Ilevens kunye nePhelps, ngo-2008; 2010). Ilungelo le-OFC (i-33 24 −8) kunye ne-insula engaphandle ye-insula (−32 21 2) iphendula ngakumbi kwisisombululo sokuphazamiseka ngokweemvakalelo.
Kwangokunjalo, kwindlela yokusebenza eguqukayo yokuqonda okubonakalayo, ukufunyanwa kwempembelelo elungileyo, xa kuthelekiswa nokungathathi hlangothi okanye ukungachaphazeli, kukhuthaza ukuguquguquka kwengqondo kunye nokuncipha kokunyamezelana, kodwa kukhokelela kukonyuka kokungaqondakali (UDreisbach kunye neGoschke, ngo-2004). Iziphumo ezifunyenweyo ziyahambelana nomzimba okhulayo wobungqina bokuba iimeko ezakhayo nezingalunganga zemeko zinokwenza banzi okanye zithathelwe ingqalelo ngokuxhomekeke kumandla wendlela okanye iimpawu zokurhoxa kwimeko yemeko yokuma (UFredrickson kunye no-Branigan, ngo-2005; IGable kunye neHarmon-Jones, ngo-2008; 2010; IGasper, kunye neClore 2002).
Ngokucacileyo, ubungqina obukhulayo bubonisa ukuba iimpembelelo zempembelelo ezinempembelelo BONAKUphendula kwi-DLPFC (BA 9) ngexesha lomsebenzi wenkumbulo yokusebenza (Ngwevu, iBraver, kunye neRaichle, 2002; I-Savine kunye neBraver, ngo-2010). Kanjalo, I-Savine kunye neBraver (2010) bonisa ukuba kwi-DLPFC yasekhohlo (i-BA 9), umvuzo wokukhuthaza imali ngokunyanzelekileyo ophuculwe umsebenzi-onxulumene nokusebenza kwe-cue, kwaye oku kusebenze kuye kwaxela kwangaphambili ukuba ityala liza kwenziwa ngokugqibeleleyo. Ukuthathwa kunye, izifundo ezinjalo zifuna ukuphinda zitshintshe kwakhona iimbono ezingacacanga zobudlelwane phakathi kweenkqubo zokuqonda kunye neemvakalelo.
Izifundo zeseli ezizodwa zibonelela ngeminye imbonakalo eyongezelelweyo kwikhosi yexesha lokunxibelelana phakathi kwemimandla yangaphakathi kunye nangaphezulu, ngokubhekisele kumvuzo. Idatha evela kwimimandla yokuqala iphakamisa ukuba ikhowudi ye-orbital ikhowudi yoxabiso olucekeceke lomvuzo kunenye imimandla engaphambili, nokuba i-OFC ibonelela ngolu lwazi lokuxabisa kwiindawo ezingakumbi zangaphambili (IHikosaka kunye neWatanabe, 2000; IWallis kunye neMiller, ngo-2003; URushworth et al, 2005). Ngokubalulekileyo, i-neuron ye-OFC ibonisa iimpendulo zomvuzo wolwazi owandulelayo kwiimpendulo ezinxulumene nomvuzo kwi-DLPFC (IWallis kunye neMiller, ngo-2003). Oku kudibana nombono wokuba i-OFC yokuqala ikhowudi ixabiso lomvuzo kwaye ke itya le ngcaciso kwiindawo ezinokwazi ukunxibelelanisa olu lwazi ngezenzo okanye kolunye ulwazi oluyimfuneko ukufumana ukufikelela kumvuzo. Siqaphele, nangona kunjalo, ukuba ayicacanga ukuba le ndlela yokukhuthaza ifikelela kangakanani kwi-DLPFC ngokubhekisele kwingxelo yohlobo lwengxelo, okanye inokuthathwa njengento ehambelana nendalo, njengoko ezinye iingqikelelo ze-30% ze-OFC, DLPFC zingathathelwa ingqalelo. indalo (IBarbas kunye neRempel-Clower, ngo-1997). Ngokobume bemodeli yokumahluko oku kungacacisa ukuba uvakalelo lomvuzo weeseli ze-DLPFC lubonakalisa uhlobo lokuphendula ngokuthanda i-DLPFC okanye lubonisa uhlobo olulula (lokuhambisa) lolwazi kuxabiso lokusetyenzwa kwexabiso le-DLPFC. Nceda ujonge I-Mitchell (2011) uphononongo lwendlela imixokelelwano ye-neural yomvuzo enokudlula ngayo kulawulo lweemvakalelo.
