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I-Post-traumatic stress disorder (PTSD) ibonakala ngokukhubazeka ekhonweni lokunciphisa izimpendulo zokwesaba ezinkomba ezihlobene nokuhlukumezeka.Ucwaningo lwabantu nezilwane lukhomba umehluko ekubandakanyekeni kwezindawo ezithile ze-cortex yangaphambili njengabalamuli abayinhloko abanquma ukuphumelela kokunciphisa ukwesaba, kodwa ukusebenzisana kwama-neural circuits anquma ukubandakanyeka okuhlukile kwalezi zindawo akucaci.Ukuze uqonde kangcono ukuthi umehluko ngamunye ekukhunjulweni kokuqothulwa ubonakala kanjani kumehluko womsebenzi wesekethe ye-neuronal, sifake ama-projections asuka ku-rat sublimbic cortex (IL) enama-retrograde tracers futhi saqhathanisa ukuqagela kwe-neuronal ngaphakathi nangaphandle kwe-IL projection neurons.Sihlaziye le datha, sayihlanganisa ngokwezinga lokulondolozwa kwenkumbulo efiphalayo kumagundane.Sithole ukuthi kumaseli e-IL-projecting, ama-neurons ku-posterior parathalamus abonise umsebenzi owengeziwe kumagundane, abonise inkumbulo enhle yokushabalala.Ngokungeziwe kumaseli e-IL-projecting, ukwanda komsebenzi we-Fos kwabonwa ezindaweni ezikhethiwe ze-rat claustrum kanye ne-ventral hippocampus enokulungiswa okuhle.Imiphumela yethu ibonisa ukuthi umehluko ekukhunjulweni kokunyamalala uhlotshaniswa namaphethini athile omsebenzi we-neural ngaphakathi nangaphandle kokuqagela kwe-IL.
Isimo sokwesaba senzeka lapho isikhuthazo sokungathathi hlangothi sihlotshaniswa ne-aversive unconditioned stimulus (UCS), kangangokuthi isisusa sokuqala sokungathathi hlangothi, manje isivuseleli esinesimo (CS), senza impendulo yokwesaba okunesimo (CR) lapho ingekho i-UCS.Ukuguqulwa kokwesaba okunesimo kwaqhutshwa ukwehla kwe-CR kuya ku-CS ngenxa yokwethulwa okuphindaphindiwe kwe-CS ngokungabikho kwe-UCS1.Ucwaningo lwangaphambilini lubonise ukuthi i-post-traumatic stress disorder (PTSD) ihlotshaniswa nokungakwazi ukukhumbula ukushabalala kwezimpendulo zokwesaba okunesimo2.Isisekelo sokwelashwa kokuziphatha kwengqondo yokwelashwa kwe-post-traumatic stress disorder ukwelashwa kokuchayeka okusekelwe ekuqothulweni kwezimpendulo zokwesaba ezifundiwe3,4.Ngakho-ke, izifundo zomehluko ngamunye ekwesabeni kokuqothulwa kwamagundane kanye nezindlela eziyisisekelo ze-neural zingasiza ukucacisa umehluko ezimpendulweni zabantu ekuhlukumezekeni nasekulapheni ukuphazamiseka kwengcindezi yangemva kokuhlukumezeka.Ngaphandle kwenqubekelaphambili ekuhlonzeni izinqubo ze-neural ezihlukanisa ngempumelelo nezinkumbulo zokushabalala ezingaphumelelanga, kuningi okusazotholakala.
Amamodeli wamagundane awusizo kulo msebenzi ngoba kunomehluko omkhulu ngamunye ekukhunjulweni kokushabalala kwamagundane7,8,9,10.Umsebenzi wangaphambilini ophenya izindlela ze-neural zokushabalala kokwesaba ezingeni labantu ubonise ukuthi ukuqaliswa kwe-infralibic cortex (IL) kuyadingeka ukuze kukhunjulwe ukushabalala (refs 11, 12, 13, kodwa bona 14), futhi ezinye izifundo zithole ukuncipha umsebenzi wamagundane ku-IL obonisa inkumbulo embi mayelana nokushabalala uma kuqhathaniswa namagundane, asatshwa kahle.Kodwa-ke, izindlela ama-ILs abandakanyeka ngazo ngokuhlukile ekusizeni ukuqedwa kokwesaba kumagundane uma kuqhathaniswa nalawo abonisa ukushabalala okubuthakathaka azicacile.
Okunye okungenzeka ukuthi umehluko wenkumbulo yokushabalala kokwesaba phakathi kwabantu wumphumela wokwenza kusebenze okuhlukile kwama-IL ahlukile.Ucwaningo lwe-anatomical18 lubonise ukuthi izindawo ezihlukahlukene ze-cortical kanye ne-subcortical yobuchopho zithumela ukuqagela okuminyene ku-IL, okubuye kuthumele ukuqagela okunamandla ezindaweni eziningi zobuchopho.Ucwaningo lwezinga labantu lubonise ukuthi ukuqagela kwe-IL ku-amygdala kubalulekile ekutholeni ukushabalala kokwesaba20,21,22 kanye nokufaka kwe-IL okuvela ku-basolateral amygdala (BLA) nakho kuhlotshaniswa nokufunda kokushabalala.Kunocwaningo oluncane ngokubandakanyeka kwamasekethe agxile ku-IL ekukhunjulweni kokuqothulwa, yize umsebenzi wakamuva uphakamisa ukuthi zombili i-ventral ne-dorsal hippocampus ziyabandakanyeka ekubikezelweni kwe-IL.Ukuqagela okusebenzayo kwe-IL kuya ku-nucleus ehlanganisa kabusha yethalamus, ngokusobala, nakho kubamba iqhaza kwinkumbulo yokuqedwa kokwesaba.
Lezi zifundo zangaphambilini seziqala ukudweba isithombe sokusebenzisana kwamasekhethi e-neural ahilelekile ekukhunjulweni kokuqothulwa, kodwa kunedatha encane kakhulu yokuthi umsebenzi kumasekethe e-neural agxile ku-IL uthonya umehluko ngamunye ekukhunjulweni kokuqothulwa.Lapha, besifuna ukunquma ukuthi ingabe umehluko wenkumbulo yokushabalala kokwesaba phakathi kwabantu uhlotshaniswa nezinguquko ekusebenziseni okokufaka kwe-IL ezindaweni ezithile zobuchopho.Ikakhulukazi, sihlole ukwenziwa kusebenze kwamaseli e-IL afferent kunucleus ye-paraventricular ye-thalamus (PVT), i-clavicle (CLA), i-BLA, ne-ventral hippocampus (vHPC).Lezi zifunda zobuchopho zikhethwe zombili ngenxa yokuthi zithumela ukuqagela okuminyene ku-IL futhi ngenxa yokuthi kunesizathu sokusola ukuthi zingase zihileleke ekuboniseni ukushabalala kokwesaba i-18.Isibonelo, ucwaningo lwakamuva lubonise ukuthi i-PVT, isifunda esaziwa ngokubandakanyeka ekutholeni ukwesaba nasekukhiqizeni kabusha, iyadingeka ukuze kuphinde kuphinde kunyamalale.Ngaphezu kwalokho, izifundo zangaphambilini zibonise ukwanda kwe-basal amygdala kanye nomsebenzi we-vHPC kumagundane aveza inkumbulo yokuqothulwa.Okokugcina, ukuhlaziywa kwe-claustrum kuwukuhlola okwengeziwe njengoba kungekho nowodwa umsebenzi wangaphambili ohlole indima yawo ekuqothulweni.Nokho, umsebenzi wakamuva uphakamisa ukuthi udlala indima esimweni sokwesaba29.
I-Viral GFP-conjugated retrograde tracers yafakwa ku-IL yamagundane ngaphambi kokuhlolwa kokuziphatha, futhi umsebenzi we-Fos kuma-IL afferents ukalwa phakathi nokudlalwa kabusha kokuqothulwa, ukukhumbula ukwesaba, kanye namagundane angazange ahlolwe ukuziphatha.Imiphumela yethu ibonisa ukuthi ukuqagela kusuka ku-thalamus engemuva kwe-paraventricular kuya ku-IL kubonisa umsebenzi owengeziwe kumagundane akhumbula ngempumelelo ukushabalala.Ngokungeziwe ekubikezelweni kwe-IL, umsebenzi we-neural ezindaweni ezithile ze-clavicle kanye ne-ventral hippocampus unyuswe kumagundane ahlehla kahle.Imiphumela yethu ibonisa ukuthi amaphethini omsebenzi we-intrinsic kanye ne-extrinsic neural evezwa ku-IL ahlotshaniswa nomehluko ngamunye kwinkumbulo yokushabalala kokwesaba.
Amagundane angama-Sprague-Dawley amaduna amadala angamashumi amahlanu nane (300-325 g lapho efika) atholwe kwa-Charles River Laboratories (Raleigh, NC) asetshenziswa njengezifundo.Amagundane ahlaliswa ngababili, futhi ukufinyelela mahhala kokudla namanzi, ngomjikelezo wamahora angu-12 wokukhanya/ubumnyama (izibani zikhanyisa ngo-7 ekuseni).Amaqembu amabili amagundane (n = 28 kanye n = 26) asetshenziselwa lezi zivivinyo.Ngemuva kokukhishwa kokufa, iphutha lokuhlinzwa, ukungabi nenkulumo ye-GFP endaweni ehlosiwe, ikhwalithi yezicubu ezimbi, kanye nezinkinga zokuziphatha (kuchazwe ku-Izindlela), iqembu lokukhumbula ukuqothulwa lalihlanganisa amagundane angu-21 kanye neqembu lokukhumbula ukwesaba lalihlanganisa amagundane angu-7, amakheji ezindlu.leli qembu lalinamagundane angu-7 (amagundane angu-35 afakwe ekuhlaziyweni kokugcina).Zonke izinqubo zagunyazwa I-Stony Brook University Institutional Animal Care and Use Committee futhi zazihambisana Nemihlahlandlela YOKUFIKA (https://arriveguidelines.org) kanye Nemihlahlandlela ye-NIH Yokunakekelwa Nokusetshenziswa Kwezilwane Zaselabhorethri.
Amagundane alashwa ezinsukwini ezimbili ngaphambi kokuhlinzwa.Amagundane abulawa izinzwa nge-ketamine (87 mg/kg) kanye ne-xylazine (10 mg/kg), afakwe kumshini we-stereotaxic (i-Stoelting, Woodale, IL) futhi athola imijovo ehlangene ye-AAVrg-CAG-GFP (Addgene, 30) ku-IL.(bhalansi imijovo yesokunxele nesokudla).Ngomjovo, i-cannula ye-gauge engu-22 yehliswa endaweni (AP: + 3.00, ML: ± 0.6, DV: - 5.2).Faka i-cannula yangaphakathi engu-28G (exhunywe kupompo yokufaka ngeshubhu ye-PE 20) ku-cannula engumhlahlandlela ukuze ulethe u-0.6 µl wegciwane ngenani elingu-0.15 µl ngomzuzu futhi uyishiye endaweni imizuzu emi-5 ngemva kokuba ukumnika sekuphelele..Ngemva kokuthunga, amagundane ajovwe nge-meloxicam (1 mg/kg) futhi ngokushesha nje lapho esekwazi ukunyakaza abuyiselwa emakhejini awo.Amagundane agcinwa emakhejini awo cishe amasonto ayi-7 ukuze avumele ukutholakala kwegciwane kanye nokuthutha kabusha.Amagundane amathathu afa ngaphansi kwe-anesthesia, okuholele ekutheni amagundane angu-51 (94%) alulame ngempumelelo ngemva kokuhlinzwa.
