Neurosci Biobehav Rev. 2012 Januwale; 36 (1) : 479–501. Yofalitsidwa pa intaneti 2011 Ogasiti 25. doi: 10.1016/j.neubiorev.2011.08.005
Rebecca Ray 1 ndi David H. Zald 2
Nkhani yomaliza yosinthidwa ndi wofalitsa ikupezeka pa Neurosci Biobehav Rev
Onani nkhani zina mu PMC zomwe zikufotokoza nkhani yofalitsidwayi.
Kudalirika
Kafukufuku wazamatsenga akuwonetsa kuti njira zamkati zimagwirizanirana ndi zochitika zina zazidziwitso. Kafukufuku wawonetsa kuthekera konse kwa zochitika zam'maganizo kuti zikope zochitika zambiri zam'malingaliro, komanso kuthekera kwa anthu kugwiritsa ntchito njira zowongolera zazidziwitso zowongolera zakumvera. Zigawo za preortal cortex zimawoneka kuti zimathandizira pakuchita uku. Komabe, momwe amathandizirana izi zimangowonjezereka pang'ono. Pakubwereza komweku tafotokozera kulumikizana kwakumakhala pakati pa malo am'kati mwa malo am'kati mwa dorsal komanso dorsal komanso momwe amalumikizirana ndi zigawo zamatsenga. Malo ochepa okha omwe angathe kutsogolera madongosolo amygdalar, ndipo machitidwe otsogola kwambiri a malingaliro ndi zitsanzo za malingaliro amomwe akuyenera kukakamizidwa kuti akhale njira zowoneka bwino. Tikuwunikanso momwe njira yolumikizirana yodutsirana ndi mayankho pakati pa zigawo izi ingatithandizire kudziwa kuchuluka kwa chidziwitso pakati pa madera a pakati ndi a dorsal asanafike ndi amygdala. Njira izi zolumikizirana sizikugwirizana ndi malingaliro angapo ofotokozedwa pokhudzana ndi chikhalidwe cholumikizirana pakati pa kutengeka ndi kuzindikira.
1. Introduction
Kafukufuku wosonkhanitsa amafufuza momwe malingaliro amagwirizanirana ndi mbali zina za kuzindikira. Ntchito yotereyi yafotokoza njira zomwe chidziwitso chogwirizana ndi malingaliro chingatsogolere kapena kukondera chidwi ( Ohman et al. 2001 ; Most et al. 2005 ; Mathews & Wells, 1999 ), ndikukhudza njira zochitira zisankho ( Knutson et al. 2008 ). Nthawi yomweyo, mabuku omwe akukula akuwonetsa kuti njira zoganizira monga kubwerezabwereza zimatha kulamulira momwe munthu amayankhira pamalingaliro ( Jackson et al. 2000 ; Kim & Hamann, 2007 ; Ochsner et al. 2002 ; Ochsner, Ray, et al. 2004 ; Ray, Wilhelm & Gross, 2008 ). Zoonadi, kuyanjana pakati pa ntchito zomwe nthawi zambiri zimafotokozedwa ngati zamaganizo kapena zamaganizo ndizokwanira kukayikira kusiyana komwe kumachitika pakati pa magawo awa (onani mwachitsanzo Pessoa, 2008 ). Komabe, kusiyana kumeneku kumapereka phindu la lingaliro chifukwa chakuti kusintha kwa malingaliro kumakhala ndi makhalidwe enieni a ntchito omwe amatha kusiyanitsidwa ndi magawo ena a chidziwitso mwanjira yomwe njira zosamalira kapena kukumbukira zimakhala ndi makhalidwe osiyanasiyana ndipo zimakhazikika m'maukonde osiyanasiyana (ngakhale nthawi zina zimalumikizana pang'ono) a madera a ubongo.
Momwe malingaliro ndi magawo ena amalingaliro amagwirira ntchito akhala ofunikira kwambiri pa zitsanzo za matenda amisala. Mwachitsanzo, malingaliro a matenda a nkhawa nthawi zambiri amayang'ana kwambiri kukonda kwambiri zinthu zowopseza ( Bishop, 2007 ; Cisler & Koster, 2010 ; Ouimet, Gawronski & Dozois, 2009 ; Williams et al. 1996 ). Mofananamo, kulephera kugwiritsa ntchito mphamvu zolamulira malingaliro kuchokera pansi kumawonedwa kwambiri ngati maziko a matenda amisala kuyambira kuvutika maganizo kwakukulu ( Fales et al. 2008 ; Johnstone et al. 2007 ; Almeida et al. 2009 ; Taylor Tavares et al. 2008 ), mpaka ku vuto la umunthu wa m'malire ( New et al. 2008 ).
Madera a Prefrontal amaonekera bwino kwambiri m'machitidwe a neurobiological a kulumikizana pakati pa malingaliro ndi mbali zina za kuzindikira. Komabe, mawonekedwe a anatomical a madera osiyanasiyana a prefrontal nthawi zambiri amangoyang'aniridwa pang'ono poganizira kuvomerezeka kwa mitundu yotereyi. Malinga ndi momwe anatomy imaganiziridwira, nthawi zambiri imakambidwa mwachidule ngati derali lili ndi kulumikizana mwachindunji kapena kwa efferent ndi madera a limbic, monga amygdala kapena hypothalamus. Komabe, tsatanetsatane wa kulumikizana kumeneku ndikofunikira kuti timvetsetse kuyanjana kwa maderawa. Mwachitsanzo, chitsanzo chomwe chimasonyeza kuti dorsolateral prefrontal cortex (DLPFC) imaletsa mwachindunji ntchito ya amygdalar chingakhale chomveka ngati zikuwonetsedwa kuti DLPFC imatumiza ziwonetsero zokwanira mwachindunji ku amygdalar. Ngati ziwonetsero zoterezo ndi zochepa kapena palibe, mitundu ina yomwe imadalira madera apakati idzakhala yofunikira kufotokoza mphamvu ya DLPFC pamayankho a amygdalar.
Ma kapangidwe kazigawo zosiyanasiyana zam'malo oyamba komanso mawonekedwe a laminar omwe amatha kulumikizana amathanso kupereka chidziwitso pakukhudzana pakati pamalingaliro ndi kuzindikira njira yolumikizidwa ndi preortal cortex (PFC). Makamaka, ma cytoaramangidwe a magawo osiyanasiyana a cortical amatsimikizira momwe amapangira chidziwitso ndi kuyanjana ndi zigawo zina. Kusanthula kwachiwiri kumeneku sikunalowe mu zokambirana za magwiridwe antchito a chidwi, ngakhale zili ndi tanthauzo lambiri pomvetsetsa njirazi.
Mu pepala lino, tikuyesera kufotokoza zinthu zingapo zokhudza kulumikizana pakati pa madera osiyanasiyana a PFC, komanso momwe zimagwirira ntchito ndi amygdala. Timayang'ana kwambiri kusiyana pakati pa orbital ndi dorsolateral PFC chifukwa cha mgwirizano wa nthawi yayitali wa orbitofrontal cortex (OFC) ndi machitidwe amalingaliro ( Zald & Kim, 1996 ) komanso mgwirizano womwewo wa nthawi yayitali wa DLPFC ndi mbali zazikulu za chidziwitso ( Fuster, 1989 ; Stuss & Benson, 1986 ). Timafotokozanso udindo wa kapangidwe ka anterior cingulate (ACC)/medial frontal muzochita izi, chifukwa kuchuluka kwa deta kukuwonetsa kuti mapangidwe awa amapereka mawonekedwe ofunikira pakati pa malingaliro ndi mbali zina za chidziwitso.
2. Zojambula pamwambapa ndi zomangamanga za PFC
Zojambula pamanja
PFC nthawi zambiri imagawidwa m'magawo 6 akuluakulu, dorsolateral, ventrolateral (VLPFC), frontopolar (FP), OFC, ventromedial (VMPFC), ndi dorsomedial (DMPFC) (onani Chithunzi 1 ). Malire enieni a malo awa amagwiritsidwa ntchito mosiyanasiyana ndi ofufuza, koma mayina onse atsimikizira kukhala othandiza ngati dongosolo lalikulu lokonzekera kumvetsetsa kapangidwe ka thupi ndi ntchito ya PFC.
Phylogeny ndi Cytoaramangidwe
PFC ili ndi njira ziwiri zosiyana, zosiyana ndi za phylogenetically architectonic ( Barbas, 1988 ; Sanides, 1969 ; Yeterian & Pandya, 1991 ). Njira ya basoventral imachokera ku fungo (allocortical) core kudzera mu OFC ndikufalikira kutsogolo kupita ku ventral frontal pole, komanso mbali ina kupita ku VLPFC (kumapeto kwa Brodmann area (BA) V46). Mosiyana ndi zimenezi, njira ya mediodorsal imayamba pa corpus collosum, imadutsa pakhoma lapakati la frontal lobe kenako imazungulira m'mphepete mwa lobe kupita ku DLPFC (kumapeto kwa BA D46). Njira iliyonseyi ikuwonetsa njira yotsatizana ya kapangidwe ka cortical yomwe ikuwonetsedwa pakukula ndi kufalikira kwa granular layer IV. Gawo lakale kwambiri la njirazi ndi agranular mwachilengedwe, pomwe madera achichepere kwambiri ali ndi granular layer yolimba komanso yodziwika bwino. Mu njira ya basoventral, kupita patsogolo kwa cortical kumeneku kumayambira mu posterior OFC (insula ya agranular pogwiritsa ntchito mawu a Carmichael ndi Price ( Carmichael & Price, 1994 )) kutsatiridwa ndi dysgranular (granular yofooka) cortex m'madera apakati a OFC, kupita ku eulaminate I cortex yokhala ndi granular layer IV yodziwika bwino pamene munthu akuyenda kutsogolo kapena kumbali, ndipo pamapeto pake kufika ku eulaminate II cortex yokhala ndi thin layer IV ndi strong supragranular layers pamene munthu akupita ku frontal pole ndi ventrolateral regions ( Barbas & Pandya, 1989 ; Carmichael & Price, 1994 ; Petrides & Mackey, 2006 ; Price, 2006a ). Njira ya mediumdorsal ikuwonetsa kupita patsogolo kofanana kwa cytoarchitectural. Kuyambira ndi periallocortex cortex m'mbali mwa rostral corpus collosum, chizolowezichi chimakhala chosalimba mu cingulate (kuphatikiza madera a subgenual, pregenual, ndi supragenual), eulaminate I pamene munthu akuyenda kutsogolo m'mbali mwa khoma la medial kapena superiorly kulowa mu superior frontal gyrus, ndipo pamapeto pake chimakhala elumaniate II m'madera a dorsolateral (BA 8 ndi 46).
Pofuna kupewa chisokonezo, tikuwona kuti kugwiritsa ntchito mawu akuti mediodorsal trend sikuyenera kusokonezedwa ndi dera la DMPFC lomwe lafotokozedwa mu Chithunzi 1. Mediodorsal trend ikuphatikizapo DMPFC, komanso ikuphatikizapo madera a VMPFC 25 ndi 32, ndi magawo a BA 10 m'mbali mwa khoma lapakati (dera la 10m mu dzina la Ongur et al. (2003) ; Chithunzi 2 ).
Kapangidwe ka chitukuko cha cytoarchitectural pamene munthu akuyenda kuchokera ku agranular kupita ku eulaminate II cortex kamakhala ndi kuwonjezeka kwa chiwerengero chonse cha ma neuron (kuchuluka kwa maselo), kukula kwa maselo a pyramidal mu zigawo II ndi V, ndi mulingo wa myelination ( Barbas & Pandya, 1989 ; Dombrowski et al. 2001 ; Chithunzi 3 ), zomwe pamodzi zimapangitsa kuti pakhale makhalidwe osiyanasiyana ogwiritsira ntchito chidziwitso m'madera osiyanasiyana. Kusiyana kwina kwakukulu pakati pa madera a prefrontal kumachitika pankhani ya histological staining, nthawi zambiri kuwonetsa mawonekedwe osiyanasiyana a interneuron. Carmichael ndi Price ( Carmichael & Price, 1994 ) amagawa Macaque OFC ndi medial PFC m'magawo angapo kutengera mawonekedwe otere (onani chithunzi 4 ), ndipo zambiri mwa izi zitha kuzindikirika mwa anthu ( Ongur et al., 2003 ). Mawonekedwe osiyanasiyana a interneuron omwe amawonedwa m'madera ang'onoang'ono a prefrontal amakhudza makhalidwe enieni a kukonza chidziwitso omwe amachitidwa ndi madera ang'onoang'ono a prefrontal ( Wang et al., 2004 ; Zald, 2007 ), koma ndi opitilira muyeso wa pepalali. Mwachidule, magawano ofotokozedwa bwino a PFC ali ndi njira zosiyanasiyana zolumikizirana mkati mwa PFC komanso m'madera ena a ubongo a cortical ndi subcortical.
Mapangidwe amtundu wa Anthu
Ngakhale pali kufanana kwakukulu mu kapangidwe ka ma primate ndi ma cytoarchitecture a anthu mkati mwa ma lobes akutsogolo, ndipo machitidwe ambiri a phylogenetic amagawidwa m'mitundu yonse ya primate, zovuta zingapo zimachitika posuntha pakati pa deta ya anthu ndi nyama. Choyamba, maphunziro a neuroimaging a anthu nthawi zambiri amatchula madera a Brodmann ( Brodmann, 1914 ), koma sakuwonetsa kupita patsogolo pakuzindikira madera a cytoarchitectural ndi malire a area omwe achitika kuyambira ntchito yoyamba ya Brodmann pafupifupi zaka zana zapitazo. Chachiwiri, kugwiritsa ntchito zilembo za madera awa nthawi zambiri kumachokera pa Talairach atlas ( Talairach & Tournoux, 1988 ), koma atlas iyi ndi pafupifupi, popeza kusanthula kwa cytoarchitectural sikunachitike pa ubongo womwe umapanga maziko a atlas. Chachitatu, pali kusagwirizana pakati pa zilembo za nyama ndi zilembo za anthu mu ventral frontal lobe, chifukwa deta ya nyama imagwiritsa ntchito mitundu yosiyanasiyana ya zilembo zomwe Walker adapanga ( Walker, 1940 ), zomwe olemba ena apereka kwa anthu ( Petrides & Mackey, 2006 ; Ongur et al., 2003 ), pomwe ofufuza ambiri a neuroimaging amagwiritsabe ntchito zilembo za Brodmann. Tsoka ilo, nthawi zina sizikudziwika bwino kuti ndi ofufuza a neuroimaging a system yolemba zilembo ati akamanena zomwe apeza. Izi zimapangitsa kusamveka bwino mu lateral OFC/VLPFC, komwe ofufuza a anthu nthawi zambiri amatchula BA 47, koma mabuku a nyama amatchula dera 12. Chizindikiro 47/12 tsopano chatengedwa ndi akatswiri ena a neuroanatomics kuti afotokoze derali mwa anthu, ngakhale malire apakati a dera lino akutsutsidwabe ndi akatswiri otsogola a neuroanatomics ( Petrides & Mackey, 2006 ; Ongur et al., 2003 ). Mofananamo, madera 13 ndi 14 ali ndi malire omveka bwino mwa anyani, ndipo madera ofanana amapezeka mwa anthu, koma Brodmann kapena Talairach, omwe adagwiritsa ntchito chizindikiro cha dera 11 kumadera onse akumbuyo ndi akutsogolo a OFC yapakati. Pofotokoza deta ya neuroimaging ya anthu, nthawi zambiri timatchula njira yayikulu yolembera yomwe yafotokozedwa ndi Petrides ndi Mackey (2006) , m'malo mwa Talairach atlas kuti tigwiritse ntchito deta yochokera ku maphunziro osakhala a anthu.