Uluvo lokuba ukuqhubekeka ngokweemvakalelo luchaphazela ukusebenza kwengqondo nako kunokuba luncedo xa kuqwalaselwa unxibelelwano olusebenzayo phakathi kwe-amygdala kunye nemimandla yangaphambili. Njengoko bekutshiwo ngaphambili, unxibelelwano olusebenzayo phakathi kwemimandla ye-PFC (ngakumbi ukwenziwa kwe-cingrate kunye ne-VLPFC) lubuye lwaphawulwa kwizifundo ezidlulileyo (IPezawas et al., 2005; I-Dolcos et al., 2006). Sicebisa ukuba kwezi meko i-amygdala inokuba ngumqalisi, kuba inokubangela ukuthambeka kwemeko kuqala, kwaye inxibelelane nalo lwazi okanye izame zokulawula imimandla ye-PFC isekwe kulolo lwazi kunenye indlela ejikeleze. Nangona kunjalo, ukuza kuthi ga ngoku bambalwa amalinge enziweyo okubonisa indlela yesalathiso yolu nxibelelaniso lusebenzayo.
8. Ingxoxo
Sikholelwa ukuba uphononongo olungentla lubonisa isidingo sokungena kwiinkcukacha zonxibelelwano lwe-anatomical ngaphakathi kwi-PFC kunye nobudlelwane babo ne-amygdala xa kujongwa ukuhlangana phakathi kwemvakalelo. Ukusilela ukwenza njalo kungakhokelela kwiimodeli ekunzima ukuzivisisanisa ne-anatomy, kwaye ngenxa yoko zinokubonakalisa ukuba azichanekanga. Ngokwahlukileyo koko, ukujongwa kweenkcukacha ze-neurocircuitry kungekuphela kokubonelela ngakumbi kwiimodeli ezinokubakho zonxibelelwano phakathi kweenkqubo zemvakalelo kunye nengqondo, kodwa kunokubonakalisa iipropathi ezisebenzayo ngenye indlela ebezingenakuhanjelwa kuzo.
Izibalo zoMmiselo woXinzelelo
Ngokusekwe kukhetho lwendlela ye-neuroanatomical phakathi kwe-PFC kunye ne-amygdala, imizekelo ebonakalayo yokuguqulwa kwe-PFC ngokwemfuno kufuneka ibandakanye ukuguqulwa, okanye ukuphinda kudluliswe, kwi-dorsal anterior cingulate, indawo engaphandle yokungena kwi-rectus ye-gyrus, okanye kwicala elingasemva lendawo. I-47 / 12. Kweli nqanaba lenkundla, iinkcazo ezilula zokuba iPFC ibandakanyeka kulawulo lweemvakalelo ibonelela ngeenkcukacha ezingonelanga ukuba ziluncedo, kwaye kwiimeko ezininzi zinokuthi zilahlekise, njengoko uninzi lweengingqi zePFC zingenalo uqikelelo olomeleleyo kwi-amygdala. Ukuvela kweemodeli zendlela ezigxila kwii-node eziphambili zeprojekthi kwi-amygdala, njengemodeli eziphakanyisiweyo kwaye zavavanywa nguWager et al. kunye noPhillips et al. luphuhliso olukhuthazayo malunga noku. Sikrokrela ukuba njengenkqubela phambili eyenziweyo ekuqondeni ukubandakanyeka kwee-PFC kulawulo lweemvakalelo, iindima ze-dorsal anterior cingulate, indawo engasemva ye47 / 12 kunye nengingqi engaphantsi kolawulo lwe-amygdala iya kudinga ukumiselwa.