Zonke izinqubo zenziwa ku-32 cm × 25 cm × 21 cm conditioning chambers (Clever Systems Inc., Reston, VA) egcinwe ku-45.7 cm × 43.2 cm × 43.2 cm amabhokisi okuzihlukanisa abamba umsindo (Clever Sys. . Inc.).).Ngesikhathi sokufunda ngokushabalala kanye nezikhathi zokukhumbula ukushabalala, umongo washintshwa ukuze wehluke kumongo wokuqala onemibandela.Isimo A (isizukulwane sokwesaba) sasihlanganisa ama-incandescent angu-28-volt, amalambu asendlini (Chicago Micro Lighting, UK), kuyilapho isimo B (ukuqeqeshwa kokushabalala, ukuhlolwa kokukhumbula ukushabalala, nokuhlolwa kokukhumbula ukwesaba) kuhlanganisa nezibani ze-infrared ze-LED (Univivi IR Illuminator, Shenzhen) ., China;U48R).Ngaphezu kwalokho, kuyilapho i-Context A inesitezi esimelene nokudlidliza esinensimbi engagqwali nezindonga ze-plexiglass, Umongo B uqukethe izinsimbi ezipendiwe ezifakwe phansi nasezindongeni.Umumo Wokuqukethwe B uphinde washintshwa ngokubeka ukufakwa kwensimbi egobile okungu-33.5 cm x 21.3 cm egumbini lokubeka isimo elijwayelekile.Ukwengeza, kumongo A amagumbi asulwa nge-acetic acid engu-5%, kuyilapho kumongo B amagumbi asulwa nge-5% ammonium hydroxide.Ekugcineni, esimeni B, amagundane alethwa egumbini lokuhlola ngamabhakede kunokuba agingqwe emakhejini ezinqoleni.Izikhathi zokuziphatha zarekhodwa ngekhamera ephezulu, futhi isignali yevidiyo evela kukhamera ngayinye yafakwa kusofthiwe (FreezeScan 2.00, Clever Sys. Inc., Reston, VA) ehlole ukuziphatha okufiphalayo ngokusekelwe ezinguqukweni zamaphikseli.Amapharamitha akhethiwe ukuze ukuziphatha kokufiphala okulinganiselwe ikhompuyutha kuhambisane eduze nokuziphatha kwezibukeli eziqeqeshiwe ezilinganiselwe mathupha.Inani elibonisa iphesenti lesikhathi sokuqanda esigoqwe ngesikhawu samasekhondi angama-30.
Zonke izinhlelo zokuziphatha zenziwa phakathi nengxenye yokukhanya yomjikelezo wokukhanya/wobumnyama.Amagundane aphathwa izinsuku ezingu-5 ngaphambi kokuqala kwezinqubo zokuziphatha futhi athuthelwa ekamelweni lokuziphatha izinsuku ezintathu zokugcina zokwelashwa.Ngosuku lokuqala lokuhlolwa kokuziphatha, iqembu lamagundane akhumbula ukushabalala lafakwa ku-reflex enesimo sokwesaba, labe selibekwa kumongo A, linikezwe isikhathi sokujwayela esingashukunyiswa yimizuzu engu-6, lase linikezwa zombili izinhlanganisela ze-4 kHz, 76 dB, 30 s. .ithoni nokuqedwa sekukonke, 1.0 mA, kick 1 s (2 amaminithi ITI).Kukho konke ukuqeqeshwa kokuziphatha, amagundane abuyiselwe emakhejini amaminithi angu-2 ngemva kokwethulwa kokugcina kwesikhuthazo.Ngosuku olulandelayo, amagundane avela eqenjini lokukhumbula ukuqothulwa abekwe ekamelweni lokuqukethwe B futhi enza izethulo zomsindo we-20 (i-2 min ITI) njengokuqeqeshwa kokuqothulwa ngemva kwesikhathi se-6 min.Ngosuku olulandelayo, amagundane eqenjini lokuzala lokuqothulwa adalulwe kumathoni angu-4 kumongo B ngemva kwemizuzu engu-6 yokujwayela isikhathi njengokuhlolwa kokuqothulwa.Amagundane eqenjini lenkumbulo yokuqothulwa ayenziwe ama-60 imizuzu ngemva kweseshini yokuziphatha.Iqembu lamagundane alawulayo avusa izinkumbulo zokwesaba angaphansi kwenqubo efanayo ngosuku lokuqala lokusabela okunesimo sokwesaba kumongo A. Ngemva kwamahora angamashumi amane nesishiyagalombili, amagundane abekwe egumbini lomongo B futhi angaphansi kokwethulwa komsindo okungu-4 (2- iminithi ITI) njengesivivinyo sokukhumbula.ukwesaba ngemva kwenkathi yokujwayela eyimizuzu engu-6.Amagundane afakwe amakha emizuzwini engu-60 ngemva kweseshini yokuziphatha.Iqembu lamagundane okulawula asekhaya lahlala emakamelweni awo asekhaya ngesikhathi sonke sokuhlolwa futhi afakwa amakha ngosuku olufanayo namagundane okuhlola.Iqembu ngalinye kulawa amabili amagundane lahlukaniswa laba uchungechunge olubili, futhi inani lezilwane eqenjini ngalinye lalilinganiselwe phakathi kochungechunge.Igundane elilodwa eqenjini lezinkumbulo zokwesaba alizange lifakwe ekuhlaziyweni ngoba alizange libonise izimpawu zokwesaba (ukubanda ngaphansi kuka-15% wesikhathi phakathi nokuhlolwa kwezinkumbulo zokwesaba).Bheka Umfanekiso 2A ukuze uthole umdwebo womugqa wesikhathi wokuziphatha.
Amagundane aqiniswe ngokweqile ngesisombululo se-Fatal Plus (100 mg/kg), abese egcotshwa ngeqhwa elingu-10% PBS elilandelwa yi-formalin ye-buffered engu-10%.Ubuchopho basuswa futhi bagcinwa ku-30% yesisombululo se-sucrose ku-formalin ku-4 ° C cishe isonto elilodwa.Khona-ke ubuchopho buqandisiwe futhi bunqunywe ku-cryostat engu-40 µm obukhulu.Izigaba zigcinwe ngokulandelana ku-10% PBS ku-4°C.Khona-ke, i-immunofluorescence yenziwa ezigabeni ezintantayo mahhala eziqukethe isifunda sobuchopho esithakaselwayo.Izigaba zigezwe izikhathi ezingu-3 ku-10% PBS imizuzu emi-5 ngayinye.Izingxenye zabe sezifukanyelwa ku-5% yokuvinjwa kwe-serum yembuzi amahora angu-2 ekamelweni lokushisa, bese zigezwa izikhathi ezintathu ngaphezulu imizuzu emi-5 ngayinye ngo-10% we-PBS.Izigaba zibe sezifukanyelwa ngobusuku obungu-4° C. kumasosha omzimba ayisisekelo (c-Fos, #2250, 1:500) (I-Cell Signaling, Danvers, MA) exutshwe ngo-1% BSA ku-10% we-PBS.Ngosuku olulandelayo, izigaba zagezwa ngo-10% PBS imizuzu engu-30 ngo-4°C, kwase kuthi izikhathi ezingu-3 imizuzu engu-5 ku-10% PBS futhi zafakwa i-antibody yesibili (i-Alexa Fluor 594 imbuzi elwa nogwaja, i-conjugate ebomvu, 1:500 ).) (I-Invitrogen, Carlsbad, CA) ekamelweni lokushisa amahora angu-2.Ngemva kokugeza oku-3 okwengeziwe ku-10% PBS imizuzu engu-5, izigaba zafakwa kumaslayidi engilazi futhi zavalwa nge-Fluoromount-G (Invitrogen).Bona izithombe ezimele ze-immunostaining ku-Figure 3G.
Isibonakhulu esikhanyayo sisebenzisa ikhamera yedijithali ye-Infinity3 (i-Lumenera, i-Ottawa, i-Ontario, i-Canada) kanye nenjini yokukhanya (i-Lumencor, i-Beaverton, NOMA) exhunywe kusibonakhulu se-Zeiss yasetshenziswa ukuze kutholwe izithombe esifundeni ngasinye sobuchopho bentshisekelo, okuhlanganisa nezigaba eziqukethe. I-IL ngaphandle kwe-immunofluorescence.kwenziwa ukuze kuqinisekiswe ukubekwa kahle kwendawo yomjovo.Izithombe ezisetshenziselwa ukubala amaseli zitholwe ekukhuliseni okungu-20x.Esigabeni ngasinye sethishu, thatha isithombe esisodwa ngesihlungi esivumela ukubonwa kwe-GFP nesithombe esisodwa esinesihlungi esivumela ukubonwa kwe-Alexa Fluor red conjugate ku-antibody yesibili, kanye nesofthiwe yokucabanga (Infinity Analyze, inguqulo 3) yasetshenziselwa isithombe. imbondela.Thola zonke izithombe zazo zonke izindawo zobuchopho usebenzisa isikhathi sokuchayeka futhi uthole izilungiselelo.Amagundane ayisithupha awafakwanga ekuhlaziyweni ngoba ukudluliselwa okukhulu kwaleli gciwane kwenzeka ngaphandle kwe-IL (88% hit rate).Amanye amagundane ayisishiyagalombili awazange afakwe ngoba, naphezu kokuthi leli gciwane lihlasela i-IL, awazange abonise ukuvezwa kwe-GFP okwanele kuzo zonke izifunda zobuchopho ezihlosiwe ezithakaselwayo.Ngaphezu kwalokho, igundane elilodwa alizange lifakwe ngenxa yekhwalithi yezicubu ezimbi.