3. Maulalo
Zambiri zomwe zilipo pa kulumikizana kwa prefrontal zimachokera ku maphunziro a nyama. Komabe, poganizira za cytoarchitectural homology pakati pa anyani ( Petrides & Mackey, 2006 ; Ongur et al., 2003 ), nthawi zambiri zimaganiziridwa kuti kulumikizana kwa madera awa kumasungidwa kwambiri pakati pa mitundu ya anyani. Chifukwa chake, ndikoyenera kugwiritsa ntchito mabuku a anyani osakhala anthu pa kulumikizana ngati maziko owunikira kulumikizana mwa anthu. Tikuyang'ana kwambiri mitundu iwiri ya kulumikizana apa: kulumikizana kwa amygdala-PFC, ndi kulumikizana pakati pa madera osiyanasiyana a PFC.
Kuyika kwa Amygdalar ku PFC
OFC ndi medial PFC zimalandira zambiri kuchokera ku amygdala ( Amaral et al., 1992 ; Carmichael & Price, 1995 ; Barbas & Zikopoulos, 2006 ). Izi zikusiyana kwambiri ndi DLPFC, yomwe imalandira zochepa zoyerekeza kuchokera ku amygdala. Kuwunikanso mabuku kukuwonetsa kuti zina mwa zoyerekeza za PFC ya m'mimba ndi yamkati zimasiyana kutengera nucleus kapena subnucleus yochokera ( Amaral & Price, 1984 ; Barbas & De, 1990 ; Amaral et al., 1992 ; Carmichael & Price, 1995 ). Komabe, tsatanetsatane uwu ndi woposa zomwe zili mu pepalali, ndipo chithunzi chokwanira cha kulumikizana chimaonekera m'ma nuclei osiyanasiyana kuti chidziwitse kukambirana kwakukulu kwa njira zolumikizirana. Chithunzi 5 chikuwonetsa chithunzithunzi cha amygdalar projections (zochokera ku ma nuclei angapo a amygdalar) kupita pamwamba pa ubongo wa macaque ndi medial ndi Carmichael ndi Price nomenclature. Chithunzichi chikuwonetsa kuti pamwamba pa orbital sipali mgwirizano wofanana ndi amygdala. Chofunika kwambiri ndichakuti palibe malo okwanira olowera m'malo 13m, 13l, 12m, 11l, ndi 10o pamwamba pa orbital. Khoma lapakati limalandiranso malo okwanira olowera m'malo a amygdalar, koma kachiwiri silili lofanana, chifukwa dera la 10m kapena dera la 9 sililandira malo okwanira olowera m'malo a amygdalar.
Malingaliro awiri atha kuchokera pamalingaliro awa. Choyamba, momwe amygalar athandizira ku PFC ndizokhazikika pamangidwe ndipo zimakhazikitsidwa m'malo omwe ali ndi zomangamanga pang'ono. Izi zikuwonetsa kuti kungakhale kulakwitsa kuchitira onse a OFC kapena a PFC amisala ngati kuti alumikizidwa kwambiri ndi amygdala. M'malo mwake, chidwi ndi malo omwe ali mkati mwa OFC ndi PFC yamankhwala amalangizidwa musanalowe m'malingaliro okhudzana ndi amygdalar. Chachiwiri, DLPFC ndi FP zimalandila kulowerera kotsika kwambiri kwa amygdalar (kwenikweni njira zowoneka bwino kwambiri ndizomwe zimawonetsa umboni wa kulowerera kwa amygdalar). Zotsatira zake, zochitika za amygdalar pa DLPFC ndi FP pokonza zikuwoneka kuti sizikuyenda mwachindunji, mwina kudzera mu ma cingate kapena posterior OFC (kapena kudzera m'njira zina zowonjezera, monga kusinthira kwa ma system a neurotransmitter).
Kutulutsa koyambirira kupita ku amygdala
Zotsatira za PFC ku amygdala nazonso ndi zachigawo ( Price, 2006b ; Ghashghaei et al., 2007 ; Stefanacci & Amaral, 2002 ; Stefanacci & Amaral, 2000 ). Kawirikawiri, madera oyambilira omwe amalandira ma projection kuchokera ku amygdala amatumiza ma projection kubwerera ku amygdala, pomwe madera omwe salandira ma amygdalar ambiri (monga DLPFC ndi FP) ali ndi ma projection ofooka kwambiri ku amygdala. Kuchuluka kwa ma projection kumawonetsa kwambiri cytoarchitectonics, ndi kufooka kwa ma projection density pamene munthu akuyenda kuchokera kumadera a agranular kupita ku eulaminate isocortex yopangidwa bwino kwambiri. Kachitidwe aka kakusonyeza kuti madera a isocortical (DLPFC ndi FP) sangapereke mphamvu yolunjika pa amygdala, ndipo momwe amakhudzira amygdala, mphamvuyo ikhoza kukhala yosalunjika. Izi sizikutanthauza kuti palibe ma specifications ochokera ku DLPFC kupita ku amygdala, chifukwa kafukufuku wambiri wawona ma specifications ochokera ku dera 9 ndi 46 ( Stefanacci & Amaral, 2002 ; Aggleton et al., 1980 ; Amaral & Insausti, 1992 ). M'malo mwake, ma specifications nthawi zambiri amakhala opepuka kwambiri kuti apereke malamulo ambiri okhudza amygdalar processing.
Ngakhale kuti ma cytoarchitectonics ambiri amapereka njira yolimba yokonzekera zinthu mogwirizana ndi amygdala-prefrontal, kugawa kwa zigawo ndi zotuluka m'madera osiyanasiyana sikugwirizana ( Ghashghaei et al., 2007 ). Chodziwika bwino n'chakuti, kulowetsa kwakukulu kwa amygdalar mu PFC kuli m'dera la agranular insular motsatira posterior OFC, pomwe, kutulutsa kwakukulu kwa amygdala kumachokera ku posterior subgenual cingulate region (BA 25) ndi magawo a dorsal anterior cingulate (BA 24). Mwachidule, madera a medial wall ali ndi kutulutsa kwakukulu kuposa kulowetsa kuchokera ku amygdala, pomwe madera a posterior OFC ali ndi kulowetsa kwakukulu kuposa kutulutsa. Chochititsa chidwi n'chakuti, madera a lateral PFC omwe ali ndi kulumikizana pang'ono, madera a DLPFC (BA 8, 9 ndi dorsal 46) ali ndi kulowetsa kwakukulu kuchokera ku kutulutsa kupita ku amygdala, pomwe mawonekedwewo amasinthidwa mkati mwa VLPFC. Chodziwika bwino pankhaniyi, chigawo chakumbuyo cha dera la 12l mkati mwa VLPFC chimapereka ziwonetsero zolimbitsa thupi ku amygdala, zomwe zimapangitsa kuti ikhale dera lokhalo la lateral PFC lomwe lili ndi kulowetsa kwakukulu mwachindunji ku amygdala. Zoonadi, ma projekiti awa ndi amphamvu kuposa omwe amaoneka m'madera a anterior orbital, omwe ali ndi madera okwana 12l omwe amatuluka kwambiri kuposa omwe amalowa, koma nthawi zambiri amawonetsa milingo yofooka yolumikizirana kuposa dera la 12l.
Mitundu yosiyanasiyana ya amygdala subnuclei imalandira PFC input. Basal ndi accessory basal ndi medial nuclei imalandira densely projections, komanso kulandira projections kuchokera ku madera ambiri a PFC, pomwe lateral, central ndi cortical nuclei imalandira PFC projections, koma pamlingo wochepa komanso wofalikira ( Stefanacci & Amaral, 2002 ). BA 25 ndi yodziwika bwino chifukwa sikuti imatumiza densely projections ku amygdala, komanso imatumiza projections ku generalized nuclei array, monga amygdala subnucleus iliyonse yomwe yatchulidwa pamwambapa imalandira input kuchokera ku BA 25. Ngakhale ikuwoneka ngati kuwala mu column B ya Chithunzi 6 , ndikofunikira kudziwa kuti BA 32 imapereka projections yolongosoledwa bwino ku amygdala. M'njira zambiri BA32 imawoneka yofanana ndi prelimbic cortex m'makoswe ( Price, 2006a ). Mu makoswe, prelimbic cortex imapita ku magawo a basolateral ndi central nucleus ya amygdala ( Vertes, 2004 ). Mu anyani omwe si anthu, ma projection awonedwanso kuchokera ku BA 32 kupita ku gawo lina la accessory basal nucleus ( Chiba et al., 2001 ). Chifukwa chake, ngakhale kuti ndi ochepa kwambiri komanso ofala kuposa ma projection ochokera ku BA 25, BA 32 imawoneka kuti ikugwirizana ndi njira zosankhidwa za amygdalar.
Barbas ndi Zikopoulos (2006) amanena kuti zomwe zimachitika pakati pa prefrontal ndi OFC ku amygdala zitha kukhala ndi zotsatira zosiyana pa ntchito ya amygdalar. BA25 pamwamba pa medial imapereka mphamvu yotulutsa mphamvu kwambiri ku magawo a basolateral a amygdala, zomwe zimapangitsa kuti hypothalamus iwonetse mphamvu. Mosiyana ndi zimenezi, posterior agranular OFC imasunga kwambiri minofu yolumikizana yomwe imazungulira basal nucleus (Onani Chithunzi 7 ). Mitsempha yolumikizana imapereka njira yoletsa kulowa mu central nucleus. Ikalimbikitsidwa, minofu yolumikizana imayimitsa njira yoletsa mphamvu kuchokera ku central nucleus kupita ku hypothalamus, zomwe zimapangitsa kuti hypothalamus isatseke. Mitsempha yopepuka yotsitsimula imafikanso ku central nucleus mwachindunji kuchokera ku posterior OFC, zomwe zimathandiza kuti OFC iwonjezere kapena kuchepetsa central nucleus fireing.
Zomwe zimayambira kutsogolo kwa hypothalamus ndi ubongo
Madera a OFC ndi medial PFC omwe ali ndi ma projection ku amygdala nthawi zambiri amafikiranso ku hypothalamus ndi autonomic brainstem/periaqueductal imvi regions ( An et al., 1998 ; Barbas et al., 2003 ; Price, 2006b ; Rempel-Clower & Barbas, 1998 ), zomwe zimapangitsa kuti zikhale zosavuta kusintha madera omwe amakhudzidwa ndi zotsatira zamaganizo (onani Chithunzi 8 ). Ma projection awa amawoneka olimba kwambiri kuchokera ku makoma apakati, komanso amachokera kudera lofanana ndi crescent pamwamba pa orbital komwe amygdalar input ndi yayikulu. Monga momwe zilili ndi kusowa kwake mwachindunji ku amygdala, DLPFC ndi FP sizimafikira mwachindunji ku malo awa. Kuphatikiza apo, magawo ambiri akutsogolo a OFC sawonetsa zotsatira zochepa mwachindunji ku malo awa odziyimira pawokha.
Maulalo mkati mwa lobe yamtsogolo
Monga tafotokozera pamwambapa PFC ikhoza kugawidwa m'magulu awiri apamwamba a phylogenetic. Maulalo apamwamba kwambiri mdera lililonse ali ndi madera omwewa, makamaka madera oyandikana ndi omwe omwe si gawo limodzi lachitukuko kutali ndi dera lomwe likukambidwa. Chifukwa chake, mwachitsanzo, zigawo za agranular zosakhazikika kumbuyo kwa OFC zimalumikizana kwambiri ndi zigawo zina za agranular ndi dysgranular orbital, koma nthawi zambiri sizimalumikizana ndi zigawo za isocortical monga dera la 46 m'mbali mwake. Komwe kulumikizana kwapakati kumachitika sikumadumpha zoposa gawo limodzi la zomangamanga. Mwachitsanzo, isocortical ventral area 46 yolumikizidwa mwamphamvu ndi isocortical dorsal area 46 mumadongosolo apakati, koma samalumikizana ndi madera ena opanda bwino a makoma monga subgenual cingulate (BA 46). Madera ena anterior ndi ofananira a OFC ali ndi kulumikizana kwakukulu ndi dera la 25 ndi dera loyandikana ndi 46, koma kulumikizana komwe kumadumpha sulfure wamkulu kumunsi kwa dera la 45 sikusowa kwenikweni.
Komabe, madera angapo a OFC akuwoneka kuti ali ndi kulumikizana mwachindunji ndi DLPFC. Makamaka madera 11m, 12o, 13a ndi 14r aliwonse ali ndi kulumikizana ndi DLPFC. Gyrus rectus (yomwe ikuphatikiza dera 14r) ikhoza kuwonedwa ngati gawo la njira yapakati kapena ngati dera losinthira pakati pa njira, kotero kulumikizana kwake sikuyimira kulumpha kwapakati pa njira. Komabe, madera 11m, 12o, ndi 13a amagawidwa ngati gawo la njira ya basoventral, kotero kulumikizana kwawo ndi DLPFC kumayimira kulumikizana kwapakati pa njira. Kuti mumvetse malo akuluakulu a netiweki m'madera awa, ndikofunikira kuganizira njira ina yogawa madera ozungulira ndi apakati. M'malo moyika zitsanzo pa phylogeny, Carmichael ndi Price (1996) amagawa OFC ndi khoma lapakati kukhala netiweki yozungulira ndi yapakati kutengera mphamvu ya kulumikizana pakati pa madera (Onani Chithunzi 9 ). Ndondomeko ya mtundu uwu wa magulu ikuwonetsa kuyanjana kwakukulu ndi kugawanika kwa phyolgenetic pakati pa machitidwe a basoventral ndi mediumdorsal, zomwe sizosadabwitsa poganizira za kapangidwe ka maulumikizidwe omwe afotokozedwa kale. Komabe, machitidwe awiriwa a magulu si ofanana kwathunthu. Chochititsa chidwi n'chakuti, madera onse ozungulira omwe amalumikizana ndi DLPFC ndi gawo la netiweki yapakati ya Carmichael ndi Price, kapena amaonedwa kuti ndi pakati pa ma netiweki. Mwachitsanzo, dera la 11m limaonedwa kuti ndi gawo la netiweki yapakati, chifukwa lili ndi maulumikizidwe ambiri ndi madera apakati a khoma kuposa ndi ena onse a OFC. Carmichael ndi Price amagawa madera 12o ndi 13a ngati madera olumikizirana chifukwa ali ndi maulumikizidwe olemera kumadera apakati ndi ozungulira. Mawonekedwe osiyanasiyana awa a kulumikizana akuwonetsa kuti padzakhala kusiyana kwakukulu kwa madera, kapena ngakhale ang'onoang'ono, pakutha kwa OFC kuyanjana ndi madera ena a prefrontal. Makamaka, gyrus rectus, komanso 11m, 12o, ndi 13a ali pamalo oti azitha kuyanjana ndi madera onse a makoma apakati (monga cingulate) ndi madera a DLPFC, pomwe madera ena ozungulira alibe ubale wolunjika uwu.