Umbuzo ophambili usahleli malunga nendlela ukwenziwa kwe-PFC ebanzi kakhulu evelayo ngexesha lolawulo lweemvakalelo inxulumene nale ndawo zingamanyani, kuba zimbalwa nje ezihlolisisa unxibelelwano lwe-intra-PFC. I-Anatomically, ezi ndawo ze-PFC azinxibelelani ngokulinganayo kwi-dorsal anterior cingate, posterior 47 / 12 okanye kwingingqi engaphandle, kwaye ke zinokwenzeka ukuba zidibane ngokukhetha kunye neendlela ezahlukileyo kwi-amygdala. Sikrokrela ukuba ukuqonda ngokupheleleyo ukubandakanyeka kwe-PFC kulawulo lweemvakalelo kuya kufuna ukuba kucaciswe ukuba mangaphi la mimandla ye-PFC engenalo ulungelelwaniso ngqo olusebenzayo inxibelelana neminye imimandla ye-PFC enolingano olwaneleyo lokuguqula ukulungiswa kwengalo.
Ukukhanya kokujonga kwicala lempembelelo
Siphikise ukuba imifuziselo ephambili ye-intra PFC, kunye ne-PFC-amygdala intsebenzo echaza ulawulo olungqongqo lwe-unidirectional top-Down kwiinkqubo zemvakalelo azingqinelani neempawu ze-laminar zonxibelelwano phakathi kwale mimandla. Impikiswano yethu malunga nale modeli iphezulu ye-PFC-amygdala kunye ne-intra-PFC ixhomekeke kakhulu kwimodeli eyakhiweyo echazwe nguBarbas kunye noogxa, apho ipatheni ye-laminar yoqikelelo iyalela ukuba iiprojekthi zibonisa impendulo-njengokukhetha uchuku, okanye ukuhambisa ulwazi Ukuba kuchanekile, iindawo ezinxulumene nemvakalelo zibonakala zibonelela ngolawulo oluphezulu olukumgangatho ophantsi wokuhambisa ulwazi kude neendawo ezaziwa njenge-PFC ngokwesiko.
Sikholelwa ekubeni isigama somgaqo ongaphezulu-ezantsi kukhokelele kwingcamango engqiqweni yokuqonda ubudlelwane phakathi kwemimandla yobuchopho kunye nenkqubo yemvakalelo. Oku kukhetha icala kuhambelana nefilosofi yeendima zeenkqubo "zokuqonda" kunye "neemvakalelo" ezibeka ukuqonda ngaphezulu kweemvakalelo zesilwanyana. Kodwa oku kuthatha icala kungaphazamisana nokukwazi kwethu ukufumana ulwazi olupheleleyo lwendlela ingqondo eqhuba ngayo ulwazi. Ukuba iinkqubo zemvakalelo zilawula kunye nokusebenza ngoku “kuthathelwa ingqalelo”, ngokungaphaya okanye ngaphezulu kwayo yonke enye indlela ejikelezayo, isigama esingaphezulu nesezantsi sinokuba sesingafanelekanga xa kuthathelwa ingqalelo unxibelelwano lokuqonda.
Izithintelo ekungeniseni umsebenzi kwisakhiwo
Ukugqwesa kwimodeli eyakhiweyo kukuba kukhokelela kuqikelelo olomeleleyo malunga nonxibelelwano lwengingqi. Nangona kunjalo, ukugxeka okuninzi kunokuphakanyiswa kwangoko ekuzoba umqukumbelo wokusebenza ngokusekwe kwiimpawu ze-anatomical. Okokuqala, nangona imodeli yolwakhiwo ixhaswe ngokuqinileyo ngokwembono yayo yeepatheni zonxibelelaniso ze-laminar esekwe kwisiseko se-cytoar, ulwaziso ngokubhekisele ekusebenzeni kwezi mpembelelo zeendlela zokuxhuma ze-laminar alufumananga uvavanyo olusesikweni kwiziphaluka ngaphandle kwemijelo yokuqhubekeka kokusebenza. Nangona kubonakala kusengqiqweni ukucinga ukuba izinto ezifanayo ezisebenzayo zibonisa iipateni ze-laminar zokuqikelela kuyo yonke ingqondo, oku akuyonyani. Ngenxa yoko, ugculelo malunga nepropathi yokusebenza konxibelelwano kwiPFC lusebenza kuphela ukuba iimpawu zomsebenzi zentsingiselo kunye nengqikelelo yengxelo ziye zangqinwa zibambelele kwii-cortices zombutho.