Lungisa ukukhanya nokugqama ukuze unciphise umsindo ongemuva esithombeni J (NIH) usebenzisa inqubo efanayo yesithombe ngasinye.Izibalo zamaseli engqikithi yamaseli anelebuli wokubuyisela emuva, isamba samaseli anamalebula amabili we-Fos, kanye nengqikithi yamaseli anamalebula amabili enziwe mathupha umhloli, ongazange azihlonze izilwane, esebenzisa ipulagi ye-Image J cytometer.Izibalo zamaseli zenziwe zajwayelekile ukuze zibe amaseli/mm2.Ukuze kuhlaziywe isisho se-Fos kumaseli e-IL-projecting, inani lamaseli anamalebula aphindwe kabili liye lenziwa lajwayelekile ukuze libe inani eliphelele lamaseli anamalebula abuyiselwe emuva.Ngokuhlaziywa kwe-mBLA, mvHPC kanye ne-pvHPC, ukubalwa kwamaseli kusuka ezithombeni eziningi ezingama-20x kwafingqwa futhi kwajwayezwa amaseli/mm2.Ukuze kuhlaziywe zonke ezinye izifunda zobuchopho, isithombe esingu-20x noma ingxenye yesithombe esingu-20x siye sahlaziywa futhi sajwayezwa ukuze sibe amaseli/mm2.Ukuhlaziywa kwe-vHPC kufaka i-CA1, i-CA2 kanye nezifunda ze-vHPC ezingaphansi.Umfanekiso 1 ubonisa izifunda zobuchopho ezihlaziywe ngezithombe ezichaza umngcele ongaphambili-ngemuva wendiza.
Izifinyezo kanye nendawo yezindawo zobuchopho ozithandayo.Incazelo yezifinyezo nezindawo zezifunda zobuchopho ezinikezwe kumbhalo wesandla.Imephu yobuchopho besizinda somphakathi ithathwe ku-Swanson (2004) Imephu Yobuchopho: Isakhiwo Sobuchopho Begundane, Ushicilelo lwesi-3, olunikezwe ilayisense ngaphansi kwe-Creative Commons Attribution-NonCommerce 4.0 International License (https://creativecommons.org/licenses/by-nc/ 4.0) ./), iyatholakala ukuze ilandwe ku-https://larrywswanson.com.
Iphesenti lesikhathi sokufriza lilinganiselwa esikhathini sokudlala ithoni yesibili engu-30, okungafaki izikhawu ezimaphakathi.Izilinganiso zokukhumbula ukushabalala zibalwe ngokuveza iphesenti lesikhathi sokunyamalala phakathi nokukhumbula ukushabalala njengephesenti lokufiphala phakathi nezilingo zokuqala ezi-4 zokuqeqeshwa kokushabalala (ifiphale phakathi namathoni okukhumbula ukushabalala okune phakathi kwamathoni amane okuqala okuqeqeshwa kokushabalala*100).Amaphuzu aphansi akhombisa inkumbulo enhle yokufiphala, futhi amaphuzu aphezulu akhombisa inkumbulo efiphalayo empofu.Amagundane ahlungwe ngomphumela wokukhumbula ukushabalala, namagundane akunani eliphezulu lesithathu lokukhumbula ukushabalala achazwa ngokuthi "amagundane aqothulayo amabi" kanye namagundane engxenyeni yokugcina emibili kwezintathu yemiphumela yokukhumbula ukushabalala achazwe ngokuthi "mahle".Inkumbulo iyafiphala amagundane.
Ukuhlolwa kwe-nonparametric kuyasetshenziswa ngoba idatha ivamise ukwephula ukuqagela mayelana nokusabalalisa okuvamile kanye/noma i-homogeneity yokuhluka.Ukuhlotshaniswa kwezinga lika-Spearman kwasetshenziswa ukuze kutholwe ukuthi ingabe kukhona ukuhlotshaniswa okubalulekile phakathi kwezikolo zokukhumbula ukushabalala nomaka be-Fos nomaka abakabili ezindaweni zobuchopho zentshisekelo kuwo wonke amagundane angaphansi kovivinyo lokukhumbula ukushabalala.I-Mann-Whitney U-test yasetshenziswa ukuze kutholwe ukuthi ngabe ukhona yini umehluko phakathi kwamaqembu amabili azimele.Ukuhlolwa kwe-Kruskal-Wallis kusetshenziselwa ukunquma ukuthi amaqembu amabili noma ngaphezulu ahlukile kwelinye, futhi ukuhlolwa kokuqhathanisa okuningi kwe-Dunn kusetshenziswa uma izibalo ze-Kruskal-Wallis zibalulekile.Ukufiphala ngesikhathi sokufunda kokushabalala kwahlolwa kusetshenziswa ukulinganisa okuphindaphindiwe kokuhlaziywa kokuhluka neqembu njengento ephakathi kwesihloko kanye nokuhlola njengento engaphakathi kwesihloko. Imiphumela ithathwe njengebalulekile lapho i-p <0.05 yazo zonke izivivinyo zezibalo. Imiphumela ithathwe njengebalulekile lapho i-p <0.05 yazo zonke izivivinyo zezibalo. Результаты считались значимыми при p < 0,05 для всех статистических тестов. Imiphumela ithathwe njengebalulekile ku-p <0.05 kukho konke ukuhlola kwezibalo.当所有统计检验的p < 0.05 时，结果被认為是显着的。当所有统计检验的p < 0.05 时，结果被认為是显着的。 Результаты считались значимыми при p < 0,05 для всех статистических тестов. Imiphumela ithathwe njengebalulekile ku-p <0.05 kukho konke ukuhlola kwezibalo.
Umfanekiso we-2 ubonisa umugqa wesikhathi wokuhlola (Umfanekiso 2A) kanye nokusabalalisa imvamisa yawo wonke amagundane angaphansi kokushabalala (Umfanekiso 2B). Amagundane emaqenjini okuhle nokubi okuqothulwa ahluke kakhulu kulawa maphuzu okukhumbula ukushabalala okubaliwe (U = 0, p <0.001) (Fig. 2C). Amagundane emaqenjini okuhle nokubi okuqothulwa ahluke kakhulu kulawa maphuzu okukhumbula ukushabalala okubaliwe (U = 0, p <0.001) (Fig. 2C). Крысы в ​​группах с хорошим и плохим угашением значительно различались по этим рассчитаным показателям припоминания уга,00, 20. Amagundane emaqenjini okuhle nokubi okuqothulwa ahluke kakhulu kulawa mazinga okukhumbula ukushabalala okubaliwe (U=0, p<0.001) (Umfanekiso 2C).在這些计算的灭绝回忆分数中，良好和不良灭绝组中的大鼠存在显着差异（U = 0,p <0.001）C). U = 0,p <0.001) (图2C, В этих рассчитанных показателях припоминания угашения крысы в ​​группах с хорошим и плохим угашением значительно различа, 0.00. Kulawa mazinga okukhumbula ukushabalala okubaliwe, amagundane emaqenjini amahle nampofu ahluke kakhulu (U = 0, p <0.001) (Umfanekiso 2C).Kwakungekho mehluko obalulekile esikhathini sokuqanda phakathi kwamaqembu anokushabalala okuhle, ukushabalala okungekuhle, kanye nokukhumbula ukwesaba phakathi nenkathi eyisisekelo yeseshini ye-reflex enesimo sokwesaba (X2(2) = 2.746, p = 0.253) (Umfanekiso 2D).Ngaphezu kwalokho, phakathi nokwethulwa kwethoni yokuqala ye-reflex enesimo sokwesaba, awubanga khona umehluko obalulekile esikhathini sokuqhwa phakathi kwamaqembu anokushabalala okuhle, ukushabalala okungekuhle kanye nokukhumbula ukwesaba (X2(2) = 1.107, p = 0.575), njengoba kanye nangesikhathi sokwesaba phakathi namathoni wesibili.Phakathi neseshini yokubeka isimo, kube nomehluko omkhulu esikhathini sokuqanda phakathi kwamaqembu anokushabalala okuhle, ukushabalala okungekuhle, kanye nokukhumbula ukwesaba (X2(2) = 2.214, p = 0.331) (Umfanekiso 2D).Kwakungekho nomehluko obalulekile esikhathini sokufiphala phakathi kwamaqembu okuhlela nokubi ngesikhathi sokuqeqeshwa kokushabalala kwesisekelo (U = 45.00, p = 0.799) (Umdwebo 2D). Okulandelayo, kube nomthelela obalulekile obalulekile webhulokhi yesilingo (amathoni angu-5 ibhulokhi ngayinye) ngesikhathi esichithwe iqhwa ngesikhathi seseshini yokuqeqeshwa kokuqothulwa (F (2.884, 54.80) = 8.331, p <0.001), okubonisa ukuthi ukufunda ngokushabalala kwenzeke (Fig. 2D ). Okulandelayo, kube nomthelela obalulekile obalulekile webhulokhi yesilingo (amathoni angu-5 ibhulokhi ngayinye) ngesikhathi esichithwe iqhwa ngesikhathi seseshini yokuqeqeshwa kokuqothulwa (F (2.884, 54.80) = 8.331, p <0.001), okubonisa ukuthi ukufunda ngokushabalala kwenzeke (Fig. 2D ). Затем наблюдался значительный основной эффект пробного блока (5 тонов на блок) на время, затрачиваемое на замирание во вренив, 8,8,8 0) = 8,331, p <0,001), что указывает на то, что обучение угашению происходило ( isiqephu 2D). Kwabe sekuba nomphumela obalulekile obalulekile webhulokhi yesilingo (amathoni angu-5 ngebhulokhi ngayinye) ngesikhathi esithathwe ukuze kumiswe iqhwa ngesikhathi sokuqeqeshwa kokuqothulwa (F(2.884, 54.80) = 8.331, p <0.001), okubonisa ukuthi ukufunda kokuqothulwa kwakwenzeka (Fig . 2D). ).接下來，在消退训练期间，试块（每块5 音）对冻结时间有显着的主效应（F（4.8803（4.8803（4.8803). ，表明发生了消退学习（图2D）） .接下來，在消退训练期间，试块（每块5 音）对冻结时间有显着的主效应（F（4.8803（4.8803（4.8803). ，表明发生了消退学习（图2D）） . Затем, во время обучения угашению, пробные блоки (5 тонов на блок) оказали значительное основное влияние на время замирания,8,8,4 = F 01), waveza ukuthi yini, waveza ukuthi ungubani происходило (рис . 