Kulumikizana kwa ma network koyambirira kumatsata njira zamtundu wa amygdala
Pa madera omwe alibe mphamvu yolunjika ku amygdala, kuthekera kokhudza kukonza kwa amygdalar kuyenera kudalira njira zosalunjika, ndipo njira izi zidzadalira kwambiri malo awo mkati mwa ma network akuluakulu a prefrontal. Popeza mphamvu ya ma projection a subgenual cingulate (BA 25) ku amygdala, ikhoza kupereka njira yofunika kwambiri yomwe madera osiyanasiyana a PFC amakhudzira amygdala. Monga momwe tikuonera kuchokera pa Chithunzi 9 , BA 25 imalandira ma projection akuluakulu kuchokera kumadera a medial network ndi madera omwe ali pamwamba pa orbital omwe amagwirizana ndi medial network. Mosiyana ndi zimenezi, ma projection ambiri a dorsolateral ndi ochepa. Vogt ndi Pandya (1987) akunena kuti BA 25 imalandira ma projection kuchokera ku DLPFC, ndipo makamaka imafotokoza zomwe zaperekedwa kuchokera ku dera 9 mu gawo lapamwamba la DLPFC. Komabe, mphamvu ya kulumikizanaku ikuwoneka yofooka, ndipo sinawonekere bwino m'maphunziro ena ( Barbas & Pandya, 1989 ). Komabe, BA 9 imagwirizana bwino ndi BA 32 pakhoma lamkati, lomwe limalumikizana kwambiri ndi BA 25 ( Carmichael & Price, 1996 ; Barbas & Pandya, 1989 ), motero imapereka njira yotheka yosalunjika yogwiritsira ntchito DLPFC kuti ikhudze BA 25. Mofananamo, dorsal BA 46 ilibe kulumikizana kwakukulu ndi BA 25, ndipo mwina ingafunike kugwiritsa ntchito BA 32, kapena magawo ena a cingulate cortex, kuti ilumikizane ndi BA 25.
Dorsal ACC (BA 24) imaperekanso malo ofunikira kwambiri ku amygdala. Chigawochi chili ndi mawonekedwe ambiri a PFC ( Carmichael & Price, 1996 ; Vogt & Pandya, 1987 ; Barbas & Pandya, 1989 ). Izi zikuphatikizapo kulowetsedwa kwakukulu kuchokera ku BA 9, komanso pang'ono kuchokera ku BA 46 mu DLPFC, magawo a BA 32, ndi BA 10 pakhoma la medial, ndi madera angapo a OFC (makamaka madera a medial/intermediate network 13a ndi Iai, ndi 12o). Chifukwa chake, dorsal ACC ikuwoneka kuti ili pamalo olimba kwambiri kuti iphatikize mbali za PFC zomwe zimagwira ntchito m'madera ambiri.
Ngakhale madera akunja a OFC ndi VLPFC amawoneka kuti ali ndi kuchuluka kwakukulu kwa zotsatira za amygdala kuposa zochokera ku amygdala, chifukwa magwirowa ndi ochepa, zigawo zakunja zitha kugwiritsa ntchito njira zosalunjika kuchita nawo amygdala. Kwa zigawo zakunja kwa OFC, izi zitha kutsogoleredwa kudzera kumadera akutsogolo kwa OFC. Mosiyana ndi izi, zigawo zoyendetsedwera, dera lakutali 12l ikhoza kupereka njira yokhayo yochitira nawo amygdala, atapatsidwa mawonekedwe apadera mkati mwa maukonde a pre-pre-pre-prealal and pre-prealal-amygdala.
4. Chitsanzo cha Zomanga
Makhalidwe a cytoarchitectural a dera la cortical amakhudza kwambiri momwe derali limagwirira ntchito ndi madera ena a ubongo. Makamaka, kuchuluka kwa granularity ndi kukula kwa laminar kumakhudza kuchuluka kwa ma feedforward ndi feedback projections ( Barbas & Rempel-Clower, 1997 ; Barbas 2000 ). Mkati mwa chitsanzo chomwe chaperekedwa ndi Barbas, ma feedforward projections amatanthauzidwa mwa kapangidwe kake kuti amachokera ku zigawo zapamwamba ndikufikira ku zigawo zakuya za cortex. Mu machitidwe a sensory, magawo oyambirira a processing stream amapereka chidziwitso ku magawo otsatira kudzera mu mtundu uwu wa feedforward projection ( Rockland & Pandya, 1979 ; Pandya, 1995 ). Mkati mwa machitidwe omwe ali ndi utsogoleri womveka bwino wa mayendedwe azidziwitso, monga machitidwe a sensory, ma feedforward projections amathanso kufotokozedwa ngati akukwera pamene akuyenda kuchokera kudera loyambirira kupita ku milingo yapamwamba mu processing stream (monga V1 kupita ku V2). Ponena za njira zodziwira, kutumiza chidziwitso kotereku kumagwirizana ndi zomwe akatswiri a zamaganizo amatcha kuti njira yoyambira pansi kupita mmwamba ( Kastner & Ungerleider, 2000 ).
Mosiyana ndi zimenezi, ma feedback projection amayamba m'magawo akuya a cortex ndipo amapita ku zigawo zapamwamba za cortex (onani Chithunzi 10 ). Mu machitidwe a sensory omwe ali ndi kapangidwe komveka bwino ka hierarchical, ma feedback projection awa angatanthauzidwe ngati akutsika, pamene akuyenda kuchokera ku magawo otsiriza kupita ku oyambirira a mtsinje wokonza sensory (monga, V2 mpaka V1). Ma feedback projection amagwira ntchito yosintha kapena kutsanzira ma computations omwe akuchitidwa m'magawo oyambilira okonza ( Raizada & Grossberg, 2003 ). Mwachitsanzo, ma feedback projection awa amagwira ntchito yothandiza kukulitsa mayankho a maselo omwe amalemba zinthu kapena malo omwe alipo, pomwe akuchepetsa kapena kuletsa mayankho ku zinthu zosayang'aniridwa ( Mehta et al., 2000 ; Saalmann et al., 2007 ). Ma feedback otere amathandiza mu njira zoyambira zowonera monga kusankhana kwa anthu ( Domijan & Setic, 2008 ; Roland et al., 2006 ), komanso kulola kuwongolera kuchokera pamwamba pa zomwe zikukonzedwa mu mtsinje wa chidziwitso ( Grossberg, 2007 ). Ponena za chidziwitso, kulamulira kumeneku kuchokera pamwamba mpaka pansi kumalola kusintha kwa magwiridwe antchito kutengera zomwe akuyembekezera, zolinga zomwe zilipo komanso kuyang'aniridwa ( Glibert ndi Sigman 2007 ).
Kuti timvetse bwino, ndikofunikira kusiyanitsa pakati pa mawu akuti kukwera/kutsika, feedforward/feedback, ndi pansi-mmwamba/pamwamba-pansi, chifukwa amatanthauza zinthu zosiyana (onani Penny et al. 2004 kuti mukambirane). Ma projection okwera ndi kutsika amatanthauza mawonekedwe enieni a hierarchical, ndipo mawuwa ndi othandiza kwambiri pankhani ya mitsinje yokonzedwa bwino. Timagwiritsa ntchito mawu akuti bottom-up ndi top-down ponena za njira zodziwira, pomwe bottom-up ikutanthauza njira zodziwira zokha, monga mayankho omwe amayendetsedwa ndi kuzindikira kwa chilimbikitso, ndi top-down ikutanthauza njira zomwe zimalola kusintha kosinthika kwa processing komwe kukugwirizana ndi zolinga ndi ziyembekezo zomwe zilipo. Mawu akuti feedforward ndi feedback munkhaniyi ali ndi matanthauzo awiri, chifukwa amatanthauzidwa ndi makhalidwe enaake a ma projection, koma amatanthauzanso mawonekedwe ogwiritsira ntchito chidziwitso.
Popeza kuti ma feedback projection ndi ma feedforward projection amatanthauzidwa ndi mawonekedwe a laminar, nkhawa ikhoza kubuka yokhudza njira zinazake za laminar zomwe Barbas ndi anzake amagwiritsa ntchito pofotokoza ma projection ngati feedforward kapena feedback kunja kwa mitsinje yogwiritsira ntchito sensory. Mu zitsanzo za mawonekedwe, ma feedforward projection nthawi zambiri amatanthauzidwa molingana ndi ma termination a laminar IV, ndi ma feedforward projections okwera omwe amachokera m'magawo apamwamba ndikutha mu layer IV (mosiyana ndi ma layer sheet ambiri; Felleman & Van Essen, 1991 ). Mosiyana ndi zimenezi, Barbas amagwiritsa ntchito tanthauzo lalikulu lomwe silikusiyanitsa mwachindunji pakati pa layer IV ndi infragranular layers. Kukulitsa kumeneku kuli pamwamba koyenera chifukwa cha kukhalapo kwa madera a prefrontal omwe alibe granular layer yolimba, ndipo ma dispersion laminar terminal patterns omwe amapezeka m'maderawa. Komabe, zotsatira zonse za extension iyi sizikufotokozedwabe.
Nkhani yosiyana pang'ono imabwera pa tanthauzo la ma feedback projections. Tanthauzo la Barbas la feedback limayang'ana kwambiri ma projections ochokera ku zigawo zakuya (za infragranular) ndikutha m'zigawo zapamwamba, mogwirizana ndi ntchito yoyambirira ya Rockland ndi Pandya (1979) . Komabe, Felleman ndi Van Essen (1991) akunena kuti ma downward feedback projections ena owonjezera akhoza kukhala ndi chiyambi cha bilaminar ndi kuphatikiza kwa ma infra- ndi supra-granular origins. Popeza, Barbas akupitirizabe kutanthauzira kosamala, kuvomereza critieria yake ndikosavuta. Komabe, zimapangitsa kuti chiwerengero cha ma projections omwe amadziwika kuti feedback mu PFC chikhale chokwera pogwiritsa ntchito tanthauzo la liberal.
Chinthu chofunika kwambiri pa chitsanzo cha kapangidwe kake ndichakuti mulingo wa mayankho ndi ma feedforward projection pakati pa madera umatsimikiziridwa kwambiri ndi kuchuluka kwa chitukuko cha cytoarchitectural cha madera. Ma projection ochokera ku cortex yosiyana kwambiri (monga, yosiyana kwambiri, ndi granular layer yokhuthala) kupita ku cortex yocheperako yopangidwa ndi cytoarchitectural amatsatira chitsanzo cha feedforward, pomwe omwe amachokera ku cytoarchitectural cortex yocheperako kupita ku cytoarchitectural cortex amatsatira mtundu wa feedback. Chitsanzochi chikugwirizana ndi zomwe zimawonedwa mu masensa, koma chitsanzocho chimawoneka chofala ku machitidwe angapo. Mu PFC, chitsanzo cha kapangidwe kake chimaneneratu kulinganiza kwa feedforward ndi ma feedback projection pafupifupi 80% ya nthawi, pomwe kulinganiza kwa mayankho ndi ma feedforward connections kukukulirakulira kusiyana kwa chitukuko cha cytoarchitectural pakati pa madera awiri omwe akukambidwa ( Barbas & Rempel-Clower, 1997 ).
Chofunikira pazomwe zimapangidwira pamitu yatsopano ndikuti zimatsogolera mwamphamvu zokhudzana ndi kulumikizana pakati pa zigawo zaubongo ngakhale pakakhala kuti palibe chidziwitso chachindunji. Zachidziwikire, pomaliza, njira zamagetsi kapena njira zina zomwe zimatha kuyesa mayendedwe a laminar ndizofunikira kuti zitsimikizire kuti njira zowonetsera zolimbitsa thupi mu PFC ndizofanana ndi zomwe zikuwoneka m'malingaliro am'malingaliro (mwachitsanzo, momwe amafotokozedwera mwatsatanetsatane ndi zomwe mukuganiza zikugwirizana. zofanana zokhudzana ndi chidziwitso zimayenda mosasamala kanthu mwa dongosolo lomwe likufunsidwa). Kafukufuku wamtunduwu amathandizanso kukonza njira zotanthauzira mozama mafayilo ndi mayankho ake. Pakadali pano, mtundu wa zomangamanga umapereka maziko olimba kwambiri omwe alipo pakulosera zamtundu wa kutuluka kwa chidziwitso mu PFC. Ngati kapangidwe kake kadali kolondola pakudziwitsidwa kwake kwa chidziwitso mkati mwa PFC, kumakhala ndi tanthauzo lofunika modabwitsa.
Mitundu ya Laminar ndi kulumikizana kwazinthu zoyambirira
Mogwirizana ndi chitsanzo cha kapangidwe kake, kusanthula kwa mapangidwe a laminar a ma projection kukuwonetsa kuti OFC yopanda ma granular imadziwika ndi mawonekedwe amphamvu a feedback mu kulumikizana kwake ndi madera opangidwa ndi cytoarchitectural kwambiri a PFC ( Barbas, 2000 ). Poyerekeza ndi machitidwe a sensory, izi zikutanthauza kuti ma projection a OFC amayang'ana kwambiri kutsanzira kapena kusintha ma computations. Mosiyana ndi izi, eulaminate DLPFC ili ndi milingo yayikulu kwambiri ya ma feedforward projections, omwe amalola kuti ipereke zotsatira kapena zotuluka za ma computations ake ku madera otsatira a ubongo. Kachitidwe kameneka ka ma feedforward ndi ma feedback projections kamasonyezanso kulumikizana kwapadera pakati pa OFC ndi DLPFC. Kulumikizana kwa lateral prefrontal ku OFC kumayambira makamaka m'magawo apamwamba a cortical (2-3) ndipo ma axon awo amatha makamaka m'magawo akuya (4-6), omwe amagwirizana ndi ma feedforward pattern ( Barbas & Rempel-Clower, 1997 ). Mosiyana ndi zimenezi, ma projection a OFC ku lateral PFC amachokera makamaka m'magawo akuya (5-6) ndipo ma axon awo amathera makamaka m'magawo apamwamba (1-3), omwe ndi mawonekedwe a feedback. Ma projection awa akuwoneka kuti amagwira ntchito pafupifupi 70-80% ya ma projection. Chifukwa chake, flow of information from OFC to granular PFC imakhala ndi feedback, pomwe flow of information in the sideline imagwirizana makamaka ndi feedforward pattern.
Mtundu wa kapangidwe kake ndizopatsa chidwi chifukwa umaganiza kuti kulumikizana kwachigawo kumatha kulumikizidwa molumikizana ndi kulumikizana kwa laminar. Ngati mtundu wa kapangidwe kake ndi yolondola, umatikakamiza kuti tisonkhane ndi njira yolumikizirana ndi yankho pakati pamagawo. Ma Model omwe akuwonetsa kuti PFC yotsogola imachita makamaka kapena kukhazikitsa njira zapamwamba ndizovuta kuyanjanitsa ndi mawonekedwe ake otchuka. Mofananamo, zitsanzo za OFC zomwe zimawona kuti zikungopereka zotsatira za ndalama (mwachitsanzo phindu lamalipiro) ku PFC yotsata, zimalephera kutengera kuthekera kwa kusankhana kwakanthawi mu PFC yoyambilira. Komabe, monga tafotokozera pambuyo pake m'nkhaniyi, kulumikizana komwe kulipo pakati pa madera a PFC, komanso pakati pa madera omwe akukhudzidwa ndi "kukhudzika" ndikuwonetsetsa nthawi zonse pazomwe zimachitika. Zowonadi, zitsanzo zomwe zilipo kale zimagwiranso ntchito zogwirizira za PFC, makamaka ntchito za DLPFC, potengera kuwongolera kwapang'onopang'ono, ndipo nthawi zambiri samaganizira zotheka kuti madera omwe sanapangidwe monga OFC angapangitse zigawo zoyambira za PFC.