Sithumele unxibelelwano oluluqilima phakathi kwempendulo yokusebenza kunye nommiselo we-top-down, kunye nekhonkco elifanayo elomeleleyo phakathi kweenkqubo zokutya kunye nezantsi. Ingxelo yengxelo kunye ne-feedirect ivela kwi-theory yolawulo, ethi izame ukuchaza ukusebenza kweenkqubo eziguqukayo. Ukuphunyezwa kwala magama ngoososayensi beengqondo kunye nochwephesha bezengqondo akothusi njengoko ingcinga yeendlela zokunikwa kwengxelo inikezela ngolawulo lolawulo kunye neendlela ezilandelanayo ezinikezela ukuhanjiswa kolwazi kwiindawo eziphakamileyo kumjelo wokulungisa kuyinto enomdla. Nangona kunjalo, i-equation elula yolawulo oluphezulu olusezantsi kunye nengxelo kunye nezantsi-phezulu kunye ne-feedxpo kuyingxaki kude kufikelele kwinqanaba lokuba amanqaku ongezelelweyo aboniswe phezulu kunye naphantsi kwe-conceptualizations. Ezo zinto zongezelelweyo azifane zichazwe ngokucacileyo, kodwa zinokungqina ubucaka ekucingeni kolwazi ekuqhubekeni kolwazi. Siyarhanela ukuba ezinye iingcali zisebenzisa amabinzana phezulu-ezantsi nezantsi-iindlela ezingangqinelaniyo nengxelo kunye neendlela ezichazayo njengokuchazwa kwthiyori yolawulo, kodwa ukungangqinelani kunqabile ukucaciswa kuncwadi.
Ekubonakaliseni ingxelo kunye noqikelelo lwePFC, siqaphela ukuba asithethi ukuba zonke iingqikelelo zohlobo olufanayo. Iindawo ezinendibaniselwano yengxelo, eyondlayo, kunye nokunxibelelana okucwangcisiweyo, kodwa ulwalamano lolu nxibelelwano luhluka ngokungafaniyo kwiindawo zonke. Ngale ndlela siphawula iipateni eziphambili zonxibelelwano, kodwa oku akuthethi ukuba uxhulumaniso oluseleyo alubalulekanga ngokusebenzayo. Umzekelo, imimandla ye-PFC eulaminate ngokuqinisekileyo ineengxelo ezaneleyo zokwenza ingxelo zokunceda ukulawula imimandla engacacanga granular, nokuba le ayisiyiyo indlela ebalaseleyo yonxibelelwano phakathi kweendawo.
Ngaphaya koko, udibaniso loqikelelo lweendlela ezinokutya lunokuthi kwezinye iimeko lutshintshe ukuqhubekeka kwemimandla ekujoliswe kuyo kunokuhambisa nje ulwazi. Mhlawumbi ngowona mzekelo mhle olu hlobo lwendlela yokuphinda uguqulwe kwiimodeli zokhuphiswano oluhlanganisiweyo (I-Desimone kunye ne-Duncan 1995; UDuncan et al. 1997) apho ukufunyanwa kommeli omnye kukhokelela ekucinezelweni komnye. Kwiimodeli ezinjalo, ukuhambisa phambili ngophawu olunikezelweyo kungakhokelela ekuphuculeni kokusebenza kwento enyusayo, kunye nokucinezelana okuthelekeneyo kolunye ukhuthazo (I-Desimone kunye ne-Duncan 1995). Ngale ndlela, into eyondlayo iya phambili isebenze ukumodareyitha ukuqhubekeka kwemimandla ekujoliswe kuyo. Kwimeko yokusebenza kwe-PFC, umqondiso we-DLPFC unakho ukutshintsha ukhuphiswano phakathi kwabameli abakhoyo kwi-OFC kolu hlobo lwendlela yokuqikelela. Olu hlobo lweendlela zokhuphiswano luyathakazelisa kuba luya kuthetha ukucaciswa kweempawu ezithile (UWalther kunye noKoch, 2006), ezingakhange zibandakanywe kwiimodeli zokulawulwa kweemvakalelo.