2D). Khona-ke, phakathi nokufunda kokuqothulwa, amabhulokhi wokulinga (amathoni angu-5 ngebhulokhi ngayinye) abe nomthelela omkhulu obalulekile ngesikhathi sokufiphala (F(2.884, 54.80) = 8.331, p <0.001), okubonisa ukuthi ukufunda kokuqothulwa kwakwenzeka (Fig. .2D) .Kodwa-ke, iqembu lokuqothulwa (F(1, 19) = 3.091, p = 0.095) alizange libe nomthelela omkhulu ngesikhathi sokufiphala kuso sonke isikhathi sokuqeqeshwa kokuqothulwa, futhi kwakungekho ukusebenzisana phakathi kwebhulokhi yesilingo neqembu lokuqothulwa (F(4) , 19)).76) = 1.890, p = 0.121) (Fig. 2D). Phakathi neseshini yokuhlola, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, namaqembu okukhumbula ukwesaba ngesikhathi esichithwe iqhwa ngesikhathi sokuqala (X2 (2) = 8.569, p = 0.014) kangangokuthi iqembu lokukhumbula ukwesaba liqhwale kakhulu. ngaphezu kweqembu elihle lokuqothulwa (Mean Rank Diff. = 10.57, p = 0.017), kodwa hhayi iqembu elibi lokuqothulwa (Mean Rank Diff. = - 3.714, p > 0.999) (Fig. 2D). Phakathi neseshini yokuhlola, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, namaqembu okukhumbula ukwesaba ngesikhathi esichithwe iqhwa ngesikhathi sokuqala (X2 (2) = 8.569, p = 0.014) kangangokuthi iqembu lokukhumbula ukwesaba liqhwale kakhulu. ngaphezu kweqembu elihle lokuqothulwa (Mean Rank Diff. = 10.57, p = 0.017), kodwa hhayi iqembu elibi lokuqothulwa (Mean Rank Diff. = - 3.714, p > 0.999) (Fig. 2D).Phakathi neseshini yokuhlola, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, namaqembu okukhumbula ukwesaba ngesikhathi esichithwe iqhwa phakathi nenkathi eyisisekelo (X2(2) = 8.569, p = 0.014), kangangokuthi iqembu lokukhumbula ukwesaba limile. ngokuphawulekayo .больше, чем в группе хорошего вымирания (средняя разница рангов = 10,57, p = 0,017), но не в группе плохого плохого врамирания, 3, 7, p 0,999) (рис. 2D). okukhulu kuneqembu elihle lokuqothulwa (isilinganiso somehluko wezinga = 10.57, p = 0.017) kodwa hhayi eqenjini elibi lokuqothulwa (isilinganiso somehluko wezinga = −3.714, p > 0.999) (Umfanekiso 2D).在测试期间，良好消退组、不良消退组和恐惧回忆组在基线期冻结时间方面存在显兰方面存在显山 X2 (2) = 6. 014），因此恐惧回忆组冻结显着超过良好的灭绝组（平均秩差= 10.57，p = 0.017），但不是差的灭绝组（平均秩差= – 3.714，p > 0.999）（图2D。在 测试 期间 , 良好 消退组 、 消 退组 和恐惧 在 基线期 冻结 时间 方面 方间 方面 方间 方间 方 方 方 方 方 方 方 方 方 方 方 方 方 方 方 方组 存 存在 显 存在 显 2) 569 , p = 0.014） , 恐惧 回忆组 冻结 显着 良好 的 组 组 组 组 组组 组 组（平均秩差= 10.57，p = 0.017），但不是差绝组（平均秩差= – 3.714，p > 0.9999））） В течение периода тестирования наблюдалась значительная разница между группой с хорошим угашением, группой с плохительная разница между хорошим угашением, группой с плохельная плохительная угашением раха с точки зрения времени замирания на исходном уровне (X2 (2) = 8,569, p = 0,014), поэтому припоминание страха группа замерзачает , чем группа с хорошим вымиранием (средняя разница рангов = 10,57, p = 0,017), но не группа с плохим вымиранием вымиранием вымиранием (срезница, 9, 9, 9, 9, 9, 9, 9 ) (isifanekiso 2D). Phakathi nenkathi yokuhlola, kube nomehluko omkhulu phakathi kweqembu elihle lokuqothulwa, iqembu elibi lokuqothulwa, kanye neqembu lokukhumbula ukwesaba ngokwesikhathi sokuqhwa ngesikhathi sokuqala (X2(2) = 8.569, p = 0.014), ngakho-ke ukwesaba ukukhumbula iqembu liqina kaningi kakhulu kuneqembu elishabalala kahle (isilinganiso somehluko wezinga = 10.57, p = 0.017) kodwa hhayi iqembu elinokushabalala okungekuhle (umehluko wezinga elisho -3.714, p > 0.999) (Umfanekiso 2D).Iqembu elihle lokuqothulwa, iqembu elimpofu lokuqothulwa, kanye neqembu lokukhumbula ukwesaba nakho kube nezikhathi ezihluke kakhulu zokufiphala phakathi nokwethulwa kwethoni yeseshini yokuhlola (X2(2) = 14.93, p = 0.001), ngakho iqembu elihle lokuqothulwa libe nokuncane kakhulu isikhathi.isikhathi sokuqandisa kuneqembu lokuqothula elibuthakathaka (umehluko wezinga elisho = 9.286, p = 0.044) kanye neqembu lenkumbulo yokwesaba (umehluko wezinga elisho = 13.86, p = 0.001) (Fig. 2D).
Umehluko ngamunye ekukhunjulweni kokushabalala.(A) Uhlaka lwezinqubo zokuhlinzwa nokuziphatha.(B) Ukusatshalaliswa kwemvamisa okubonisa umehluko ngamunye kuzikolo zenkumbulo ezishabalalayo.(C) Ubufakazi bokuthi amaqembu aklanywe ngokusekelwe kuzikolo ezikhumbulayo zokushabalala ezibaliwe amele ama-phenotype amabili ahlukene.(D) Isilinganiso sephesenti lesikhathi samagundane aba yiqhwa ukuze anyamalale okubi, ukushabalala okuhle, kanye nokukhumbula ukwesaba kumaseli wesibili angu-30 weseshini ye-reflex yokwesaba okunesimo, ngamathoni angu-20, 30 s, agoqeka abe amabhulokhi angu-5 phakathi neseshini yokufunda yokushabalala (amathoni ama-4) .ngayinye), futhi ngamathoni amane ngezikhathi zezinkumbulo ezifiphalayo nezinkumbulo zokwesaba.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo. *p < 0.05, **p < 0.01, ***p < 0.001, ****p < 0.0001. *p < 0.05, **p < 0.01, ***p < 0.001, ****p < 0.0001. *р < 0,05, **р < 0,01, ***р < 0,001, ****р < 0,0001. *p < 0.05, **p < 0.01, ***p < 0.001, ****p < 0.0001. *p <0.05,**p <0.01,***p <0.001,****p <0.0001. *p <0.05,**p <0.01,***p <0.001,****p <0.0001. *р < 0,05, **р < 0,01, ***р < 0,001, ****р < 0,0001. *p < 0.05, **p < 0.01, ***p < 0.001, ****p < 0.0001.
Isikhombi sokubuyisela emuva sajovwa ku-IL (Fig. 3A) kanye nenani lamaseli e-GFP+ eduze kwe-anterior-posterior axis yesifunda sentshisekelo yanqunywa (Fig. 3B-F).Kube nomehluko omkhulu enanini lamaseli e-GFP+ phakathi kwe-PVT yangaphambili, emaphakathi nengemuva (X2(2) = 8.200, p = 0.017), ngakho-ke i-mPVT ibonise ngokuphawulekayo amaseli e-GFP+ amaningi kune-aPVT (isilinganiso sezinga) Diff.= 18.37, p = 0.035) kanye ne-pPVT (isilinganiso sezinga Diff. = 17.71, p = 0.045) (Fig. 3C).Nakuba izilwane ezimbalwa zingazange zithole noma yimaphi amaseli e-GFP+ ku-pCLA futhi ngenxa yalokho azikwazanga ukwenza imephu yomsebenzi kulesi sifunda, awukho umehluko obalulekile phakathi kwe-CLA yangaphambili, emaphakathi, neyangemuva (X2(2) = 5.596, p = 0.061).Inombolo yamaseli e-GFP+ (Umfanekiso 3D).Bese, njengoba amanye amaseli e-GFP+ atholakala ku-aBLA noma ku-avHPC kumagundane amaningana, indawo emaphakathi nengemuva kuphela yalezi zindawo eyahlaziywa.I-BLA emaphakathi nengemuva (U=393, p=0.009) ihluke kakhulu enanini lamaseli e-GFP+, ngakho-ke i-pBLA ibonise ukuqagela kwe-IL okuningi kune-mBLA (Umfanekiso 3E).Ngokufanayo, kube nomehluko omkhulu phakathi kwama-vHPC amaphakathi nangemuva, ngakho-ke ama-pvHPC abonise ukuqagela kwe-IL okuningi kunama-mvHPC (U = 403.5, p = 0.014) (Umfanekiso 3F).Umfanekiso we-3G uyisibonelo sesithombe esibonisa i-Fos, i-aavRG-GFP, namaseli anamalebula amabili.
Linganisa ama-IL afferents kuyo yonke indawo yobuchopho enentshisekelo kuyo.(A) Ukumelwa okuhleliwe kokusatshalaliswa kwe-aavRG-CAG-GFP kulo lonke irat IL.(B) Izithombe ezimele zomaka okubuyisela emuva emuva ezindaweni ezihlukile ze-anteroposterior endaweni yobuchopho oyithandayo.Ukulinganisa inani lokulebula kwe-retrograde ku-anteroposterior axis (C) paraventricular thalamus, (D) clavicle, (E) i-basolateral tonsil, kanye (F) ne-ventral hippocampus.(G) Izithombe ezimelele ezibonisa ukulebula kabusha kwe-aavRG, ukulebula kwe-Fos, kanye nokulebula okuphindwe kabili kwe-aavRG kanye ne-Fos ku-aPVT.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo. *p <0.05, **p <0.01. *p <0.05, **p <0.01. *р < 0,05, **р < 0,01. *p <0.05, **p <0.01. *p <0.05,**p <0.01. *p <0.05,**p <0.01. *р < 0,05, **р < 0,01. *p <0.05, **p <0.01.Ibha yesikali 100 µm.Imephu yobuchopho besizinda esisesidlangalaleni kuphaneli A ikhiqizwe kabusha kusukela ku-Swanson (2004) Imephu Yobuchopho: Isakhiwo Sobuchopho Begundane, Uhlelo lwesi-3, lunikezwe ilayisense ngaphansi kwe-Creative Commons Attribution-NonCommons 4.0 International License (https://creativecommons.org/licenses/by-nc )./4.0/) iyatholakala ukuze ilandwe ku-https://larrywswanson.com.