Mitundu ya Laminar yolumikizana ndi prefrontal-amygdalar
Ma amygdala projections ku posterior OFC amayendetsa zigawo zonse za cortex, motero sizingakhale zongopeka pa ma feedforward kapena feedback type projections ( Ghashghaei et al., 2007 ). Komabe, n'zoonekeratu kuti pali gawo lamphamvu la feedforward ku ma projections awa kutengera laminar termination. Mosiyana ndi zimenezi, ma projections a OFC ku amygdala makamaka amachokera ku layer 5, zomwe zikusonyeza kuti ali ngati ma feedback projections (kutanthauza kuti amachita zinthu motsatira bias amygdalar processing m'malo mopereka chidziwitso china monga momwe zimakhalira ndi stimuli). Chochititsa chidwi n'chakuti, ma feedforward projections ochokera ku lateral PFC amalunjika ku layer 5 ya OFC, yomwe ndi layer yoyamba yotulutsa yomwe ma projections a OFC ku amygdala amachokera.
Kodi kuzindikira kwa maatomiki kungapangitse kukambirana pazokonzekera? Zaka zingapo zapitazi zawonapo kuphulika kwa chidwi mwa njira yomwe magawo osiyanasiyana aubongo amalumikizirana. Chidwichi chakhala chifukwa cha njira zomwe zatulukira njira zopangira kulumikizana kwa magwiridwe antchito ndi fMRI, ndikupereka koyamba kwa mwayi wofufuza mogwirizana pakati pa zigawo zaubongo mwa anthu athanzi. Komabe, kukambirana kwa izi, ndi mitundu yomwe imachokera muzilembedwezi, sikuti nthawi zonse kumakhala kovuta chifukwa cha anatomy. Pamene mitundu iyi yatchuka kwambiri, tikhulupirira kuti ndikofunika kuwunika momwe angagwirizane bwino ndi neuroanatomy yomwe afotokozeredwa pamwambapa. Tikukhulupirira kuti mitundu yotereyi imayenera kukhala yogwirizana ndi njira zonse zodzigwirizanitsira zomwe zimagwirizanitsa zigawo zosiyanasiyana zam'madera amtundu wathunthu komanso momwe lingagwiritsire ntchito njira / mayankho amachitidwe awa. Mitundu ikakhala kuti siyikugwirizana ndi izi, imalephera kufotokoza, kapena ingafunike malongosoledwe amomwe angagwiritsire ntchito bwino chifukwa cholingana ndi ubongo.
Mabuku a zamaganizo omwe akukula akuyesera kumvetsetsa momwe machitidwe a "kuzindikira" amagwirira ntchito ndi machitidwe a "maganizo". Ngakhale pali zoletsa zenizeni pakugawikana kopangidwa pakati pa machitidwe amalingaliro ndi malingaliro ( Pessoa, 2008 ), kusiyana kumeneku kwakhala kothandiza pofotokoza machitidwe osiyanasiyana monga kulamulira malingaliro, chilimbikitso, kupanga zisankho zachuma komanso njira yowunikira. M'magawo otsatirawa, tikufotokoza deta yomwe ikubwera ndi mitundu yowongolera malingaliro, kukumbukira kogwira ntchito ndi kuyanjana kwa dorsal-ventral prefrontal, ndikuyang'ana kwambiri kusagwirizana kwawo ndi deta ya anatomical. Timayang'ana kwambiri mabuku owongolera malingaliro, chifukwa mabukuwa akuchulukirachulukira akukambirana za psychopathology ndi chithandizo cha psychotherapeutic.
5. Malangizo Akumvera
Kulamulira maganizo kwatanthauzidwa ngati njira zomwe zimakhudzidwa ndi kusintha chiyambi, nthawi, mphamvu kapena zomwe zili mu yankho la maganizo ( Gross, 1998 ; Gross, 2008 ). Njira zowongolera malingaliro zimayambira pa zochita zomwe zimachitika nthawi yayitali malingaliro asanayambe, monga kusankha zochitika, mpaka njira zomwe zimachitika asanayambe kapena pamene malingaliro ayamba kuonekera, monga kuyika chidwi kapena kuwunikanso chidziwitso ( Gross, 1998 ). Ndi njira zomalizazi zomwe kufufuza za ubale pakati pa madera okhudzana ndi kuwongolera kuzindikira kwa malingaliro ndi omwe amagwirizana ndi yankho la maganizo kumakhala kofunikira kwambiri. Kafukufukuyu amafotokoza mosapita m'mbali kapena momveka bwino malamulo a malingaliro monga kuyika madera olamulira 'ozizira' a PFC kuti ayang'anire pansi njira zoyambira pansi kupita mmwamba, 'zotentha' zokhudzana ndi madera a subcortical limbic monga amygdala. Kulephera kwa kugwiritsa ntchito bwino njira zowongolera za PFC kuchokera pansi kupita pansi kapena njira zogwirira ntchito mopitirira muyeso za amygdala kwanenedwa kuti zikuthandizira mitundu ingapo ya matenda amisala ( Rottenberg & Gross, 2003 ; Rottenberg & Johnson, 2007 ).
Njira yowongolera malingaliro yomwe yakhala ikusamalidwa kwambiri m'mabuku ofufuza za ubongo ndi kuwunikanso malingaliro. Njira yowongolera iyi imaphatikizapo kutanthauziranso malingaliro mwanzeru kuti musinthe momwe munthu amayankhira ( Gross, 1998 ). Kuwunikanso kumaphatikizapo njira zambiri zokhudzana nazo. Mwachitsanzo, kuwunikanso kungayang'ane pa kutanthauziranso tanthauzo la munthu la chinthu chamaganizo kuti chikhale chodziyimira pawokha kapena chosakhala chofunikira. Kapenanso, kuwunikanso kungayang'ane pa kutanthauziranso chifukwa, zotsatira zake, kapena zenizeni za zoyambitsa malingaliro popanda kusintha ubale wa munthu ndi zoyambitsazo. Mwachitsanzo, munthu akhoza kuwunikanso ngozi yagalimoto m'mbali mwa msewu ngati kuti mwina imatha ndi magulu onse omwe achoka pachiwopsezocho mosamala. Maphunziro angapo ogwira ntchito a neuroimaging tsopano achitika panthawi yowunikiranso ntchito, ndipo alembedwa mu Gome 1 , pomwe malo a PFC akuwonetsedwa mu Chithunzi 11. Pogwiritsa ntchito mawu ofunikira kuwongolera malingaliro, kusokoneza ndi kuwunikanso, nkhani zoyeserera zoyesa kuwongolera malingaliro mwaufulu zidaphatikizidwa. Maphunziro a fMRI awa anali ndi maphunziro ophunzitsidwa owunikiranso zamaganizo, kuletsa malingaliro ndi kusokoneza anthu omwe si achipatala. Mndandanda uwu wa maphunziro okhudza malingaliro siwokwanira; mwachitsanzo, suphatikizapo mfundo zokhudzana ndi malingaliro monga kulamulira maganizo. Tikuwona kuti m'matebulo onse tasunga mayina (malembo a Brodmann omwe adagwiritsidwa ntchito, kapena mafotokozedwe a topographical/regional) omwe adagwiritsidwa ntchito ndi olemba mapepala oyambilira. Pali zochitika zina pomwe mafunso angabuke okhudza kugwiritsa ntchito zilembo, koma kusowa kwa dongosolo lofalitsa la "golide standard" la madera ambiri a prefrontal, sitinasinthe zilembo, kupatula kuti m'malemba timalemba makamaka ma activation a VLPFC omwe amagwirizana ndi gawo la posterior la BA 47/12. Popanda malire omveka bwino a gawo la BA 47/12 lomwe lili ndi maulumikizidwe ofunikira a amygdala mwa anthu, timaganizira gawo la dera lomwe lili posterior mpaka y = 32 monga momwe nthawi zambiri limayimira posterior BA 47/12. Timasonyezanso m'malemba pomwe OFC foci imagwirizana ndi malo a BA 13 (mosasamala kanthu za dzina lawo loyambirira).
Njira yodziwika bwino yophunzirira kubwerezabwereza imafuna ophunzira kuti aone zithunzi zosasinthika, zodzutsa chidwi kwambiri, komanso zosasinthasintha (monga kudula ziwalo, kuukira, kuwola ndi kutuluka m'mimba) ndikuyerekeza kuyambika kwa mitsempha panthawi ya mayeso omwe amaperekedwa kuti ayesedwenso ndi malingaliro ndi mayeso omwe amaperekedwa kuti ayang'ane mopanda chidwi (Eippert et al., 2006; Kim et al., 2007 ; Ochsner et al. , 2002; Ochsner et al., 2004 ; Phan et al., 2005 ; Urry et al ., 2006 ; Van Reekum et al., 2007 ). Ngakhale pali kusiyana kwa tsatanetsatane wa malangizo obwerezabwereza kuyambira pakuphunzira mpaka kuphunzira, nthawi zonse amafuna ophunzira kuti apange kutanthauzira kwatsopano kwa tanthauzo, chifukwa, zotsatira kapena kufunika kwa chithunzicho panthawi ya mayeso obwerezabwereza. Kuwunikanso kosiyana ndi kuwona zithunzi zoipa mosalamulirika kumabweretsa madera akuluakulu a PFC, kuphatikizapo DLPFC ndi VLPFC (nthawi zambiri zimakhala kumanzere kwambiri), ndi madera a ACC ya dorsal ndi/kapena PFC yapakati ngati zothandizira mbali zowongolera kuzindikira pakuwunikanso. Chithunzi 11 chikuwonetsa malo omwe kuwunikiranso kogwirizana ndi (zizindikiro za cyan zochepetsera zoyambitsa zoyipa, ndi zachikasu zochepetsera zoyambitsa zabwino) kuchokera ku maphunziro omwe atchulidwa pamwambapa.
Njira yofananira imagwiritsa ntchito zithunzi za kanema wosinthika m'malo mwa zithunzi zosasunthika. Maphunziro awa akuwonetsanso kulembedwa kwa DLPFC yapawiri panthawi yowunikiranso zamaganizo koma amasiyana ngati madera a ACC ndi PFC yapakati amalembedwanso kuti achepetse chisoni, kunyansidwa kapena chilakolako chogonana ( Beauregard et al., 2001 ; Goldin et al., 2008 ; Levesque et al., 2003 , 2004 ).
Mu maphunziro angapo obwerezabwereza pogwiritsa ntchito zithunzi zokhazikika kapena zosinthika, kuchepa kwa amygdala kunagwiritsidwa ntchito ngati njira yodziwira kusintha kwa valence yoyipa ndi kudzuka pamodzi ndi kuchepa kwa kusankhidwa kwa insula mu maphunziro ena ( Goldin et al., 2008 ; Levesque et al., 2003 ; Ochsner et al., 2002 ; 2004 ; Phan et al., 2005 ). Tikuwona kuti kufananiza kosavuta kwa zochita za amygdalar ndi zotsatira zoyipa kumakhala kovuta, chifukwa 1) amygdala imakhala yogwira ntchito m'mikhalidwe yomwe si yoyipa, ndi 2) zokumana nazo zoyipa zimakhudza zigawo za cortical ndi subcortical zomwe zimapitilira amygdala. Komabe, chifukwa cha chidwi chathu pakuyanjana kwa ubongo wachigawo, kutsika kwa ntchito ya amygdalar kumapereka chizindikiro chothandiza poyesa kuyanjana kwa prefrontal-limbic mosasamala kanthu kuti ntchito yake imagwirizana bwanji ndi zotsatira zoyipa. Kafukufuku wambiri amapeza kuchepa kwa amygdala yakumanzere, komanso nthawi zambiri amygdalae yapawiri, pogwiritsa ntchito kubwerezabwereza kuti muchepetse kukhudzidwa koyipa. Maphunziro ochepa okha ndi omwe adafufuzanso momwe zinthu zomwe zili ndi mphamvu zogonana zimayendera. Akafunsidwa kuti ayesenso kapena kuchepetsa mphamvu zogonana zomwe zimapangitsa kuti munthu ayambe kuchita zinthu zabwino kapena zogonana, kuchuluka kwa mphamvu zogwirira ntchito za amygdala kumanja kwa mphamvuzo kunachepa ( Beauregard et al., 2001 ; Kim & Hamann, 2007 ). Izi zitha kuyambitsa malingaliro okhudza momwe zinthu zilili pamaganizo, koma kawirikawiri, maphunziro oyesa momwe zinthu zilili ndi mphamvu zogonana za amygdala sizikuyenderana.
Njira ina yowongolera malingaliro imaphatikizapo kukumbukira zithunzi zabwino kapena zotonthoza kuchokera ku chilengedwe kapena zakale za munthu kuti zilowe m'malo kapena kuthana ndi zotsatira zoyipa. Kuyesa kwa machitidwe kukuwonetsa kuti kukumbukira kukumbukira kapena zithunzi zosagwirizana ndi malingaliro kumachepetsa zotsatira zoyipa ( Erber & Erber, 1994 ; Joormann, Seimer & Gotlib, 2007 ; Parrott & Sabini, 1990 ; Rusting & DeHart, 2000 ). Kafukufuku awiri wokhudza ubongo anayerekeza kuwongolera momwe munthu amakhudzidwira mwa kukumbukira chithunzi chotonthoza kapena kukumbukira ndi kuyembekezera kosalamulirika kwa mantha. Kalisch ndi anzake (2005) adayesa mayeso ndi mawu osonyeza ngati panali mwayi wogwedezeka pa mayesero amenewo kapena ayi. Mu mayeso owongolera, ophunzira adalangizidwa kuti asiyane ndi nkhawa zawo ndikuganiza za malo apadera omwe adadziwika kale. Mu mayeso osawongolera, ophunzira adalangizidwa kuti achite nawo mayankho awo amalingaliro. Kusanthula kwa ROI kunasonyeza kuti mtundu uwu wa malamulo unatenga dera la right anterolateral frontal cortex (MNI: 42, 48, 18) ndi malamulo pamaso pa nkhawa zomwe zinatenga madera a medial PFC ndi rostral ACC (−4, 46, 28). Mu kafukufuku wofanana, Delgado ndi anzake (2008b) adagwiritsa ntchito mabuloko amitundu yosiyanasiyana kuti asonyeze mayeso omwe kugwedezeka kunali kotheka, ndipo adapempha ophunzira kuti alamulire nkhawa zawo pokumbukira malo amodzi mwa awiri omwe adadziwika kale m'chilengedwe. Kusanthula kwawo kwa ROI kukuwonetsa kuti kukumbukira zithunzi za chilengedwe poyembekezera kugwedezeka kumatenga gyrus wapakati wakumanzere wakutsogolo (Talairach: −43, 28, 30). Kukula kwake kunagwirizana ndi kupambana kwa malamulo. Kulamulira kunapangitsanso kuti pakhale kuyambika kwa khoma lamkati lamkati ndi subgenual cingulate (BA 32; −3 36, −8 ndi BA 25; 0, 14, −11), zomwe olembawo akunena kuti zakhala zikugwirizana ndi kutha ( Phelps et al., 2004 ) komanso kuchepa kwa ntchito ya kumanzere kwa amygdalar. Ngakhale kuti maphunziro onsewa amagwiritsa ntchito njira zofanana, njira zawo zowunikira kuphatikizapo kusankha ma ROI ndi kuwonetsa zotsatira za tonic motsutsana ndi phasic zitha kukhala zomwe zimayambitsa kusiyana kwa madera omwe adanenedwa kuti akugwiritsa ntchito zithunzi zabwino kapena zotonthoza kuti athetse nkhawa zomwe zimagwirizanitsidwa ndi kuyembekezera kugwedezeka komwe kungachitike.