Ekuqwalaseleni imodeli yolwakhiwo, kubalulekile ukuphinda ubalule ukuba iikhrayitheriya ezisetyenziswa nguBarbas kunye noogxa bakhe ukuchaza okuphambili kunye nokunxibelelana kwengxelo akuhambelani ngokupheleleyo nenqobo esetyenzisiweyo abanye abaphandi behlola ulungiselelo lwe-hierarchical ye-laminar. Ngokukodwa, iinkcazo zempendulo kunye nokunxibelelana ngaphambili zihlala zichazwa ngokubhekisele kumaleko IV, njengokuthi ukuphakanyiswa (kukhuphuka) uqikelelo kuchazwa ngokupheliswa kwabo kumqolo IV (okanye ubukhulu becala kumqolo IV), ngelixa ingxelo (yehla) uqikelelo luphela ngaphandle komaleko. IV. Ngelixa ukubambelela ngokungqongqo kumaleko we-IV kusenokwenzeka ukuba akucetyiswa kakuhle, njengoko kungakhethwanga kule patheni.IPandya kunye neRockland, 1979; UFelleman noVan Essen, 1991), iimpembelelo zendlela ebanzi yokuvumela uqikelelo ekuphelisweni kolwazelelelo lwe-infragranular V kunye ne-VI ukuba luphathwe njengotyekelo lokutya ayiqondwa ncam. Ngokucacileyo, izifundo zexesha lokudubula kwiindawo ezahlukeneyo ze-cCical cortical zinokuphendula lo mbuzo, kodwa idatha kulo mbandela iyasilela.
Umbuzo weenqobo ubangela ukuba ume kancinci ngaphambi kokuthatha isigqibo sokuba i-OFC-DLPFC inepateni apho kufanele ukuba i-OFC iphathwe njengomgangatho ophezulu kune-DLPFC, kwaye ayizonjongo zethu ukuphikisana. Nangona kunjalo, kunokuchazwa ngokucacileyo ukuba iipateni zentelekelelo ngokuqinisekileyo azihambelani nombutho wobuphathi obuphezulu apho i-DLPFC ikwindawo ekumgangatho wobukhosi ngaphezulu kwe-OFC, ngendlela efanayo neendawo eziphakamileyo zenqanaba lokuhlala ngaphezulu kweendawo eziphambili zoluvo. Kananjalo, imifuziselo yombutho we-PFC kuya kuba bubulumko ukuphepha indawo egcweleyo ye-DLPFC njengokuhlala kwindawo ephezulu yemimandla ye-PFC.
Imodeli yokudityaniswa okuphindaphindiweyo kunye nengxelo
Ukuvavanywa kweemodeli ezikhoyo zokunxibelelana nemvakalelo akukho mfuneko yokuba izifundo ezimbalwa ezipapashiweyo ukuza kuthi ga ngoku zibandakanye kuvavanyo oluthile lokuba iingqikelelo zibonisa ingxelo, iingxelo zolohlobo okanye iingqikelelo zangaphambili (ngaphandle kokubalaseleyo Seminowicz et al. 2004). Uninzi lwezifundo ezinomdla ze-neuroimaging zezi zifundo aziboneleli ngolwazi oluthile lwe-laminar olunokuphendula lo mbandela. Nangona kunjalo, uphuhliso lwamva nje kubuchule bokumanyanisa unxibelelwano olusebenzayo lubonelela ngezixhobo ezinokusetyenziselwa ukuxelisa ubunjani kunye nokuya kunxibelelwano phakathi kwemimandla. Umzekelo, umfuziselo oshukumayo we-causal (DCM) usebenzisa ukuthambeka kwinqanaba losapho kunye ne-Bayesian imodeli yokulinganisa inokufakwa kuvavanya ii-hypotheses ngokubhekisele kulwazelelo kunye nobunjani bokuhamba kolwazi kunye noshintsho lwembangela yemimandla eyahlukeneyo yobuchopho (UFriston et al. 2003; U-Chen et al. 2009; UDaunizeau et al. I-2009; UFriston kunye neDolan 2010; UPenny et al. 2010). I-DCM inokuvavanya kwakhona imodeli yokhuphiswano njengokubonelela ngentloko ukuya kuthelekiso lwentloko ukuba ingaba i-DLPFC isezantsi-ilawula i-amygdala ngokuthe ngqo okanye ngesakhiwo esiliqela. Ukuza kuthi ga ngoku, zimbalwa kuphela izifundo zeDCM ezinxulumene nokusebenza ngokweemvakalelo ezipapashiweyo (Ethofer et al. 2006; USmith et al. 2006; Rowe et al. 2008; UAlmeida et al. 2009), kwaye kulwazi lwethu akukho zifundo zipapashiweyo zijongana ngqo nolawulo lweemvakalelo. Nangona kunjalo, ukusetyenziswa kwezobuchwephesha kunokwenzeka ukuba kuphucule kakhulu ukuqonda kwethu kokunxibelelana kweemvakalelo kwiminyaka ezayo.