Umsebenzi we-Fos we-Global kanye ne-IL othize we-Fos wahlaziywa ku-aPVT, mPVT kanye ne-pPVT kuwo wonke amagundane.Kwakungekho mehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya kusisho se-Fos ku-aPVT (X2(3) = 3.888, p = 0.274) (Fig. 4A), futhi kwakungekho ukuhlobana okubalulekile phakathi kwe-Fos ku-aPVT phakathi kwenkulumo nokukhumbula kokushabalala (rs = 0.092, p = 0.691) (Umfanekiso 4B) noma phakathi kwenkulumo ye-Fos kuma-afferents we-PVT IL kanye nokukhumbula kokuqothulwa (rs = 0.143, p = 0.537) (Umfanekiso 4D).Nokho, kuma-afferents e-aPVT IL, isisho se-Fos sihluke kakhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya (X2(3) = 15.05, p = 0.002), ngakho iqembu lokukhumbula ukwesaba libonise ukushabalala okuhle kakhulu.ukuhlehla (isilinganiso somehluko wezinga = 11.54, p = 0.003), ukuhlehla okungekuhle (umehluko wezinga osho = 10.57, p = 0.034), kanye neseli lasekhaya (umehluko wezinga = 12.79, p = 0.005) amaqembu (Fig. 4C).Ngaphezu kwalokho, awukho umehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya okuveza i-Fos ku-mPVT (X2(3) = 2.272, p = 0.518) (Fig. 4E) kanye nokusho kwe-Fos ku-mPVT..Ukuxhumana okubalulekile nokukhumbula kokushabalala (rs = 0.168 p = 0.468) (Umfanekiso 4F).Nakuba bekunomehluko omkhulu phakathi kwamaqoqo amahle, amabi, okukhumbula ukwesaba, namaqembu eseli lasekhaya kusisho se-Fos kumaseli e-IL afferent mPVT (X2(3) = 9.252, p = 0.026), ukuqhathanisa kwe-post hoc akuzange kuvezwe noma yikuphi noma okubili.Umehluko omkhulu phakathi kwamaqembu (Umfanekiso 4G).Ngaphezu kwalokho, kwakungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos kumaseli e-IL-afferent mPVT kanye nokukhumbula kokushabalala (rs = 0.174, p = 0.450) (Umfanekiso 4H). Okulandelayo, kube nomehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya ekukhulumeni kwe-Fos ku-pPVT (X2 (3) = 13.89, p = 0.003), ukuze iqembu elihle lokuqothulwa (Inani Lenani Umehluko = 14.96, p = 0.010), kodwa hhayi ukushabalala okungekuhle (Mean Rank Diff. = 12.86, p = 0.113) noma iqembu lokukhumbula ukwesaba (Mean Rank Diff. = 2.571, p > 0.999), ebonisa ukubonakaliswa kwe-Fos okuningi kune-Fos iqembu lekheji lasekhaya (Fig. 4I). Okulandelayo, kube nomehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya ekukhulumeni kwe-Fos ku-pPVT (X2 (3) = 13.89, p = 0.003), ukuze iqembu elihle lokuqothulwa (Inani Lenani Umehluko = 14.96, p = 0.010), kodwa hhayi ukushabalala okungekuhle (Mean Rank Diff. = 12.86, p = 0.113) noma iqembu lokukhumbula ukwesaba (Mean Rank Diff. = 2.571, p > 0.999), ebonisa ukubonakaliswa kwe-Fos okuningi kune-Fos iqembu lekheji lasekhaya (Fig. 4I). Далее, наблюдалась значительная разница между группами с хорошим угасанием, плохим угашением, отзывом страхай приксай домашение (X2 (3) = 13,89, p = 0,003), так что группа с хорошим угашением (средний ранг Umehluko = 14,96, p = 0,010), но не в группе плохого угашения (средняя ранговая разница = 12,86, p = 0,113) или группого угашения . разница = 2,571, p > 0,999), демонстрировалась более выраженная экспрессия Fos, чем в группе группа домашних клеток (рис. 4I). Ngaphezu kwalokho, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, kanye namaqembu amaseli asekhaya kusisho se-Fos ku-pPVT (X2(3)=13.89, p=0.003), ukuze iqembu elihle lokushabalala (lisho izinga elithi Diff. = 14.96, p = 0.010), kodwa hhayi eqenjini lokushabalala elimpofu (umehluko wezinga osho ukuthini = 12.86, p = 0.113) noma iqembu lenkumbulo yokwesaba (umehluko wezinga elisho = 2.571, p > 0.999), libonise ukuphimisela kwe-Fos okuphimiseleke kakhulu kunaku- iqembu lamaseli asekhaya (Umfanekiso 4I).其次 , ppvt 中 fos 表达 好 消 、 差消 、 恐惧 和 家笼组 之间 存在 显着 差异 （3x2 09） = 3 x 0 9).使得 消组 （(isho i-Rank Difff.= 14.96, p = 0.010)，但不是较差的消退(IMean Rank Diff. = 12.86, p = 0.113) 或恐惧回忆组(IMean Rank Diff. = 2.571, 纭纭廄 廄 廄 廄 9, p 9, p. 4I ）.= 14.96, p = 0.010)，但不是较差的消退(IMean Rank Diff. = 12.86, p = 0.113) ）.Okwesibili, kube nomehluko omkhulu ekukhulumeni kwe-Fos ku-pPVT phakathi kokuhle, okubi, ukukhumbula ukwesaba, namaqembu amaseli asekhaya (X2(3) = 13.89, p = 0.003), okwenza iqembu elihle lokuthatha (kusho umehluko wezinga = 14.96)., p = 0,010), futhi akukho хуже по угашению (средняя разница рангов = 12,86, p = 0,113) noma группе отзыва страха (средня разница , 9, 9, 9 = в группе домашней клетки (рис. 4I) . , p = 0.010), kodwa akukho okubi kakhulu ekuqothulweni (isilinganiso somehluko wezinga = 12.86, p = 0.113) noma iqembu lokukhumbula ukwesaba (umehluko wezinga elisho = 2.571, p > 0.999) kuneqembu lamaseli asekhaya (Umfanekiso 4I)..Nokho, kwakungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-pPVT Fos kanye nokukhumbula ukushabalala (rs = 0.051, p = 0.825) (Umfanekiso 4J).Ekugcineni, kube nomehluko omkhulu ekukhulumeni kwe-Fos kuma-pPVT IL afferents phakathi kwamaqembu anokushabalala okuhle, ukushabalala okungekuhle, izinkumbulo zokwesaba, namaseli asekhaya (X2(3) = 12.34 p = 0.006), ngakho ukusho okuhle kwe-Fos ku-IL- kwaba kubi kakhulu kunasemaqenjini okuqothulwa (umehluko wezinga osho ukuthini = 12.54, p = 0.014) nasesitokisini sasekhaya (umehluko osho isikhundla = 12.89, p = 0.049) (Fig. 4K) futhi uhlotshaniswa kakhulu nama-IL afferents ngaphakathi kwe-pPVT Phakathi kokuqalisa nokusebenza ukuhoxiswa kokushabalala, ukukhunjulwa kokuqothulwa okungcono kwakuhlotshaniswa nokusebenza okukhulu kwalawa ma-afferents e-IL (rs = -0.438, p = 0.047) (Umfanekiso 4L).
Umsebenzi we-Fos unyuswe kuma-IL afferents we-posterior paraventricular thalamus (PVT) kumagundane, abonise ukwehla okuhle.(A) Bekungekho mehluko omkhulu phakathi kwamaqembu ekukhulumeni kwe-Fos ku-APVT.(B) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos kanye nokukhumbula ukushabalala ku-aPVT.(C) Iqembu lokukhumbula ukwesaba libonise ukukhula kwenkulumo ye-Fos kuma-IL afferents uma kuqhathaniswa nawo wonke amanye amaqembu.(D) Bekungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-Fos kuma-IL afferents kanye nokukhumbula ukushabalala ku-aPVT.(E) Bekungekho umehluko obalulekile we-intergroup ekukhulumeni kwe-Fos ku-mPVT.(F) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos nenkumbulo yokushabalala ku-mPVT.(G) Inkulumo ye-Fos kumaseli e-IL ahlukile ku-mPVT ayizange ihluke kakhulu phakathi kwamaqembu.(H) Bekungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-Fos kuma-IL afferents kanye nokukhumbula ukushabalala ku-mPVT.(I) Iqembu elashabalala kahle, kodwa alikho elinye iqembu, elibonise ukwanda komsebenzi we-Fos ku-pPVT uma kuqhathaniswa neqembu lekheji yasekhaya.(J) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos nokukhumbula ukushabalala ku-pPVT.(K) Iqembu elihle lokuqothulwa libonise ukukhula kwenkulumo ye-Fos kumaseli ahlukile we-IL uma kuqhathaniswa neqembu lokuqothulwa elibuthakathaka kanye neqembu lamaseli asekhaya.(L) Kukhona ukuhlobana okubalulekile phakathi kwesisho se-Fos kuma-IL afferents kanye nokukhumbula ukushabalala, ngakho-ke ukukhumbula okuhle kokushabalala kuhlotshaniswa nokusho okukhulu kwe-Fos kuma-IL afferents.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo. *p <0.05, **p <0.01. *p <0.05, **p <0.01. *р < 0,05, **р < 0,01. *p <0.05, **p <0.01. *p <0.05,**p <0.01. *p <0.05,**p <0.01. *р < 0,05, **р < 0,01. *p <0.05, **p <0.01.