Mofanana ndi njira yakale yolamulira malingaliro, kusokoneza kumaphatikizapo kusunga chidziwitso chosalowerera komanso chosafunikira mu kukumbukira kwa munthu. Kafukufuku wamakhalidwe akuwonetsa kuti kuchita izi kumachepetsa kukhudzidwa koipa mwa anthu omwe ali ndi vuto la dysphoria komanso omwe alibe vuto la dysphoria ( Fennell, Teasdale, Jones, & Damle, 1987 ; Lyubomirsky, Caldwell, & Nolen-Hoeksema, 1998 ; Teasdale & Rezin, 1978 ). Mwa kutenga mphamvu yokumbukira yogwira ntchito ndi malingaliro osagwirizana ndi malingaliro, malingaliro ofanana ndi malingaliro amalepheretsedwa kupeza zinthu zowunikira ( Siemer, 2005 ). Kafukufuku wa Neuroimaging wa kusokoneza wagwiritsa ntchito njira ziwiri zosiyana. Yoyamba, yogwiritsidwa ntchito ndi Kalisch et al. (2006) , idagwiritsa ntchito njira yoyembekezera kugwedezeka, pokhapokha m'malo mopangitsa wophunzirayo kukumbukira kukumbukira kosangalatsa kapena kotetezeka, panali malangizo otseguka osokoneza momwe wophunzirayo adalimbikitsidwira kuganizira china chilichonse kupatula kugwedezeka komwe kungatheke. Njira imeneyi inapeza dera la kumanzere kwa PFC (MNI: −56, 30, 22) lomwe linali lochita bwino kwambiri m'mayesero omwe ophunzira adalangizidwa kuti azidzisokoneza okha kuposa m'mayesero osasokoneza. Njira yachiwiri yosokoneza inali ndi ntchito yosokoneza yomwe adapatsidwa (Sternberg working memory task) momwe wophunzirayo amasunga zilembo zingapo mu kukumbukira kogwira ntchito pamene akuwona zithunzi zoyipa kapena zosasinthasintha kenako kutsatira chithunzicho kuyenera kuyankha ngati chilembo chimodzi chinali mu seti yomwe anali kukumbukira. McRae et al. (2009) anena kuti kuchita ntchito yokumbukira ntchito pamene mukuyang'ana zithunzi zoyipa poyerekeza ndi kuwonera passive kumawonjezera yankho LOWONDA mu gyri yakumanzere ndi yakumanja yapamwamba komanso yapakati (MNI: BA6; −6, 10, 62 ndi −56, −4, 48 ndi 48, 42, 32; BA 9; −42, 22, 30 ndi 42, 30, 34; BA 10; −36, 62, 12 ndi 38, 64, 14) komanso PFC yotsika kumanja (BA47/12p; 36, 20, −4).
Malipoti ambiri okhudza ubongo okhudza kulamulira maganizo amasonyeza momveka bwino kuti madera a DLPFC akuchita zinthu zina zowongolera kuzindikira ndipo amasamala kuti asachepetse mayankho a amygdala mogwirizana mwachindunji ndi amygdala. Pankhani yowunikiranso ndi kusokoneza, chenjezoli ndilofunika kwambiri chifukwa njirazi zimapanga foci zomwe zimagawidwa ku PFC ( Chithunzi 11 ). Monga tanenera kale, mawonekedwe a anatomical projections kuchokera ku cortex akusonyeza kuti njira zolunjika kuchokera kumadera a DLPFC sizingakhale ndi mphamvu yolamulira kwambiri amygdala processing. Madera a PFC omwe ali ndi ma projection okhuthala pang'ono mu lateral PFC amapezeka kokha m'gawo laling'ono la VLPFC, makamaka m'madera akumbuyo kwambiri a BA 47/12. Tsoka ilo, monga tanenera kale, mayina omwe amagwiritsidwa ntchito pofotokoza za kuyambitsa m'derali m'maphunziro ambiri amapanga kusamveka bwino pankhani yokhudza kulumikizana ndi amygdala. Kafukufuku wofufuzanso, kukumbukira bwino kapena kukhudzidwa kwa zithunzi ndi kusokoneza nthawi zambiri amanena za kuyambitsa kwa zinthu m'madera onse a VLPFC ndi medial OFC ( Eippert et al., 2007 ; Goldin et al., 2008 ; Kim & Hamann, 2007 ; Lieberman et al., 2007; McRae et al., 2009; Ochsner, Ray et al., 2004 ). Makamaka, maphunziro ambiri obwerezabwereza amanena za kuyambitsa kwa BA 47/12 kwa mbali ziwiri pamene kumachepetsa malingaliro oipa kapena abwino. Monga taonera pamwambapa, BA 47/12 ndi dera lalikulu komanso losiyana ndipo madera okhawo a BA 47/12 ndi malo omwe ali ndi ziwonetsero zazikulu za amygdalar. Chifukwa chake, mawu amphamvu okhudza mphamvu yozindikira mwachindunji pa amygdala amakhala omveka bwino m'maphunziro amenewo ndi kuyambitsa kwa zinthu mu gawo ili la BA 47/12.
Madera apakati a PFC nthawi zambiri amaonedwa ngati ali ndi mwayi wopeza madera a subcortical monga amygdala. Komabe, malinga ndi ma mapu a medial direct connections omwe ali ndi amygdala, madera okhawo a subgenual cingulate (BA 25) ndi dorsal ACC (BA 24) ndi omwe ali ndi maulumikizidwe olunjika ndi amygdala. Maphunziro a Delgado ndi anzake (2008a , 2008b ) okha ndi omwe amanena za foci pamwamba pa medial omwe ali m'madera omwe amakhudza kwambiri amygdala. Popeza deta ya anatomical, zitha kuoneka zodabwitsa kuti kuyambitsa kwa BA25 sikumachitika kawirikawiri m'maphunziro awa. Komabe, n'zotheka kuti kusiya chizindikiro mu posterior VMPFC kwaletsa maphunziro kuwonetsa kuyambitsa kokhazikika m'derali. Kawirikawiri, maphunziro oletsa/kuletsa, kusokoneza ndi kuwunikiranso amangonena za foci mu BA 32, zomwe zingasonyeze kusintha kwapadera kwa amygdala, chifukwa cha momwe BA 32 imalowera ku amygdala.
Maphunziro a Correlational deactivation amygdala
Kuti timvetse bwino momwe kulamulira kwa malingaliro kuchokera pamwamba mpaka pansi kumagwirira ntchito ndi amygdala, kafukufuku wokhudza kulamulira kwa malingaliro wapita patsogolo kuposa kusiyanitsa ntchito ndi kulamulira kuti afufuze zokhudzana ndi kuchepa kwa ntchito ya amygdala (Onani Gome 2 ). Izi zikutanthauza kuti, m'malo mofunsa madera omwe akugwira ntchito yomwe imadziwika kuti imachepetsa kulamulira kwa ntchito ya amygdala, adayesa momveka bwino kulumikizana kapena kulumikizana kogwira ntchito/kogwira ntchito pakati pa amygdala ndi ubongo wonse panthawi ya magwiridwe antchito a kulamulira kwamalingaliro. Kapenanso, maphunziro ena adagwirizanitsa kuchepa kwa amygdala ndi madera omwe adadziwika kale kuchokera ku kusiyanitsa kwakukulu kwa kulamulira. Maphunzirowa akuwonetsa kuti kuchepa kwa amygdalar kumagwirizana molakwika ndi madera ambiri a ntchito ya PFC. Chofunika kwambiri ndi kuyambitsa mu VMPFC, kuphatikiza BA 11m/14r (5, 37, −12; −6, 46, −20: Urry et al., 2006 , Ochsner et al., 2002 motsatana). Kuphatikiza apo, madera a subgenual ndi pregenual cingulate adawonedwa kuti ali ndi mgwirizano woipa ndi ntchito ya amygdala panthawi ya kulamulira. Mwachitsanzo, Urry ndi anzake (2006) adanenanso kuti dera la BA 32/10 (lomwe limakhala pa −23, 43, −10) linafalikira m'mimba komanso pakati. Delgado et al. (2008b) adanenanso kuti pali mgwirizano wotsutsana pakati pa ntchito ya BA 32 (0, 35, −8) ndipo amygdala imachepa. Madera a Posterior (BA 13) a OFC adagwirizananso ndi kuletsa kwa amygdala (−24, 28, −14; 26, 24, −22: Banks et al, 2007: −30, 22, −16; 34, 24, −16: Ochsner, Ray et al., 2004 ). Madera ochepa a PFC mu BA 47 (34, 54, 12) ndi BA46 (−54, 12, 12: Urry et al., 2006 ; −54, 42, 12: Ochsner et al., 2002 ), nawonso adawonekera m'maphunzirowa. Maphunziro awiri adagwirizanitsa ziwerengero za madera enaake a DLPFC ndi madera apakati, zomwe zidagwirizana ndi kuchepa kwa mayankho a amygdala. Mu kafukufuku wopangidwa ndi Urry et al. (2006) , kusanthula kwapakati kunawonetsa kulumikizana pakati pa amygdala, BA 10 (3, 63, 18) ndi chigawo cha DLPFC (−50, 23, 19). Delgado et al. (2008b) m'malo mwake adagwiritsa ntchito chigawo chapakati cha BA 32 ngati mbewu ya kusanthula kwawo kwa PPI komwe kenako kudazindikira chigawo chamanzere cha amygdala ndi chigawo cha DLPFC. Chofunika kwambiri, maphunzirowa apeza madera ofanana ndi kuchepa kwa amygdala omwe atchulidwanso pamwambapa kuti akuwonetsa ku amygdala monga dorsal anterior cingulate, subgenual cingulate ndi posterior orbitofrontal cortex.
Mwa madera omwe adanenedwa kuchokera ku kusanthula kwa mgwirizanowu kapena kusanthula kwambiri kwa regression, ochepa mwa iwo ali ndi kulumikizana mwachindunji ndi amygdala. Madera ofala kwambiri omwe ali ndi mgwirizano wolakwika ndi mayankho a amygdala ndi madera a posterior OFC ndi subgenual cingulate ndi VLPFC ( Chithunzi 12 ). Mwa madera a lateral prefrontal, gawo lokha la posterior lateral la BA 47/12 lili ndi ma projection amphamvu ku amygdala. Madera a anterior BA 32 amadziwikanso mu ma projection a correlational, omwe angawonetse ma projection ku assessory ndi basal lateral nucleus ya amygdala ( Cheba et al, 2001 ).
Mitundu yamalingaliro amomwe akumvera
Mpaka pano, chitsanzo chapamwamba kwambiri choyendetsera malingaliro chochokera ku deta chimachokera ku kafukufuku wa kubwerezabwereza kwabwino kwa Wager ndi anzake (2008) . Chosintha cha zotsatira zake ndi kusintha kwa zotsatira zoyipa zomwe zanenedwa ndi munthu payekha. Njira yolinganiza kapangidwe kake idagwiritsidwa ntchito pa data ya neuroimaging kuchokera ku paradigm yowunikiranso yofanana ndi yomwe idagwiritsidwa ntchito ndi Ochsner et al. (2002 ; 2004) . VLPFC yoyenera idasankhidwa ngati poyambira pa kusanthula, ndi ma coordinates omwe ali m'dera lomwe mwina limaphatikizapo gawo lakumbuyo la dera 47/12 ndi ma projection ku amygdala. Olembawo adagwiritsa ntchito njira ya ROI koyamba kuyesa udindo wa amygdala ndi nucleus accumbens ngati oyimira pakati pa VLPFC yoyenera ndi kuchepa kwa zotsatira zoyipa zomwe zidadziwika ngati muyeso waukulu wa kupambana kwa kubwerezabwereza. Mu kusanthula kwa ROI uku, mapangidwe onse awiri adawonetsedwa kuti alumikizane ndi ubale pakati pa VLPFC yoyenera ndi kuchepa kwa zomwe zanenedwa ndi munthu payekha pa zotsatira zoyipa (onani Chithunzi 13 ).
Olembawo adagwiritsa ntchito kusanthula kwa magulu onse a ubongo ndi malingaliro osasintha kuti adziwe maukonde awiri ngati oyimira ubale pakati pa VLPFC ndi kusintha kwa zotsatira zoyipa zomwe zanenedwa ndi anthu (onani Chithunzi 14 ). Network imodzi ili ndi tsankho losalunjika pakuwonjezera kusintha kwa zotsatira zoyipa. Network iyi imaphatikizapo madera a nucleus accumbens, subgenual cingulate (BA 25), pre-SMA, precuneus, DMPFC (MNI: 24, 41, 40), ndi superior frontal gyrus (24, 21, 58). Pakati pa madera awa, nucleus accumbens ndi subgenual cingulate ndizolumikizana kwambiri ndi amygdala. Network yachiwiri yomwe yadziwika ili ndi tsankho losalunjika pakuchepetsa kusintha kwa zotsatira zoyipa ndikuchepetsa kupambana kobwerezabwereza. Network iyi ikuphatikiza rostral dorsal ACC, amygdala (bilateral) ndi posterior-lateral OFC (48, 24, −18). Ntchito yamtsogolo iyenera kufotokoza momwe zigawo za maukonde zimagwirira ntchito komanso ngati maukonde awa ndi enieni a mtundu uwu wa njira yolamulira malingaliro.
Ofufuza angapo apereka zitsanzo za chiphunzitso cha njira za mitsempha zomwe zimayang'anira momwe munthu amamvera. Chitsanzo chosavuta kwambiri mwa izi chikusonyeza kuti madera ochepa amakhudza mwachindunji amygdala. Delgado et al. (2008b) , Hansel ndi von Kanel (2008) ndi Quirk and Beer (2006) aliyense akupereka lingaliro lakuti ventromedial PFC pansi imalamulira madera a amygdala. Zitsanzozi zimayesa kwambiri kutithandiza kumvetsetsa maziko a neuroanatomical a malamulo a malingaliro a anthu m'mabuku ambiri a nyama okhudza kutha kwa zinthu ndi kulumikizana kwa ventromedial PFC ndi magulu ophatikizana mu basolateral amygdala ( Morgan, Romanski & LeDoux, 1993 ; Likhtik et al., 2005 ; Quirk et al., 2000 ). Quirk and Beer (2006) amamanga pa kukhalapo kwa zotsatira zolimbikitsa komanso zoletsa za "ventral" medial PFC projections ku amygdala mwa anthu ndi makoswe. Gawo la subgenual cingulate, BA 25, likunenedwa kuti ndi loletsa kwambiri pomwe BA 32 yomwe ili dorsal ndi anterior ikuyembekezeka kukhala ndi maulumikizidwe oyambitsa ndi amygdala. Onse awiri a BA 25 ndi 32 ali ndi maulumikizidwe ndi amygdala. Komabe, BA 32 ili ndi maulumikizidwe ochepa kwambiri.