Ukuvavanywa ngokuthe ngqo kwempembelelo
Mhlawumbi yeyona ndlela ilungileyo yokuseka ubudlelwane obusebenzayo phakathi kwemimandla yobuchopho kukuhlolwa kwommandla omnye ngexesha lokukhetha komzimba phezulu okanye ezantsi kommiselo wenye indawo. Umzekelo, ukuba i-DLPFC isebenza ngokunyibilikisa ukusebenza kwe-OFC, umntu unokulindela iimpendulo ezongeziweyo kwi-OFC xa i-DLPFC isenziwa ngaphandle kweintanethi. Oku kunokuba nakho ukuqwalaselwa ngokuvavanya imisebenzi ye-OFC nge-fMRI kwizigulana ezinezilonda ze-DLPFC. Ngenye indlela, ukunyanzeliswa kwamandla kazibuthe (iTMS) kunokusetyenziswa ngaphezulu kwe-DLPFC ukutshintsha okwethutyana impembelelo ye-DLPFC kwimisebenzi ye-OFC. Kanjalo, Knoch et al. (2006) kutsha nje kuxelwe ukuba i-TMS engaphezulu kwe-DLPFC ivelise utshintsho kwimisebenzi yasemva kwe-OFC ngendlela exhomekeke rhoqo. Ngokufanayo, kuya kuba ngumdla ukwazi ukuba izilonda kwenye indawo yecortex yangaphambili ziyichaphazela njani inkqubo kwezinye iindawo zenethiwekhi. Umzekelo, ukuba i-OFC ibalulekile ekubaleni ixabiso lomvuzo eliqinisekileyo, kwenzeka ntoni kwiindawo ezingakumbi xa isuswa i-OFC? Saddoris et al. (2005) basebenzise olu hlobo lwendlela yokujonga ukuba izilonda ze-OFC zitshintsha njani ngokudubula kwi-amygdalar kudubula ezintongeni, kodwa ezinye izifundo ezithatha le ndlela zinqabile ukuba zingabikho. Uncwadi olukhulayo kunxibelelwano olusebenzayo ngokufanayo lunokwandisa ukuqonda kwethu ukuba le mimandla yobuchopho inxibelelana njani. Nangona kunjalo, ukuqonda okugcweleyo kokunxibelelana kuya kufikelelwa kuphela ngononophelo kwiinkalo ezithile zesekethe.
Amagqabantshintshi ophando
- Ukudibana okungxamisekileyo okuphambili kulawula ukuguqulelwa kweemvakalelo kwi-amygdala
- Iipatheni zokuqikelela kweLaminar zimisela ukuhamba kolwazi kwi-cortex yangaphambili
- Uqikelelo lwempendulo kunye nengxelo ngomngeni wangaphambili
Imibulelo
Lo msebenzi uxhaswe zizibonelelo T32MH018931-21, T32MH018921-20, & 5R01MH074567-04 kwiZiko leSizwe lezeMpilo yeNgqondo. Siyabulela uTawny Richardson ngoncedo ekulungiseleleni lo mbhalo-ngqangi.
Imihlathi
Iphepha elichazayo ukuba awusenanto oyifunayo: Le fayili yeFayile yombhalo wesandla ongabhalwanga owamkelwe ukushicilelwa. Njengenkonzo kumakhasimende ethu sinika le ngcaciso yokuqala kwincwadi yesandla. Umbhalo wesandla uza kufumana ukukopishwa, ukufakela, nokuphonononga ubungqina obunokubakho ngaphambi kokuba kukhutshwe kwifomu yayo yokugqibela. Nceda uqaphele ukuba ngexesha lokuveliswa kweeprogram ezinokuthi zifumaneke ezinokuthi ziphazamise umxholo, kunye nazo zonke izisemthethweni ezichasayo ezisetyenziswa kwiphephancwadi.
Uluhlu lweNkcukacha
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