Umsebenzi we-Fos womhlaba wonke kanye ne-IL eqondile ku-aCLA kanye ne-mCLA yamagundane wabe usuhlaziywa kuwo wonke amaqembu. Kube khona umehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya kusisho se-Fos ku-aCLA (X2 (3) = 8.455, p = 0.036) kangangokuthi iqembu lokukhumbula ukwesaba (Mean Rank Diff. = 14.50, p = 0.049), kodwa abampofu (Mean Rank Diff. = 10.21, p = 0.373) noma ukushabalala okuhle (Mean Rank Diff. = 4.607, p > 0.999) amaqembu, abonise ukubonakaliswa kwe-Fos okuningi kuneqembu lekheji lasekhaya ( Umfanekiso 5A). Kube khona umehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya kusisho se-Fos ku-aCLA (X2 (3) = 8.455, p = 0.036) kangangokuthi iqembu lokukhumbula ukwesaba (Mean Rank Diff. = 14.50, p = 0.049), kodwa abampofu (Mean Rank Diff. = 10.21, p = 0.373) noma ukushabalala okuhle (Mean Rank Diff. = 4.607, p > 0.999) amaqembu, abonise ukubonakaliswa kwe-Fos okuningi kuneqembu lekheji lasekhaya ( Umfanekiso 5A). Между группами с хорошим угашением, плохим угашением, припоминанием страха и домашними клетками наблюдалась значителнисся значительские значительский ) = 8,455, p = 0,036), kanti futhi группа припоминания страха (среднее ранговое различие = 14,50, p = 0,049), но ни плохая (средняя ранговая разница = 10,21, p = 0,373), ни группа с хорошим вымиранием (средняя ранговая разница = 9,90, 9, 9, 9, 9 ровали большей экспрессии Fos, чем группа в домашней клетке ( Рис 5A). Kube khona umehluko omkhulu kusisho se-aCLA Fos phakathi kokushabalala okuhle, ukushabalala okubi, ukukhumbula ukwesaba namaqembu amaseli asekhaya (X2(3) = 8.455, p = 0.036), ukuze iqembu lokukhumbula ukwesaba (kusho umehluko wezinga = 14.50, p = 0.049), kodwa abampofu (umehluko wezinga osho ukuthini = 10.21, p = 0.373) noma iqembu elihle lokuqothulwa (umehluko wezinga elisho = 4.607, p > 0.999) ubonise inkulumo ye-Fos eminingi kuneqembu leseli yasekhaya (Fig. .5A) . aCLA 中Fos 表达的良好消退、消退差、恐惧回忆和家庭笼组之间存在显着差异(X2 (3) = 槐回忆和家庭笼组之间存在显着差异(X2 (3) =槐 = 8.45,30 = 8.45.忆组(Mean Rank Diff. = 14.50,p = 0.049），但无论是差（平均秩差= 10.21，p = 0.373）还是良好灭绝（平均秩差= 4）都礮= 4）石礮= 4） 9p家庭笼组更多的Fos 表达（图5A) . Acla 中 fos 表达 的 消退 、 消退差 、 回忆 和 家庭 笼组 之间 存在 显着 差异 差异 差异 差异 差异 3 455, p = 0.036) , 因此 恐惧 回忆组 回忆组 回忆组 回忆组 (okusho Isikhundla Ukwahlukana = 14.50 , P = P = P = P = 14. 0.049） , 但 无论是 差 平均 秩差 = 10.21 , p = 0.373） 秩差 = 0.373） 秩差 , p = 0.373差 秩差 = 4.607 , p> 0.999） 组 都 出 比 家庭 笼组 更 多 的 表达 图 图 图 5a) . Была значительная разница между группами с хорошим угашением, плохим угашением, отзывом страха и домашней клеппами клеткой 4, 25, 25, 2014 = 5 = 0,036), поэтому группа отзыва страха (среднее ранговое различие = 14,50) , p = 0,049), но группы с плохим (средняя разница рангов = 10,21, p = 0,373) futhi с хорошим вымиранием (средня разница рангов, 9, 9, 9, p сокую экспрессию Fos, чем группа с домашней клеткой (рис. 5A). Kube khona umehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba kanye namaqembu amaseli asekhaya kusisho se-aCLA Fos (X2(3) = 8.455, p = 0.036), ngakho iqembu lokukhumbula ukwesaba (isho umehluko wezinga = 14.50), p = 0.049 ), kodwa amaqembu ampofu (umehluko wezinga eliphansi = 10.21, p = 0.373) nokushabalala okuhle (umehluko wezinga elisho = 4.607, p > 0.999) abonise isisho se-Fos esiphezulu kuneqembu lamaseli asekhaya (Fig. 5A). .Kwakungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos somhlaba wonke ku-(rs = 0.036, p = 0.876) (Umfanekiso 5B) noma isisho se-Fos kumaseli ahlukile we-IL aCLA (rs = -0.282, p = 0.215) kanye nokukhumbula ukushabalala (Umfanekiso 5B)..5D), awubanga khona nomehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya kusisho se-Fos kuma-aCLA IL afferents (X2(3) = 6.722, p = 0.081) (Umfanekiso 5C)..). Okulandelayo, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya ekukhulumeni kwe-Fos ku-mCLA (X2 (3) = 10.12, p = 0.018) kangangokuthi iqembu elihle lokuqothulwa (Mean Rank Diff . = 12.93, p = 0.038), kodwa akukho ukushabalala okungekuhle (IMean Rank Diff. = 5.143, p > 0.999) noma amaqembu okukhumbula ukwesaba (Mean Rank Diff. = 14.00, p = 0.063) aboniswa ngokuphawulekayo ukubonakaliswa kwe-Fos ku-mCLA isihlobo iqembu yezinyoni ekhaya (Fig. 5E). Okulandelayo, kube nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya ekukhulumeni kwe-Fos ku-mCLA (X2 (3) = 10.12, p = 0.018) kangangokuthi iqembu elihle lokuqothulwa (Mean Rank Diff . = 12.93, p = 0.038), kodwa akukho ukushabalala okungekuhle (IMean Rank Diff. = 5.143, p > 0.999) noma amaqembu okukhumbula ukwesaba (Mean Rank Diff. = 14.00, p = 0.063) aboniswa ngokuphawulekayo ukubonakaliswa kwe-Fos ku-mCLA isihlobo iqembu yezinyoni ekhaya (Fig. 5E). Затем наблюдалась значительная разница между группами с хорошим угашением, воспоминаниями осторошением, воспоминаниями осторошением, воспоминаниями осторошением mCLA (X2 (3) = 10,12, p = 0,018), так что группа с хорошим угашением (средняя разница рангов = 12,93, p = 0,038), futhi ни группы плохого угашения (средня ранговая разница = 5,143, p > 0,999), ни группы зратяпы зрапяпы зразяпысы а = 14,00, p = 0,063) не показали значительно большей экспрессии Fos ku mCLA. Kwabe sekuba nomehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, izinkumbulo zokwesaba kanye namaqembu amaseli asekhaya kusisho se-mCLA Fos (X2(3) = 10.12, p = 0.018), ukuze iqembu elihle lokuqothulwa (isilinganiso sezinga lomehluko = 12.93, p = 0.038), kodwa awekho amaqembu ampofu okushabalala (umehluko wezinga = 5.143, p > 0.999) noma amaqembu okukhumbula ukwesaba (umehluko wezinga olinganiselayo = 14.00, p = 0.063) abonise inkulumo ye-Fos enkulu kakhulu ku-mCLA.uma kuqhathaniswa neqembu lekheji lasekhaya (Umfanekiso 5E).接下來 在 mcla 中 的 fos 表达 , 良好 消退组 、 不良 消 退组 、 恐惧 回忆组 容庭 显组2 (3) = 10.12 , p = 0.018) , 良好 (((((((((()) (((() Mean Rank Diff.= 12.93, p = 0.038) , 但在mCLA 中，弱消退（平均秩差= 5.143,p > 0.999）和恐惧回忆组（平均0（平均0 （平均0) 6未显示出更多的Fos 表达相对于家庭笼组(图5E). = 12.93, p = 0.038) , 在 mcla 中 , 弱消退 （平均 秩差 秩差 = 5.143 , p> 0.999） 秩差 = 0.999） 秩差 ＆ 1 .00 , p = 0.063） 未 显示 出 更 多 的fos 表达 表达 的 fos 表达 表达 的 fos 表达相对于家庭笼组（图5E）. Далее, в экспрессии Fos ku mCLA наблюдалась значительная разница между группой с хорошим угасанием, группой с плоханим угасанием рахе и группой с домашней клеткой (X2(3) = 10,12, p = 0,018), так, группа хорошего угашения (средняя разность рангов = 12,93, p = 0,038), но в mCLA ни слабое угасание (средня разница рангов = 5,143, pы 9рга, pы9р няя разница рангов = 14,00, p = 0,999) = 0,063) показали лучшую экспрессию Много Fos по сравнению с группой с домашней клеткой (Рисунок 5E). Ngaphezu kwalokho, kube nomehluko omkhulu ekukhulumeni kwe-Fos ku-mCLA phakathi kweqembu elihle lokuqothulwa, iqembu elibi lokuqothulwa, iqembu lempendulo yokwesaba, neqembu lamaseli asekhaya (X2(3) = 10.12, p = 0.018), ngaleyo ndlela, okuhle ukushabalala (umehluko wezinga olinganisiwe = 12.93, p = 0.038), kodwa ku-mCLA akukho ukushabalala okubuthakathaka (umehluko wezinga osho ukuthini = 5.143, p > 0.999) noma iqembu lokukhumbula ukwesaba (umehluko wezinga = 14.00, p = 0.999) = 0.063) ubonise kangcono Inkulumo ye-Multi Fos uma iqhathaniswa neqembu lamaseli asekhaya (Umfanekiso 5E).Nokho, isisho se-Fos somhlaba wonke ku-mCLA (rs = 0.321, p = 0.156) (Fig. 5F) noma kumaseli e-IL mCLA ahlukile (rs = -0.121, p = 0.602) kanye nokukhumbula kokunyamalala (Fig. 5H), awukho umehluko obalulekile phakathi amaqembu anokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, kanye neseli yasekhaya yesisho se-Fos kumaseli ahlukile we-IL mCLA (X2(3)=4.923, p=0.178) (Umfanekiso 5G).
Umsebenzi we-Fos uphakanyiswe phakathi ne-claustrum kumagundane anenkumbulo enhle yokushabalala.(A) Iqembu lokukhumbula ukwesaba, kodwa hhayi amanye amaqembu, libonise ukwanda komsebenzi we-Fos uma kuqhathaniswa neqembu lamaseli asekhaya ku-aCLA.(B) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos ku-aCLA kanye nokukhumbula ukushabalala.(C) Inkulumo ye-Fos kumaseli e-IL afferent aCLA ayizange ihluke kakhulu phakathi kwamaqembu.(D) Bekungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-Fos kuma-IL afferents kanye nenkumbulo yokushabalala ku-aCLA.(E) Iqembu elishabalala kahle, kodwa hhayi amanye amaqembu, libonise ukwanda komsebenzi we-Fos ku-mCLA uma kuqhathaniswa neqembu lamaseli asekhaya.(F) Bekungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos nokukhunjulwa kokushabalala ku-mCLA.(G) Inkulumo ye-Fos kumaseli ahlukile we-IL mCLA ayizange ihluke kakhulu phakathi kwamaqembu.(H) Bekungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-Fos kuma-IL afferents kanye nenkumbulo yokuqothulwa ku-mCLA.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo. *p <0.05. *p <0.05. *р <0,05. *p <0.05. *p <0.05. *p <0.05. *р <0,05. *p <0.05.
Ngemva kwalokho, umsebenzi we-Fos womhlaba wonke kanye ne-IL ethile ku-mBLA naku-pBLA wahlaziywa kuwo wonke amaqembu amagundane.Awukho umehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya kusisho se-Fos ku-mBLA (X2(3)=0.944, p=0.815) (Umfanekiso 6A).Awukho futhi umehluko obalulekile phakathi kwamaqembu anokuhlehla okuhle, ukuhlehla okungekuhle, ukukhumbula ukwesaba, kanye nokuvezwa kweseli yasekhaya ye-Fos kumaseli ahlukile we-IL mBLA (X2(3)=0.518, p=0.915) (Umfanekiso 6C).Ngaphezu kwalokho, kwakungekho ukuhlobana okubalulekile phakathi kwenkulumo ye-Fos yomhlaba wonke ku-mBLA (rs = 0.126, p = 0.588) (Umfanekiso 6B) kanye nenkulumo ye-Fos kumaseli ahlukile we-IL mBLA (rs = 0.200, p = 0.385) (rs = 0.200, p = 0.385).p = 0.385).Umfanekiso 6D) kanye nokukhumbula ukushabalala.Kwakungekho nomehluko obalulekile ekuqothulweni okuhle, ekuqothulweni okungekuhle, inkumbulo yokwesaba, namaqembu amaseli asekhaya kusisho se-Fos ku-pBLA (X2(3) = 4.246, p = 0.236) (Fig. 6E), futhi awukho umehluko obalulekile ku-pBLA Kuhle.ukushabalala, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya kusisho se-Fos kumaseli ahlukile we-IL (X2(3)=1.954, p=0.582) (Umfanekiso 6G).Okokugcina, isisho se-Fos somhlaba wonke ku-pBLA (rs = 0.070, p = 0.762) (Fig. 6F) kanye nenkulumo ye-Fos kumaseli ahlukile we-pBLA IL (rs = 0.122, p = 0.597) kanye nokukhumbula ukushabalala (Fig. 6H).