Phillips et al (2008) apanga chitsanzo cha dera chomwe chikuyesera kufotokoza maziko a mitsempha ya mitundu yosiyanasiyana ya malamulo okhudza malingaliro (onani Chithunzi 15 ). Chitsanzochi chili ndi zigawo za DLPFC, OFC, VLPFC, DMPFC ndi ACC. Chochititsa chidwi kwambiri, olembawo amasiyanitsa madera omwe akukhudzidwa ndi malamulo okhudza malingaliro okha (mu ACC yapansi pa dziko ndi ya rostral) ndi madera omwe amasankhidwa kuti azitsatira malamulo okhudza malingaliro mwaufulu (DLPFC ndi VLPFC). Amanena kuti madera otsirizawa ndi atsopano mwachibadwa ndipo amapereka mayankho ku njira zakale zopangira malingaliro. Koma OFC, DMPFC ndi ACC, ndi madera akale mwachibadwa omwe amafotokozedwa kuti amagwira ntchito kudzera mu njira zoperekera uthenga kuti atumize zambiri zamkati mwa DLPFC ndi VLPFC. Olembawo amaika DMPFC ngati njira yomwe OFC imaperekera chidziwitso chamtengo wapatali ku madera a ubongo a neocortical kuti achite zisankho.
Mbali imodzi yapadera ya chitsanzo ichi ndi kufotokoza momveka bwino njira zoperekera ndemanga ndi mayankho. Chitsanzochi ndi chokongola mwachibadwa ndipo chikugwirizana bwino ndi malingaliro achikhalidwe okhudza DLPFC yomwe imagwiritsa ntchito mphamvu kuchokera pamwamba kupita pansi pa madera "omwe amakhudzidwa ndi maganizo". Komabe, n'zovuta kugwirizanitsa lingaliro ili ndi chitsanzo cha kapangidwe kake, chifukwa cha kufalikira kwa kulumikizana kwa PFC ( Barbas & Rempel-Clower, 1997 ; Barbas, 2000 ). Zoonadi, chitsanzo cha kapangidwe kake chikusonyeza kuti kuyenda kwa chidziwitso pakati pa DLPFC ndi OFC kwenikweni kuli kosiyana ndi njira zomwe zimayambira mu OFC ndikupita ku DLPFC zomwe zimafotokozedwa makamaka ngati mayankho, ndipo zomwe zimachokera mu DLPFC ndikupita ku OFC zomwe zimafotokozedwa kwambiri ngati mayankho.
Phillips et al. mtunduwu umadziwikanso poyika madera ena otchedwa "automatic regulation" madera monga subgenual cingulate ndi OFC ngati njira yayikulu yomwe zigawo zatsopano za phylogenetically zimakhudzira madera amphamvu monga amygdala. Izi ndizofanana kwambiri (makamaka gawo lokhalitsa la cingrate) ndi makonzedwe amtaneti omwe afotokozedwa pamwambapa. Titha kuwerengetsa, komabe, kuti pakhoza kukhala njira zopitilira imodzi momwe madongosolo odzifunira amomwe angakhudzire zochitika za amygdala. Makamaka, VLPFC yaposachedwa imatha kuwongolera machitidwe a amygdala popanda kugwiritsa ntchito gawo limodzi mwazigawo “zodziwikiratu”, potengera zolingalira zake za amygdala nuclei.
Mwachidule, deta yambiri ikuwonetsa kutenga nawo mbali kwa madera a PFC panthawi ya ntchito zowongolera malingaliro, ndi zochitika m'magulu osankhidwa bwino a madera (BA 47/12, BA25 ndi BA 32) zomwe zikuwonetsa kuyanjana ndi kuthekera kochepetsa zochita za amygdala. Ma model okhwima kwambiri aperekedwa kuti afotokoze deta iyi. Kutuluka kwa ma model awa ndikokopa, monga momwe olemba awo akusonyezera nkhawa yakuti ma network olumikizirana omwe akuperekedwa ndi otheka. Komabe, tikuzindikira kuti palibe ma model mpaka pano omwe avomereza momveka bwino njira yolumikizirana pakati pa madera osiyanasiyana a PFC. Mwachitsanzo, Wager et al (2008) amapereka chitsanzo chovuta kwambiri cha njira inayake yowongolera malingaliro, koma sichikukhudza mtundu wa kayendedwe ka chidziwitso pakati pa madera a zigawo. Phillips et al. akuphatikiza momveka bwino lingaliro la chidziwitso cha feedforward ndi mayankho, koma sagwirizanitsa malingaliro awa ndi mawonekedwe omwe awonedwa a mayankho ndi malingaliro a feedforward m'madera omwe akukambidwa. Tikukhulupirira kuti kuyanjanitsa nkhanizi kumapereka chimodzi mwazovuta zazikulu kwa ofufuza omwe akuyesera kumvetsetsa magawo a neural a malamulo a malingaliro.
6. Kuwongolera kuzindikirika kwa kusokonezedwa kwa malingaliro
Ngakhale kuti kusanthula kwathu kwakukulu kwayang'ana kwambiri pa maphunziro okhudza kulamulira malingaliro, nkhani zambiri zofanana zimabuka poganizira mabuku okhudza kulamulira maganizo. Mwambiri, kulamulira maganizo kumatanthauza njira zapamwamba zoyendetsera ntchito zomwe zimalimbikitsa kukonza zolinga, pomwe zikuletsa kukonza kosafunikira cholinga. Mawuwa amagwiritsidwa ntchito makamaka pofotokoza ntchito zomwe zimafuna chidwi chosankha chidziwitso chokhudzana ndi cholinga komanso kuletsa chidziwitso chosafunikira cholinga, komanso kusankha mayankho olimbikitsa cholinga komanso kuletsa mayankho otsutsana ndi cholinga. Njira yosankha yotereyi nthawi zambiri imaperekedwa momveka bwino pankhani ya kusintha kwa pamwamba ndi kusankha njira zoyendetsera. Maphunziro okhudza kulamulira maganizo angawonedwe ngati gulu linalake la kulamulira maganizo lomwe limayang'ana kwambiri pakusintha yankho logwirizana nalo. Mosiyana ndi zimenezi, mitundu ina yambiri ya maphunziro okhudza kulamulira maganizo okhudza kumvera imayang'ana kwambiri kuthekera kogonjetsa zosokoneza zomwe zimachitika chifukwa cha kukhudzidwa kwa maganizo. Chifukwa cha makhalidwe awo enieni (nthawi zambiri odzipangira okha) omwe amakopa chidwi ( Most et al., 2005 ; 2007 ; Pessoa, 2008 ), kukakamiza maganizo nthawi zambiri kumabweretsa kufunikira kwakukulu kwa kulamulira maganizo kuti tisunge kusankha koyenera kwa chidziwitso chogwirizana ndi cholinga. Kufunika kopewa kusokonezedwa ndi zinthu zomwe zimapangitsa munthu kukhala ndi maganizo, makamaka kumachitika m'maphunziro omwe zinthu zomwe zimapangitsa munthu kukhala ndi maganizo zimachitikira nthawi imodzi ndi zinthu zina, zomwe sizikugwirizana ndi ntchito zina, kapena panthawi yogwira ntchito yokumbukira, komwe kusokonezeka kungasokoneze kusunga chidziwitso pa intaneti. Tikuwunikanso mwachidule maphunzirowa kuti tiwonetsetse kugwirizana kwawo ndi mabuku okhudza malingaliro. Kuti mumve zambiri za izi, owerenga mabukuwa atumizidwa kwa Banich et al. (2009).
Kuponderezedwa kwamalingaliro amakhudzidwa panthawi yodziwa ntchito
Kafukufuku wambiri wagwiritsa ntchito njira zomwe ophunzira ayenera kuyankha ku chinthu cholimbikitsa chomwe sichili ndi malingaliro (monga mtundu) ndipo osasokonezedwa ndi zomwe zili mumtima (monga mawu amalingaliro), kapena kusamalira zomwe sizili ndi malingaliro (monga nyumba) pomwe akunyalanyaza zomwe zimawalimbikitsa (nkhope yoopa). Mwachitsanzo, madera a rostral (dorsomedial, pregenual ndi dorsal ACC) komanso madera onse a DLPFC ndi VLPFC onse awonedwa mu njira zamaganizo za Stroop momwe anthu ayenera kupewa kusokonezedwa ndi zomwe zili mumtima mwa mawu ( Whalen et al., 1998 ; Compton et al., 2003 ; Herrington et al., 2005 ; Mohanty et al., 2007 ). Kuti mumve zambiri za momwe kulamulira chidwi ndi kulamulira malingaliro kungakhudzire magawo omwewo a neurocognitive, owerenga atumizidwa ku Blair & Mitchell (2009) ndi Mitchell (2011).
Kuchepa kwa kutanthauzira kwa ambiri mwa malingaliro amenewa kumachitika chifukwa chakuti sizikudziwika nthawi zonse ngati madera awa akukhudzidwa chifukwa akugwiritsa ntchito mphamvu zowongolera kuzindikira, kuyang'anira mikangano, akugwiritsidwa ntchito chifukwa cha mikangano/kusokoneza kwakukulu popanda kulamulira mikangano/kusokoneza, kapena akungoyankha momwe zinthu zilili. Mwachitsanzo, Mohanty ndi anzake (2007) akuwonetsa bwino kuti dera la pregenual cingulate (pafupifupi BA 24/32) likuwonetsa kuwonjezeka kwa kuyambitsa panthawi ya ntchito ya Stroop ndi mawu okhudza mtima, ndipo izi zikugwirizana ndi kuwonjezeka kwa nthawi yochitapo kanthu pantchitoyo. Izi zitha kutanthauziridwa ponena za rACC yomwe ikugwira ntchito kuti igwiritse ntchito mphamvu zowongolera kuzindikira pa zosokoneza zamaganizo. Komabe, popeza kuyambitsa dera lino kumagwirizana ndi nthawi yochulukirapo yochitirapo kanthu, kuchuluka kwake kwa kuyambitsa sikukuwoneka kuti kukugwirizana ndi kuletsa bwino zosokoneza. Kuphatikiza apo, zidawonetsa kuwonjezeka kwa kulumikizana kwa magwiridwe antchito ndi amygdala, zomwe sizikugwirizana ndi lingaliro lakuti rACC inali kuyendetsa kutsika kwa amygdala. Zoonadi n'zodziwika kuti olembawo akunena kuti m'malo mowonetsa malamulo a rACC a amygdala, kulumikizana kwakukulu panthawi yokumana ndi zosokoneza zamaganizo kungawonetse malamulo a amygdalar kapena kulowetsa ku rACC, m'malo mosiyana.
Pakati pa umboni wodabwitsa kwambiri wokhudza kulamulira kwa chidziwitso cha prefrontal pa processing emotional mu amygdala umachokera ku kafukufuku wa Etkin et al. (2006) , pomwe ophunzira adachita ntchito yofanana ndi Stroop momwe mawonekedwe a nkhope yamaganizo amatha kukhala ofanana kapena osagwirizana ndi mawu otchula malingaliro. Kapangidwe ka kafukufukuyu kanali kovuta kwambiri chifukwa olembawo sanayang'ane kwambiri kufananiza kophweka kwa mayesero amalingaliro ndi osalowerera kapena mayeso osagwirizana ndi ofanana, koma m'malo mwake adayang'ana zotsatira panthawi ya mayeso osagwirizana omwe adatsata makamaka mayeso ofanana kapena osagwirizana. Chochititsa chidwi n'chakuti, DLPFC, dera la DMPFC mu superior frontal gyrus, ndi rostral (pregenual) ACC adawonetsa kuyambitsa panthawi ya mayeso osagwirizana omwe adadalira ngati mayeso am'mbuyomu anali ofanana kapena ayi. DLPFC (ndi DMPFC) idayankha bwino kwambiri mayeso osagwirizana omwe adatsata mayeso ofanana, pomwe rostral ACC idayankha kwambiri mayeso omwe adatsata mayeso ena osagwirizana. Kafukufukuyu ndi umodzi mwa maphunziro ochepa omwe adachitika m'mabuku owongolera chidziwitso omwe adafufuza makamaka ubale wa zigawo za prefrontal cortical ndi ntchito ya amygdala (pogwiritsa ntchito kusanthula kwa psychophysiological interaction, Friston et al. 1997 ). Chodabwitsa n'chakuti, ntchito yayikulu mu rostral ACC idagwirizana ndi ntchito ya amygdala yakumanja. Kutengera ndi momwe mayankho a amygdala amachitikira, olembawo akunena kuti ntchito ya amygdala imagwirizanitsidwa ndi kuchuluka kwa mikangano pa mayeso enaake, ndipo poletsa ntchito ya amygdala, rostral ACC imapereka ulamuliro pa mkanganowu. Kuthandizira lingaliroli kumachokera ku deta ya khalidwe chifukwa iwo omwe adawonetsa kulumikizana kwakukulu kwa ntchito yotsutsana pamayeso osagwirizana adawonetsa kuthetsa mkangano waukulu monga momwe zimayesedwera ndi nthawi yochitira pa ntchitoyi. Mu kafukufuku wotsatira Etkin et al. (2010) adawona kuti kuletsa ntchito ya amygdala kumawoneka kofooka mwa odwala omwe ali ndi matenda a nkhawa omwe amakhudzana ndi zowongolera zathanzi, zomwe zimapangitsa kuti pakhale vuto lolamulira kusokonezeka kwa malingaliro kapena mkangano mwa odwala awa.
Phukusi lofunikira ndiloyenera kuyang'anira izi. Choyamba, kafukufuku wopangidwa ndi gulu la Etkin sakusonyeza kukhalapo kwa zoletsa zamdziko lonse za amygdala ndi madera a PFC panthawi yakusokonekera kwamalingaliro, kapena kusungidwa kosalekeza kwa "zigawo zazidziwitso," koma makamaka ntchito inayake yomwe imatengera mtundu wa kulimbana pakati zisanayambike. Ngati ndi zolondola, kuthekera kowona mayanjano ophatikizika pakati pa zida zodziwika bwino (kapena madera ena a PFC) ndi amygdala kungakhale kantchito komanso kosanthula kwenikweni.
Umboni wina umasonyezanso kuti madera ena a prefrontal, makamaka dorsal ACC, angagwiritse ntchito mphamvu zoletsa amygdala. Mwachitsanzo, mu kafukufuku wogwiritsa ntchito njira yomweyi monga Etkin et al. ( Chechko et al., 2009 ), odwala omwe ali ndi vuto la mantha adawonetsa kuchepa kwakukulu kuposa kulamulira bwino panthawi ya mayeso osagwirizana ndi malingaliro, komanso amygdala yapamwamba, koma ntchito yochepa ya dorsal ACC/DMPFC, zomwe zidapangitsa kuti pakhale lingaliro lakuti vuto la mantha limadziwika ndi kusakwanira kwa DMPFC/dorsal ACC. Mofananamo, Hariri et al., (2003) adawona mgwirizano woipa pakati pa amygdala ndi dorsal ACC (ndi VLPFC) pamene anthu adayenera kulemba chizindikiro motsutsana ndi zithunzi zamaganizo (ndi ntchito ya amygdala ikuwonjezeka pa mkhalidwe wa machesi, ndi VLPFC ndi dorsal ACC ikuwonjezera ntchito panthawi ya chizindikiro). Zanenedwanso kuti dACC ikhoza kugwiritsa ntchito mphamvu zowongolera amygdala ngakhale popanda mikangano yeniyeni kapena kusokoneza maganizo a ntchito. Pezawas et al. (2005) adawona mgwirizano waukulu pakati pa dACC ndi ntchito ya amygdala panthawi ya ntchito yofananira ndi zoopsa. Zingadziwikenso kuti ACC yaching'ono mu kafukufuku wa Pezawas inali yogwirizana bwino ndi ntchito ya amygdala, zomwe zikusonyeza mgwirizano wapadera pakati pa madera osiyanasiyana a cingulate ndi amygdala, ndipo zikusonyezanso, monga momwe zilili mu pepala la Monhaty et al. (2007) , kuti rACC, nthawi zina imakhala yabwino, osati yoipa, yolumikizidwa ndi amygdala.
Kukumbukira Memory
Gulu lina la zoyeserera zowongolera kuzindikira limayang'ana kwambiri kuthekera koletsa kusokonezeka kwa malingaliro panthawi yogwira ntchito zokumbukira. Chifukwa kuchuluka kwa chidziwitso chomwe chingasungidwe ndikusinthidwa pa intaneti ndi kochepa ( Cowan, 2010 ), ndikofunikira kuti anthu aziika patsogolo bwino chidziwitso chomwe chimalowa mu sitolo iyi ya pa intaneti. Mwanjira yabwino, tiyenera kusunga chidziwitso chofunikira poyerekeza ndi chidziwitso chosafunikira kwenikweni, komanso kuti tithe kutaya zomwe zili mu kukumbukira kogwira ntchito pamene chidziwitso chofunikira kwambiri chikalowa m'malo mwa zolinga zam'mbuyomu. Motero, kukumbukira ntchito kumapereka gawo lothandiza pofufuza momwe malingaliro ndi kuzindikira zimagwirira ntchito, makamaka poganizira udindo wofunikira wa DLPFC ndi VLPFC pakugwira ntchito kukumbukira ( Badre et al., 2005 ; Blumenfeld et al., 2010 ; Curtis & D'Esposito, 2004 ; Jonides et al., 2005 ; Levy & Goldman-Rakic, 2000 , Nee & Jonides, 2010 ; Postle, 2006 ; Thompson-Schill et al., 2002 ).
Malipoti awiri a Dolcos ndi anzake ndi ofunika kwambiri chifukwa amagwirizanitsa makamaka kuyambika kwa ubongo ndi magwiridwe antchito abwino kapena kuthana ndi mavuto okhudzana ndi kulumikizana kwa ntchito ( Dolcos ndi McArthy 2006 ; Dolcos et al., 2006 ). Malipoti onsewa adasanthula deta kuchokera ku ntchito yosavuta yolumikizana ndi nkhope yomwe zithunzi zamaganizo kapena zandale zidawonetsedwa panthawi yochedwa (yosamalira) ya ntchitoyi. Mu kafukufuku woyamba, adawonetsa kuti ventrolateral cortex (BA 45/47) idagwira ntchito molumikizana nthawi zonse panthawi yamalingaliro poyerekeza ndi zosokoneza zandale. Ophunzira omwe adawonetsa zochita zazikulu za ventrolateral pamaso pa zosokoneza zamaganizo adawona kuti zosokonezazo sizikusokoneza kwenikweni. Mu kafukufuku wotsatira adawonetsa kuti ntchito ya BA 45 yotsala (koma osati yoyenera BA 45) idasiyanitsa pakati pa mayeso omwe anthu adapambana motsutsana ndi omwe adalephera kunyalanyaza chosokoneza (monga momwe zasonyezedwera ndi magwiridwe antchito ochedwa kapena olakwika). Dolcos et al. (2006) nawonso adanenanso za kulumikizana kwa ntchito ya VLPFC-amygdala, ndi madera onse awiri akuwonjezeka panthawi yamalingaliro poyerekeza ndi mayeso a neutral distractor. Chofunika kwambiri ndichakuti kulumikizana kumeneku kuli m'njira yabwino ndipo sikungatanthauzidwe ngati kukuwonetsa kuletsa kuwombera kwa amygdalar.
Kafukufuku wa Dolcos ndi anzake amaperekanso umboni wa njira zolekanitsira ntchito ndi kuzimitsa ntchito m'madera akutsogolo. Makamaka, madera akutsogolo anawonjezeka ndi zosokoneza zamaganizo, pomwe DLPFC (BA 9/46) inachepa, zomwe zikusonyeza kuti pali ubale wobwerezabwereza pakati pa madera awa. Ubale wobwerezabwerezawu ukufanana ndi chitsanzo cha inverse dorsal vs. ventral chomwe Perlstein et al. (2002) adawona anthu omwe adachita ntchito yokumbukira momwe zithunzi zokhudzana ndi malingaliro zimawonekera ngati zizindikiro ndi ma probe [chosangalatsa n'chakuti, ubale wobwerezabwereza unali wogwirizana kwambiri ndi valence pomwe DLPFC ikukwera ndi zolimbikitsa zabwino ndi madera akutsogolo (BA 10/11) zomwe zikuwonetsa kuchuluka kwa zochita za zoyambitsa zoyipa]. Kachitidwe kotsutsana pakati pa madera akutsogolo ndi akutsogolo kwa PFC kawonedwanso m'ma paradigms ena okumbukira ntchito, ndi DLPFC yayikulu poyerekeza ndi zochitika zakutsogolo zamkati zomwe zikugwirizana ndi kuchuluka kwa kukumbukira ntchito ( Rypma et al., 2002 ; Woodward et al., 2006 ), ngakhale madera enieni a PFC omwe akukhudzidwa ndi maphunziro otere amasiyana. Kapangidwe kosiyana ka madera a m'mimba ndi m'mimba kumasonyeza kusamvana pakati pa madera awa, koma sikusonyeza chifukwa cha ubalewu. Ranganath (2006) akupereka lingaliro la kapangidwe ka kayendetsedwe ka ntchito ka kukumbukira komwe madera a caudal/ventral PFC amapereka ulamuliro wapamwamba pansi pa machitidwe a posterior, pomwe dorsal/rostral PFC imapereka ulamuliro wa madera a caudal ventral frontal. M'malingaliro awa, Ranganath akunena kuti njira zosankhira zomwe zimachitika ndi rostral/dorsal PFC zimaphatikizapo kusintha kwa ntchito mu caudal/ventral PFC. Komabe, monga tafotokozera pansipa, kusintha kosiyana kumafunikanso kuganizira.
7. Malangizo okhudzana ndi madera achidziwitso
Popeza chitsanzo cha kapangidwe kake chafotokozedwa m'magawo oyambirira, ziyerekezo za OFC ku PFC yolumikizana, kuphatikiza DLPFC zitha kugawidwa m'magulu monga kupereka ndemanga. Motero, ziyerekezo izi zitha kupereka tsankho ndi kuwongolera madera omwe akukula kwambiri. Ngakhale zikuwoneka kuti zikusiyana ndi malingaliro afilosofi omwe amaika nzeru kuposa malingaliro, lingaliro lakuti gawo lomwe likukhudzidwa ndi kukonza malingaliro lingapereke tsankho la mtundu wa mayankho pamadera omwe akukhudzidwa ndi zochitika zina zamaganizo likugwirizana mosavuta ndi malingaliro amakono amalingaliro omwe amagogomezera kuthekera kwa malingaliro kuyika patsogolo ndikusintha chidziwitso kuti zithandizire zolinga zofunika kwambiri pazamoyo komanso pagulu. Lingaliro ili la malingaliro limafotokozedwa bwino ndi Gray ndi anzake ( Gray, 2001 , Gray, Braver, Raichle, 2002 ), omwe amanena kuti njira ndi machitidwe odzipatula zimakhudza magwiridwe antchito ena azidziwitso, zonse zikukulitsa ndikusokoneza ntchito zosiyanasiyana zamalingaliro kuti zikwaniritse zofunikira pazochitika bwino. Umboni wa kusintha kwa malingaliro koteroko kwa chidziwitso umavomerezedwa bwino popanga zisankho ( Delgado et al., 2003 ; Grabenhorst & Rolls, 2009 ; Hardin, Pine & Ernst, 2009 ; Piech et al., 2010 ), komanso ukhoza kuwoneka m'machitidwe ena oyambira monga kukumbukira ntchito. Mwachitsanzo, magwiridwe antchito a kukumbukira ntchito m'malo ndi mawu amasinthidwa molakwika mwa kuyambitsa mkhalidwe wabwino ndi woipa wa malingaliro, pomwe kukumbukira ntchito m'malo kumakulitsidwa ndi mkhalidwe wamalingaliro wodzipatula ndikuwonongeka ndi momwe zinthu zilili, ndi kukumbukira ntchito pamawu kukuwonetsa zotsatira zosiyana ( Gray, 2001 ). Kuphatikiza apo, chidziwitso chabwino ndi choipa cha malingaliro chimachepetsa kusokoneza kukumbukira ntchito poyerekeza ndi chidziwitso chosalowerera ( Levens & Phelps, 2008 ; 2010 ). OFC yakumanja (33 24 −8) ndi insula yakumanzere yakutsogolo (−32 21 2) zimayankha kwambiri pakuthetsa kusokoneza kwamalingaliro.
Mofananamo, mu ntchito yosinthana maganizo ndi khalidwe, kuyambitsa zotsatira zabwino, poyerekeza ndi zotsatira zosalowerera kapena zoyipa, kunalimbikitsa kusinthasintha kwa chidziwitso ndi kuchepetsa kupirira, komanso kunapangitsa kuti pakhale kusokonezeka maganizo ( Dreisbach & Goschke, 2004 ). Zomwe zapezekazi zikugwirizana ndi umboni wokulirapo wakuti malingaliro abwino ndi oipa amatha kukulitsa kapena kuchepetsa chidwi kutengera mphamvu ya njira kapena mawonekedwe odzipatula a mkhalidwe wa maganizo ( Fredrickson & Branigan, 2005 ; Gable & Harmon-Jones, 2008 ; 2010 ; Gasper, & Clore 2002 ).
Mwachidule, umboni wowonjezereka ukusonyeza kuti zotsatira zolimbikitsa zimakhudza mayankho a BOLD mu DLPFC (BA 9) panthawi ya ntchito yokumbukira ( Gray, Braver, & Raichle, 2002 ; Savine & Braver, 2010 ). Inde, Savine & Braver (2010) akuwonetsa kuti mkati mwa DLPFC yakumanzere (BA 9), zolimbikitsira mphotho ya ndalama zimathandizira makamaka kuyambitsa ntchito zokhudzana ndi cue, ndipo kuyambitsa kumeneku kunaneneratu ngati kuyesa kudzachitika bwino. Ponseponse, maphunziro oterewa amafunika kusintha malingaliro a unidirectional a ubale pakati pa njira zoganizira ndi zamaganizo.
Kafukufuku wa maselo amodzi amapereka chidziwitso china chowonjezera pa nthawi yolumikizirana pakati pa madera a m'mimba ndi madera ambiri a m'mimba, pokhudzana ndi mphotho. Deta yochokera ku ma primate imasonyeza kuti madera ozungulira amalemba kuti apeze kuwerengera koyera kwa mphotho kuposa madera ena akutsogolo, ndipo kuti OFC imapereka chidziwitso ichi ku madera ambiri a m'mimba ( Hikosaka & Watanabe, 2000 ; Wallis & Miller, 2003 ; Rushworth et al, 2005). Chofunika kwambiri, ma neuron a OFC amasonyeza mayankho ku chidziwitso cha mphotho chomwe chisanachitike mayankho okhudzana ndi mphotho mu DLPFC ( Wallis & Miller, 2003 ). Izi zikugwirizana ndi lingaliro lakuti OFC choyamba imalemba mtengo wa mphotho kenako imapatsa chidziwitsochi kumadera omwe angathe kulumikiza chidziwitsochi ndi zochita kapena chidziwitso china chofunikira kuti mupeze mphothoyo. Komabe, tikuzindikira kuti sizikudziwika bwino kuti chidziwitsochi cholimbikitsa chimafikira bwanji DLPFC pankhani ya ma feedback type prospects, kapena chingaganizidwe kuti ndi feedforward mwachilengedwe, chifukwa pafupifupi 30% ya OFC, ma feedback a DLPFC angaganizidwe kuti ndi feedforward mwachilengedwe ( Barbas & Rempel-Clower, 1997 ). Malinga ndi chitsanzo cha kapangidwe kake, kusiyana kumeneku kungatsimikizire ngati kukhudzidwa kwa mphotho kwa maselo a DLPFC kukuwonetsa tsankho la mtundu wa feedback wa DLPFC kapena kuwonetsa kutumiza kosavuta (mtundu wa feedforward) kwa chidziwitso pamtengo womwe DLPFC ingagwire ntchito. Chonde onani Mitchell (2011) kuti muwone momwe magawo a neural a mphotho angagwirizanire ndi omwe amalamulira malingaliro.
Lingaliro lakuti kusintha kwa malingaliro kumakhudza magwiridwe antchito a ubongo lingakhale lothandiza poganizira kulumikizana kwa magwiridwe antchito pakati pa madera a amygdala ndi prefrontal. Monga taonera kale, kulumikizana kwabwino pakati pa madera a PFC (makamaka pregenual cingulate ndi VLPFC) kwawonedwa m'maphunziro am'mbuyomu ( Pezawas et al., 2005 ; Dolcos et al., 2006 ). Tikupereka lingaliro lakuti m'mikhalidwe iyi amygdala ikhoza kukhala yoyambitsa, chifukwa mwina ingawerengere kufunika kwa mkhalidwewo kaye, ndikufotokozera chidziwitsocho kapena kuyesa kuwongolera madera a PFC kutengera chidziwitsocho m'malo mosiyana. Komabe, mpaka pano pali zoyesayesa zochepa zomwe zapangidwa kuti zitsatire njira yolumikizirana kwa magwiridwe antchitoyi.
8. Kukambirana
Tikukhulupirira kuti ndemanga yomwe ili pamwambapa ikuwonetsa kufunika kokhudzana ndi kulumikizidwa kwa PFC ndi ubale wawo ku amygdala poganizira momwe zimakhalira ndi malingaliro. Kulephera kutero kungayambitse zitsanzo zomwe ndizovuta kuyanjanitsa ndi anatomy, motero zimawonetsa kuti sizolondola. Mosiyana ndi izi, chidwi pa tsatanetsatane wa ma neurocircuitry sikuti kungangopereka zitsanzo zabwino za mgwirizano wamalingaliro ndi kuzindikira, komanso kuwululira zinthu zomwe sizikadakhudzidwa.
Kuzindikira kwa Malangizo Akumvera
Kutengera mtundu wosankhidwa wa njira za neuroanatomical pakati pa PFC ndi amygdala, zitsanzo zowoneka bwino za kusinthika kwa PFC pofunikira kuyenera kukhudzana ndi kusinthika, kapena kufotokozeredwa, kakhomo kanyumba kanyumba, chigawo chofikira kufikira gawo la gyrus, kapena kudutsa mbali yakumalo. 47 / 12. Pakadali pano, mawu osavuta akuti PFC ikukhudzana ndi malingaliro amtunduwu imapereka chidziwitso chosakwanira, ndipo nthawi zambiri zitha kukhala zosokoneza, popeza ambiri mwa madera a PFC sakhala ndi zolimba ku amygdala. Kutuluka kwa mitundu yamayendedwe omwe amayang'ana kwambiri ma key osakhazikika ku amygdala, monga zitsanzo zomwe zimayesedwa ndikuyesedwa ndi Wager et al. ndi Phillips et al. ndi gawo lolimbikitsa pankhaniyi. Tikukayikira kuti patsogolo popitilapo kumvetsetsa kutengapo gawo kwa malingaliro a PFC, malingaliro a wachibale wa dorsal anterior cingate, posterior 47 / 12 ndi madera ena pakukhazikitsa amygdala adzafunika kutsimikizika.