Umehluko ngamunye ekukhiqizeni kabusha kokushabalala awuzange ufanekiswe umehluko ekukhulumeni kwe-Fos ku-amygdala eyisisekelo.(A) Bekungekho umehluko obalulekile we-intergroup ekukhulumeni kwe-Fos ku-mBLA.(B) Bekungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos nokukhunjulwa kokushabalala ku-mBLA.(C) Inkulumo ye-Fos kumaseli ahlukile we-IL mBLA ayizange ihluke kakhulu phakathi kwamaqembu.(D) Bekungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos kumaseli ahlukile we-IL kanye nokukhumbula ukushabalala ku-mBLA.(E) Bekungekho umehluko obalulekile we-intergroup ekukhulumeni kwe-Fos ku-pBLA.(F) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos nokukhumbula ukushabalala ku-pBLA.(G) Isisho se-Fos kumaseli e-IL pBLA ahlukene awazange ahluke kakhulu phakathi kwamaqembu.(H) Bekungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos kumaseli ahlukile we-IL kanye nokukhumbula ukushabalala ku-pBLA.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo.
Ekugcineni, umsebenzi we-Fos womhlaba wonke kanye ne-IL wahlaziywa ku-mvHPC naku-pvHPC kuwo wonke amagundane. Kube khona umehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya kusisho se-Fos ku-mvHPC (X2 (3) = 8.056, p = 0.045) kangangokuthi ukushabalala okuhle (Mean Rank Diff. = 13.29 , p = 0.031), kodwa akukho ukushabalala okungekuhle (Mean Rank Diff. = 6.857, p > 0.999) noma ukwesaba ukukhumbula (Mean Rank Diff. = 8.000, p = 0.864) amaqembu abonise inkulumo ye-Fos eminingi kuneqembu lekheji yasekhaya (Fig. 7A). Kube khona umehluko omkhulu phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu ekheji yasekhaya kusisho se-Fos ku-mvHPC (X2 (3) = 8.056, p = 0.045) kangangokuthi ukushabalala okuhle (Mean Rank Diff. = 13.29 , p = 0.031), kodwa akukho ukushabalala okungekuhle (Mean Rank Diff. = 6.857, p > 0.999) noma ukwesaba ukukhumbula (Mean Rank Diff. = 8.000, p = 0.864) amaqembu abonise inkulumo ye-Fos eminingi kuneqembu lekheji yasekhaya (Fig. 7A). Между группами с хорошим угасанием, плохим угашением, отзывом страха и домашними клетками наблюдалась значительаная 2 = 8,056, p = 0,045), так что хорошее угасание (средняя ранговая разница = 13,29) , p = 0,031), но ни в группах с плохим угасанием (средняя ранговая разница = 6,857, p > 0,999), ни в групппе с отзыям знгося знгям знгям знгям страням знгям страням стран 0, p = 0,864) экспрессия Fos была выше, чем в группе с домашней клеткой (рис. 7А). Kube nomehluko omkhulu ekukhulumeni kwe-mvHPC Fos phakathi kwamaqembu anokushabalala okuhle, ukushabalala okubi, ukukhumbula ukwesaba, namaseli asekhaya (X2(3) = 8.056, p = 0.045), ngakho ukushabalala okuhle (kusho umehluko wezinga = 13.29), p = 0.031), kodwa hhayi eqenjini lokushabalala elimpofu (isilinganiso somehluko wezinga = 6.857, p > 0.999) noma eqenjini lokukhumbula ukwesaba (umehluko wezinga elisho = 8.000, p = 0.864) ubungaguquki be-Fos obuphezulu kuneqembu lasekhaya.iseli (Fig. 7A). I-MVHPC 中 FOS 表达 的 良好 消退, 不良不良, 恐惧恐惧 和回忆 家庭家庭 存在笼组之间存在6家庭笼组更多的Fos 表达 (图2). mvhpc 中 fos 表达 的 消退 、 不良 消退 恐惧 回忆 和 家庭 笼组 之间 存在 显着 显退 巈退 差异 60 = 30 = 60 = 60 = 60 = 60 = 60 ） , 良好 ((平均 秩差 秩差 = 13.29） p = 0.031) , 但 无论是 弱消退 （平均 秩差 秩差 = 6.857 , p> 0.999） 还是 恐惧 （平均 秩差 = 0 6 p. ） 组都 出 家庭 笼组 多 的 fos 表达 图 图。。. ））））））））））））））））））））））））））））））) Имелась значительная разница между группами «хорошо», «плохо», «припоминание страха» и «домашняя клетка» для экспрессии 3PC, 8(v 6,80 = 60, 80 = 5 PC, 80 futhi, следовательно, хорошая регрессия (средняя разница рангов = 13,29), p = 0,031), но группы со слабым угасанием (средняя разница рангов = 6,857, p > 0,999) и воспоминания о страха, 80 = 0,857 = 0,999 ,864) показали более высокую экспрессию Fos, чем группа в домашней клетке (рис. 2). Kube nomehluko obalulekile phakathi kokuhle, okubi, ukukhumbula ukwesaba, namaqembu amaseli asekhaya ekukhulumeni kwe-Fos ku-mvHPC (X2(3) = 8.056, p = 0.045) ngakho-ke ukwehla okuhle (isho umehluko wezinga = 13.29), p = 0.031), kodwa amaqembu anokushabalala okubuthakathaka (umehluko wezinga osho ukuthini = 6.857, p > 0.999) nezinkumbulo zokwesaba (umehluko wezinga elisho = 8.000, p = 0.864) abonise isisho se-Fos esiphezulu kuneqembu ekhejini lasekhaya (Fig. 2).7A).Kodwa-ke, kwakungekho umehluko ophawulekayo (X2 (3) = 4.893, p = 0.180) (Umfanekiso 7C).Ngaphezu kwalokho, kwakungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos somhlaba wonke ku-mvHPC (rs = -0.233, p = 0.309) (Umfanekiso 7B) kanye nesisho se-Fos kumaseli e-IL ahlukile we-mvHPC (rs = 0.056, p = 0.810) (Umfanekiso 7D) .kanye nombiko ngokunyamalala.Ngaphezu kwalokho, awukho umehluko obalulekile phakathi kokushabalala okuhle, ukushabalala okungekuhle, ukukhumbula ukwesaba, namaqembu amaseli asekhaya kusisho se-Fos ku-pvHPC (X2(3) = 3.623, p = 0.353) (Umfanekiso 7E), futhi kwakungekho mehluko obalulekile.umehluko ekuhlehleni okuhle kwesisho se-Fos kumaseli e-IL ahlukile we-pvHPC, ukuhlehla okungekuhle, inkumbulo yokwesaba, namaqembu amaseli asekhaya (X2(3)=3.871, p=0.276) (Fig. 7G).Ekugcineni, akuzange kube khona ukuhlobana okubalulekile phakathi kwe-pvHPC global Fos expression (rs = −0.127, p = 0.584) (Figure 7F) kanye nenkulumo ye-Fos kumaseli e-IL-afferent pvHPC (rs = 0.176, p = 0.447) kanye nenkumbulo yokuqedwa (Umfanekiso 7F ).7H).
I-Fos expression iphakeme ku-ventral hippocampus yamagundane, okubonisa ukushabalala okuhle kwenkumbulo.(A) Iqembu elishabalala kahle, kodwa hhayi amanye amaqembu, libonise ukubonakaliswa okukhulisiwe kwe-Fos ku-mvHPC uma kuqhathaniswa neqembu lamaseli asekhaya.(B) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos kanye nokukhumbula kokushabalala ku-mvPHC.(C) Isisho se-Fos kumaseli e-IL ahlukile we-mvHPC awazange ahluke kakhulu phakathi kwamaqembu.(D) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos kuma-IL afferents kanye nokukhumbula kokushabalala ku-mvHPC.(E) Bekungekho mehluko obalulekile phakathi kwamaqembu kusisho se-Fos ku-pvHPC.(F) Akukho ukuhlobana okubalulekile phakathi kwesisho se-Fos nokukhumbula ukushabalala ku-pvHPC.(G) Isisho se-Fos kumaseli e-IL ahlukile we-pVHPC awazange ahluke kakhulu phakathi kwamaqembu.(H) Bekungekho ukuhlobana okubalulekile phakathi kwesisho se-Fos kuma-IL afferents kanye nokukhumbula ukushabalala ku-pvHPC.Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo. *p <0.05. *p <0.05. *р <0,05. *p <0.05. *p <0.05. *p <0.05. *р <0,05. *p <0.05.Ukuhlaziywa kwethu okuyinhloko kwazo zonke izifunda kubonise ukuqhathanisa kumaleveli amathathu ku-axis ye-anteroposterior, nakuba siphinde sahlaziya isifunda ngasinye esigoqeke eduze kwe-axis ye-anteroposterior.Imiphumela yalokhu kuhlaziya yethulwe kuthebula loku-1.
Lapha sihlole ukuthi ingabe umehluko ngamunye ekukhunjulweni kokunyamalala ungabonakala yini kumaphethini ahlukene omsebenzi ohlukile ku-limbic cortex engezansi.Kuze kube manje, sihlole umsebenzi we-Fos ekuqageleni kwe-IL kusukela ku-thalamus ye-paraventricular, i-claustrum, i-basolateral tonsil, ne-ventral hippocampus ngemva kokudlala kabusha kokunyamalala.Kumaseli e-IL-projecting, sithole umsebenzi ophezulu endaweni engemuva ye-PVT kumagundane abonise inkumbulo enhle yokushabalala uma kuqhathaniswa namagundane anokushabalala okubi.Kwakungekho mehluko kuma-IL afferents kusuka ku-clavicular nucleus, ventral hippocampus, noma i-basolateral tonsil.Ngokungeziwe kumaseli e-IL-projecting, ukwanda komsebenzi we-neural kwabonwa ezindaweni ezikhethiwe ze-rat claustrum kanye ne-ventral hippocampus enesinqumo esihle.Imiphumela yethu ikhombisa ukuthi inkumbulo yokushabalala eyimpumelelo ihlelwa ukuqagela kwe-PVT okuqondile ku-IL namaseli angewona ama-IL-targeting ku-claustrum kanye ne-ventral hippocampus.
Sithole ukuthi ukubikezela kwe-PVT IL bekusebenza kumagundane abonisa ukukhumbula okuhle kokushabalala, okuhambisana nocwaningo lwakamuva olubonisa ukuthi i-PVT iyadingeka ukuze kukhunjulwe ukushabalala.Lolu cwaningo aluzange lusebenzise ukukhohlisa okuqondene nendawo ethile, kodwa lubonise ukuthi kokubili ukuqagela kwe-PVT ku-lateral central amygdala kanye nokuqagela kwe-IL ku-PVT kwakudingeka ukukhiqiza kabusha ukushabalala.Imiphumela yethu ibonisa ukuthi, ngaphezu kochungechunge lwe-IL-PVT-CeL, ukungena kwe-post-PVT ku-IL kungase kudingeke futhi ukuze kucishwe ukukhumbula.Ngakho-ke, kuvela ukuthi kokubili ukuxhumana okusebenzayo nokuhambisanayo kwe-IL kuyabandakanyeka ekukhiqizeni kabusha kokushabalala.Isinyathelo esilandelayo esibalulekile ukunquma ukuthi yini ebangela ukuthi i-pPVT ibonise ukuzala kabusha ezingeni le-neural circuit.Ngaphezu kobudlelwane ne-IL, izifundo zangaphambilini zokulandela umkhondo31,32 zibonise ukuthi i-pPVT ithola okokufaka okuvela ku-ventral periaqueductal grey (vPAG), ehlotshaniswa nokufunda kokuqothulwa33,34,35,36.Nakuba indima ye-vPAG ekukhunjulweni kokuqothulwa ingakasungulwa, ukubikezelwa kwe-pPVT nge-vPAG kuyikhandidethi elikhangayo ngenxa yokuminyana kanye nokubandakanyeka kwazo zombili izifunda ekukhiqizeni ubufakazi bangaphambilini bokushabalala kokwesaba.