Funso lofunikirali likufunsanso momwe kufalikira kwa PFC komwe kumachitika nthawi yamalamulo kumagwirizana ndi mfundozi, popeza ndi owerengeka ochepa okha omwe adawunikira kulumikizana kwa ntchito kwa PFC. Anatomically, madera a PFC awa samalumikizidwa mofananirana ndi dorsal anterior cingate, posterior 47 / 12 kapena dera lachiberekero, chifukwa chake akhoza kuphatikizidwa mosankha ndi njira zosiyanasiyana kupita ku amygdala. Tikukayikira kuti kumvetsetsa kwathu momwe PFC ikukhudzira momwe mungagwiritsire ntchito machitidwe kumafunikira kuwunikira kuchuluka kwa zigawo za PFC zomwe sizimagwirizana ndi zigawo zina za PFC zomwe zimakwaniritsa magwiridwe antchito.
Zindikirani pakuwonekera kwa zoyeserera
Tanena kuti mitundu yayikulu ya ma intra PFC, komanso machitidwe a PFC-amygdala omwe amafotokoza mosadukizadukiza pazomwe zimayenderana pamalingaliro amtunduwu ndiosagwirizana. Mtsutso wathu wotsutsana ndi zitsanzo zamtunduwu za PFC-amygdala komanso okhudzana ndi chipani cha PFC zimadalira kwambiri kapangidwe kofotokozedwa ndi a Barbas ndi anzawo, momwe mawonekedwe amatsenga amatsimikizira ngati zomwe akuyerekezazo zikuyimira tsankho ngati lingaliro, kufotokozera kwatsatanetsatane. Ngati ndi zolondola, madera ena okhudzidwa ndi kutengeka kwawo akuwoneka kuti amapereka njira yolankhulira zapamwamba kuti ingofotokozere zidziwitso kuposa malo achikhalidwe a PFC.
Tikukhulupirira kuti tanthauzo latsopanolo lidayambitsa kubwereza kwatsatanetsatane kumvetsetsa mgwirizano pakati pa zigawo za ubongo ndi njira zamaganizidwe. Kusankhaku kumayenderana ndi malingaliro a malingaliro a magawo a "kuzindikira" ndi "kukhudzika" komwe kumapangitsa kuzindikirika kuposa malingaliro okonda nyama. Koma kusankhaku kungasokoneze kuthekera kwathu kuti timvetsetse bwino momwe ubongo umapangira chidziwitso. Ngati zochitika zam'malingaliro zimayendetsa ndikutsata "mwazidziwitso" machitidwe, mochuluka kapena kuposa njira ina yonse, matchulidwe a pamwamba ndi pansi akhoza kukhala osayenera poganizira momwe zimakhalira.
Zolepheretsa pakukumitsa ntchito pantchito
Kusangalatsa kwa kapangidwe kake ndikuti kumabweretsa maulosi olimba zokhudzana ndi kulumikizana kwa magawo. Komabe, zotsutsa zingapo zimatha kudzutsidwa pokoka magwiridwe antchito molingana ndi mawonekedwe a anatomical. Choyamba, ngakhale mawonekedwe ake amathandizidwa mwamphamvu molingana ndi zonena zake za njira yolumikizira ya laminar yochokera pa kayendedwe ka kayendedwe kazinthu zam'mizinda, zolemba zokhudzana ndi kuthekera kwazinthu zokhudzana ndi mitundu iyi ya laminar yolumikizana sizinalandiridwe mayeso oyendetsedwa m'mabwalo kunja kwa mitsinje yogwiritsira ntchito. Ngakhale zikuwoneka zomveka kuganiza kuti magwiridwe antchito amodzimodziwo amakhala ndi ubongo wolosera, izi siziri choncho ayi. Mwakutero, zokambirana zokhudzana ndi njira yolumikizirana mu PFC ndizovomerezeka ngati magwiridwe antchito amomwe amapangidwira komanso malingaliro ake atatsimikiziridwa kuti azigwira m'ma cortices onse.
Tapereka mgwirizano wamphamvu pakati pa mayankho ogwira ntchito ndi malamulo apamwamba, komanso kulumikizana kolimba pakati pa njira zotsatira komanso zotsika. Mayankho ndemanga ndi chakudya zimachokera ku malingaliro olamulira, omwe amayesera kufotokoza kagwiritsidwe kazinthu zazikulu. Kutengera kwa mawuwa ndi akatswiri a zamaubongo ndi akatswiri azamisala sizodabwitsa chifukwa lingaliro la njira zoperekera mayendedwe owongolera ndi njira zothandizira kuperekera chidziwitso kumadera akutali mumtsinje wokuthandizira ndizabwino. Komabe, kuyesa kosavuta kwa malamulo apamwamba pamwambapa ndi mayankho ndi pansi-mmwamba ndi feed Express kumakhala kovuta mpaka momwe zowonjezera zimasonyezedwera ndi conceptualizations ya pamwamba-pansi ndi pansi. Zowonjezerazi sizimafotokozedwa kawirikawiri, koma zitha kutsimikizira m'njira zowerengera anthu njira. Tikukayikira kuti ena ophunzitsa amagwiritsa ntchito mawuwo pamwamba-pansi ndi pansi m'njira zosagwirizana ndi mayankho komanso njira zina monga zimanenedwera ndi malingaliro, koma zosagwirizana ngati izi sizimafotokozedwa kawirikawiri m'mabuku.
Polemba mayankho komanso zofunikira za PFC, tikuwona kuti sititanthauza kuti zonena zonse ndizofanana. Madera ali ndi kuphatikiza kwa ndemanga, zogwirizana, komanso zolumikizana pambuyo pake, koma kuchuluka kwa kulumikiraku kumasiyana modabwitsa m'malo onse. Chifukwa chake tikuwonetsa makina olumikizana, koma izi sizikutanthauza kuti kulumikizana kwotsala sikofunikira. Mwachitsanzo, zigawo za eulaminate PFC zilidi ndi malingaliro okwanira othandiza kuwongolera madera ocheperako, ngakhale ngati iyi si njira yolankhulirana pakati pa madera.
Kuphatikiza apo, kulumikizana kwa ma feedforward type projections nthawi zina kumatha kusintha momwe zinthu zilili m'madera omwe akufunidwa m'malo mongonyamula chidziwitso. Mwina chitsanzo chabwino kwambiri cha mtundu uwu wa feedforward modulation chimapezeka m'ma model ampikisano ophatikizidwa ( Desimone ndi Duncan 1995 ; Duncan et al. 1997 ) momwe phindu la kuyimira kumodzi kumabweretsa kuletsa kwina. Mu ma model otere, kudyetsa patsogolo kwa kuyimira kwina kungayambitse kukulitsa kwa kukonza kwa chilimbikitso chimenecho, ndikuletsa chilimbikitso china ( Desimone ndi Duncan 1995 ). Mwanjira imeneyi, zomwe zimapatsidwa patsogolo zimatha kusintha momwe zinthu zilili m'madera omwe akufunidwa. Pankhani ya PFC yogwira ntchito, chizindikiro cha DLPFC chikhoza kusintha mpikisano pakati pa ma significant omwe angakhalepo mu OFC kudzera mu mtundu uwu wa feedforward projection. Mtundu uwu wa njira yampikisano ndi wosangalatsa chifukwa ungatanthauze zinthu zinazake zamakompyuta ( Walther & Koch, 2006 ), zomwe sizinaphatikizidwe mu ma model a emotional regulation.
Poganizira za kapangidwe kake, ndikofunikira kubwerezanso kuti miyezo yomwe Barbas ndi anzake amagwiritsa ntchito pofotokoza kulumikizana kwa feedforward ndi feedback sikugwirizana kwathunthu ndi miyezo yomwe ofufuza ena agwiritsa ntchito pofufuza dongosolo la ma projection a laminar. Makamaka, matanthauzidwe a ma feedback ndi ma connection a forward nthawi zambiri amatanthauzidwa ponena za layer IV, kotero kuti ma feedforward (kukwera) projection amatanthauzidwa ndi kutha kwawo mu layer IV (kapena makamaka mu layer IV), pomwe ma feedback (kutsika) projections amatha kunja kwa layer IV. Ngakhale kutsatira kwambiri lamulo la layer IV mwina sikulangizidwa, chifukwa zosiyana ndi izi zawonedwa ( Pandya ndi Rockland, 1979 ; Felleman ndi Van Essen, 1991 ), zotsatira za kukulitsa miyezo kuti zilole ma projection omwe amatha mu infragranular layers V ndi VI kuti awoneke ngati ma feedforward projections sizikumveka bwino. Mosakayikira, maphunziro a nthawi yowombera m'magawo osiyanasiyana a PFC cortical akhoza kuyankha funsoli, koma deta pankhaniyi ikusowa.
Funso la njira zimayimitsa kaye musaganize kuti OFC-DLPFC ilidi ndi njira yomwe OFC iyenera kuchitiridwa ngati yapamwamba kwambiri kuposa DLPFC, ndipo sicholinga chathu kutsutsana ndi izi. Komabe, zitha kunenedwa momveka bwino kuti mitundu yamalingaliro sikufanana ndi gulu lowongolera momwe DLPFC ili m'malo olamulira pamwamba pa OFC, mofanananira ndi malo apamwamba okhala ndi malo okhala. Mwakutero, mitundu ya bungwe la PFC ingakhale yanzeru kupeŵa kufalikira kwa DLPFC ngati kukhala pamwamba pamwambo wa zigawo za PFC.
Kufanizira kwa kulumikizana kwa feedirect ndi mayankho
Pofufuza mitundu yomwe ilipo ya kuyanjana kwa malingaliro ndi kuzindikira, ndizodziwika kuti maphunziro ochepa ofalitsidwa mpaka pano aphatikizapo mayeso enieni ngati ma projection akuwonetsa mayankho, ma feedforward kapena ma lateral projection (kupatula Seminowicz et al. 2004 ). Maphunziro ambiri a neuroimaging sapereka chidziwitso chapadera chomwe chingathetse vutoli. Komabe, chitukuko chaposachedwa cha njira zopangira ma modeling efficient connectivity chimapereka zida zomwe zingagwiritsidwe ntchito kutsanzira mtundu ndi njira yolumikizirana pakati pa madera. Mwachitsanzo, dynamic causal modeling (DCM) pogwiritsa ntchito family level inference ndi Bayesian model average ingagwiritsidwe ntchito poyesa malingaliro okhudzana ndi njira ndi mtundu wa chidziwitso choyenda komanso kusintha kwa causal m'madera osiyanasiyana a ubongo ( Friston et al. 2003 ; Chen et al. 2009 ; Daunizeau et al. 2009; Friston & Dolan 2010 ; Penny et al. 2010 ). DCM ikhozanso kuyesa mitundu yopikisana monga kupereka kufananiza kwa mutu ndi mutu ngati DLPFC imatsitsa amygdala mwachindunji kapena kudzera mu kapangidwe kena kapakati. Mpaka pano, maphunziro ochepa okha a DCM okhudzana ndi kukonza malingaliro ndi omwe afalitsidwa ( Ethofer et al. 2006 ; Smith et al. 2006 ; Rowe et al. 2008 ; Almeida et al. 2009 ), ndipo malinga ndi chidziwitso chathu palibe maphunziro omwe afalitsidwa mwachindunji okhudza kulamulira malingaliro. Komabe, kugwiritsa ntchito njira zotereku kungawonjezere kwambiri kumvetsetsa kwathu kuyanjana kwa malingaliro ndi kuzindikira m'zaka zikubwerazi.
Mayeso achindunji a chikoka
Mwina njira yabwino kwambiri yokhazikitsira ubale pakati pa madera a ubongo ndi kudzera mu kufufuza dera limodzi panthawi yosankha kusintha kwa thupi kapena kutsika kwa dera lina. Mwachitsanzo, ngati DLPFC imagwira ntchito yochepetsera kukonza kwa OFC, munthu angayembekezere mayankho okokomeza mu OFC pamene DLPFC yachotsedwa. Kuthekera kumeneku kungathetsedwe pofufuza ntchito za OFC ndi fMRI mwa odwala omwe ali ndi zilonda za DLPFC. Kapenanso, transcranial magnetic stimulation (TMS) ingagwiritsidwe ntchito pa DLPFC kuti isinthe kwakanthawi mphamvu ya DLPFC pa ntchito za OFC. Inde, Knoch et al. (2006) posachedwapa adanena kuti TMS pamwamba pa DLPFC yakumanja idapanga kusintha kwa ntchito ya posterior OFC mwanjira yodalira pafupipafupi. Mofananamo, zingakhale zosangalatsa kudziwa momwe zilonda zomwe zili mbali imodzi ya prefrontal cortex zimakhudzira kukonza m'malo ena a netiweki. Mwachitsanzo, ngati OFC ndi yofunika powerengera phindu lenileni la mphotho, chimachitika ndi chiyani kumadera ambiri a dorsal pamene OFC yachotsedwa? Saddoris et al. (2005) agwiritsa ntchito njira yamtunduwu kuti aone momwe zilonda za OFC zimasinthira kugunda kwa amygdalar m'makoswe, koma maphunziro ena omwe akugwiritsa ntchito njira iyi ndi osowa kapena palibe. Mabuku omwe akukula okhudza kulumikizana kwa ntchito akuyembekezekanso kuwonjezera kumvetsetsa kwathu momwe madera ofunikira awa amagwirira ntchito. Komabe, kumvetsetsa kwathunthu kwa kuyanjana kumeneku kudzafikiridwa pokhapokha ngati titayang'anitsitsa mawonekedwe enieni a ubongo a ma circuits awa.
Mfundo Zofufuzidwa
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Maulumikizidwe apadera asanachitike amalamula kutanthauzira kwa amygdala
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Makina olimbitsa thupi a Laminar amatsimikizira kuchuluka kwa chidziwitso mu preortal cortex
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Makanema amatsatanetsatane komanso mayankho amakumana ndi zovuta gulu loyambirira
Kuvomereza
Ntchitoyi idathandizidwa ndi mabungwe T32MH018931-21, T32MH018921-20, & 5R01MH074567-04 ochokera ku National Institute of Mental Health. Tikuthokoza a Tawny Richardson chifukwa chothandizira kukonza zolembedwazo.
Mawu a M'munsi
Chodzikanira cha Wofalitsa: Iyi ndi fayilo ya PDF ya cholembedwa chosasinthidwa chomwe chavomerezedwa kuti chifalitsidwe. Monga chithandizo kwa makasitomala athu, tikupereka mtundu woyambirira wa cholembedwacho. Cholembedwacho chidzasinthidwa, kusinthidwa, ndikuwunikanso umboni womwe wapezeka chisanafalitsidwe mu mawonekedwe ake omaliza. Dziwani kuti panthawi yopanga, zolakwika zitha kupezeka zomwe zingakhudze zomwe zili mkati, komanso zoletsa zonse zalamulo zomwe zimagwira ntchito pa magaziniyi.
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