Esinye isici esibalulekile semiphumela yethu ye-PVT ukuthi ikhethekile ngokuhambisana ne-anterior-posterior axis yayo.Ngokumangalisayo, umsebenzi we-neuronal wokuqagela kwe-PVT ku-IL uhlotshaniswa nesimo sokuziphatha esiphambene, njengokuthi umsebenzi wokuqagela wangaphambi kwe-PVT ku-IL uhlotshaniswa nokukhumbula ukwesaba, kuyilapho ukuqagela kwe-pPVT kusebenza ngemva kokuqedwa okuphumelelayo kokukhumbula (okungukuthi, ukwesaba).Lokhu kuhlukahluka okusebenzayo ngaphakathi kwe-PVT akumangazi uma kunikezwe umsebenzi wangaphambilini [okuxoxwe ngakho ku-37].Isibonelo esimangalisayo sokusatshalaliswa okusebenzayo ku-PVT sisanda kuvela ocwaningweni olubonise izakhiwo zezinhlobo ezithile zamaseli ku-PVT.Lolu cwaningo lukhombisa ukuthi amaseli e-dopamine aveza i-DRD2 avezwa kakhulu ku-pPVT, awangenisi i-forelimb cortex, futhi aphendule ku-aversive stimu.Isibalo samangqamuzana esibili sivezwa kakhulu ku-aPVT futhi siphawula ukushintshela esimweni sokuvuka okuphansi komzimba futhi kwenza i-cortex yezitho ezingezansi ingabonakali.Imiphumela yethu ayingeni nhlobo kule phethini, njengoba amaseli e-PVT akhiqiza i-IL esebenza ngesikhathi sokukhumbula ukwesaba, kuyilapho ukuqagela kwe-pPVT kusebenza futhi izilwane zibonisa amazinga aphansi okwesaba.Okungenani kunezincazelo ezimbili ezingaba khona zokwehluka okusobala.Okokuqala, izinhlobo zamaseli ezikhonjiwe azitholakali kuphela endaweni eyodwa yangaphambili-ngemuva ye-TVV.Ngakho, amaseli e-pPVT asebenzayo e-IL-projecting kumagundane anenkumbulo enhle yokushabalala angase abe ekilasini lamaseli okungenzeka ukuthi atholwe ku-aPVT futhi abonise ukushintshela esimweni esiphansi sokuvuka.Okufanayo kungase kube yiqiniso kumaseli a-IL-projecting ku-PVT enziwe asebenza ngemva kwenkumbulo yokwesaba.Okwesibili, izifundo zokulandelela zangaphambilini zithole ubukhona be-IL3-projecting pPVTs, nakuba ezinye zibonakala ziphuma kumaseli aqukethe i-DRD2, ezinye izinhlobo zamaseli zingase zivele ku-IL futhi zenziwe zisebenze ekukhiqizeni kabusha okuphumelelayo kokucisha.
Nakuba inhloso yalolu cwaningo bekuwukubona umehluko phakathi kwamagundane abonisa ama-phenotypes ahlukene okuqothulwa, lezi zivivinyo ziphinde zembule idatha entsha ehlobene nezindlela zenkumbulo yokwesaba.Kuyathakazelisa ukuthi sithole ukwanda komsebenzi we-Fos endaweni yangaphambili ye-CLA kumagundane anenkumbulo yokwesaba.
I-clavicle ibekwe njengesikhungo sokuxhumana kwe-cortical futhi ihileleke ezinkambisweni ezivela ekuhlanganisweni kwezinzwa kuya ekunakekelweni nasekulaleni40,41,42,43.Kunobufakazi obulinganiselwe bokuthi i-claustrum ihileleka kanjani esimweni sokwesaba noma ekuboniseni ukwesaba, nokho, ucwaningo lwangaphambili lubonise ukuthi ukubonakaliswa kokwesaba okungokomongo kuhileleke kumsebenzi we-Fos ku-claustrum.Muva nje kubikwe ukuthi ukuvinjelwa kwe-atresia projections ku-entorhinal cortex ngesikhathi sokwesaba isimo somongo kuphazamisa ukwakheka kwenkumbulo yesikhathi eside, nakuba isidingo sabo senkulumo yokwesaba singazange sihlolwe.Ocwaningweni olufanayo, ukwenziwa kusebenze okwengeziwe kwe-Fos kubonwe lapho izilwane zivezwe endaweni entsha uma kuqhathaniswa namagundane achayeke endaweni ejwayelekile.Unalokhu engqondweni, ukwenza kusebenze kwe-CLA esikubika lapha kungase kubangelwe ukuchayeka kukhamera entsha phakathi nokuhlolwa, esikhundleni sokwesaba ukuzikhumbula.Ukuze ukhombise ngokunembe kakhudlwana umsebenzi wezingidi ngokwesaba nokucutshungulwa kwesimo, izifundo zesikhathi esizayo kufanele zisebenzise ukukhohlisa kokukhiya okuhlosiwe.
Nakuba umsebenzi wangaphambilini ubonise ukuthi i-PVT ihlotshaniswa nenkulumo yenkumbulo yokwesaba, i-45,46,47 asizange sibone noma yiluphi ushintsho ekukhulumeni kwe-Fos okuphelele kumagundane lapho ekhumbula ukwesaba amahora angu-48 ngemva kokumiswa.Lo mehluko ungachazwa izici ezimbalwa, okuhlanganisa ukwesaba kokuhlolwa komsebenzi wangaphambilini kwezimpawu ezisobala endaweni efanayo lapho ukumiswa okwenzeke khona, kanti ekuhloleni kwethu, ukuhlolwa kwenziwa ekamelweni elisha.Ngaphezu kwalokho, sanikeza izilwane zethu amandla emizuzu engama-60 ngemuva kokuhlolwa, kanti umsebenzi wangaphambilini wasebenzisa iphuzu lesikhathi lemizuzu engama-90.Ekugcineni, ezifundweni zangaphambilini, ukuhlolwa kwenziwa ekamelweni lapho izilwane zingaphendula khona ngokudla, kanti emsebenzini wethu, amagundane ahlolwe ngaphandle kokusabela kwesifiso sokudla.Nakuba lokhu kuvumela izinga elithile lokuvinjelwa okunesimo, kunobufakazi bokuthi ukuvumela izilwane ukuthi zivimbe ingcindezi yokuthola ukudla ngenkathi zihlola ukuthi ziyesaba yini izimpawu kudala ukungqubuzana okushukumisayo (okungukuthi, ukwesaba ngokumelene nomvuzo), okuyisici esiyinhloko esishukumisayo.ukubamba iqhaza PVT48, 49. .
I-amygdala eyisisekelo yaziwa ngokubandakanyeka ekutholweni kokushabalala kokwesaba50,51 futhi kunobufakazi bokuthi ukuqagela kwe-BLA ku-IL nakho kuyabandakanyeka kule nqubo23.Kodwa-ke, akucaci ukuthi i-BLA kanye nokuxhumana kwayo kuyabandakanyeka yini ekubuyiselweni kokushabalala.Izifundo zokucabanga23,28 zibonise ukwanda kwe-BLA ezilwaneni ezikhumbula izinkumbulo eziphelile.Nakuba umsebenzi wethu wangaphambilini ungazange ubonise mehluko ekusebenzeni kwe-BLA phakathi kwamagundane okuhle nokubi okuqothulwa, imiphumela yethu lapha iphakamisa ukuthi ukukhumbula ukushabalala ngokuvamile ayithinti i-BLA noma ukwenziwa kusebenze ekubikezeleni i-BLA IL.Ngokuvumelana nokutholakele kwethu, nakuba ucwaningo lokukhohlisa lwesekethe luphakamisa ukuthi okokufaka kwe-IL ku-BLA kubalulekile ekufundeni kokushabalala, akudingekile ukuze kukhunjulwe ukushabalala.Kodwa-ke, indima ye-BLA ayinakushaywa indiva ngokuphelele njengoba ubufakazi bakamuva buphakamisa ukuthi izinhlobo ezithile zamaseli ku-BLA ziyadingeka ukuze zikhiqize kabusha ukushabalala.
Ngokuphawulekayo, ukukhumbula ukwesaba akuzange kubangele ukuthi i-Fos isebenze ku-BLA, njengoba isilonda sangaphambilini, izidakamizwa, nezifundo zokucabanga ziye zafaka le ndawo ekukhulumeni kokwesaba kanye/noma ekuhlanganisweni kabusha kokwesaba ngemva kokubuyisa54,55,56,57.Idatha evezwe lapha ihlanganisa i-basal ne-lateral subnucleus ye-amygdala, futhi idatha yangaphambilini iphakamisa ukuthi inkulumo yokwesaba iqhuba umsebenzi we-Fos ohlangothini lwe-dorsal nucleus engemuva.Sihlaziye idatha ewumsuka neseceleni ngokuhlukana, kodwa awukho umehluko kuzo zombili izimo (idatha ayiboniswanga) futhi zombili izindawo zigoqekile kudatha esiyethula lapha.Asizange sihlaziye izifunda ezingaphansi kwe-amygdala ehlangothini, ngakho-ke izinguquko ezithile kule ndawo zingafihlwa.Okunye okungenzeka ukuthi akukho shintsho kumsebenzi we-Fos ku-BLA kungenxa yesikhathi sezinkumbulo zokwesaba uma kuqhathaniswa nesimo.Umsebenzi othile wangaphambilini ubonise ukuthi umnikelo we-BLA wokwesaba inkulumo wehla ngokuhamba kwesikhathi ngemva kokumiswa, okufana nokuthi leyo nkulumo incike ku-BLA emahoreni angu-24 ekufakweni kwesimo kodwa ezimele ngezinsuku ezingu-7 (ref. 45 kodwa bheka 58).kwenzeke emahoreni angu-48 ngemva kokuqeqeshwa, okwenza ukuntuleka koshintsho kumsebenzi we-Fos kuleli phuzu cishe kubonise izinguquko ezincike esikhathini ekuhlanganyeleni kwe-BLA ekuboniseni ukwesaba.
Okokugcina, sithola ubufakazi bokuthi inkumbulo yokuqedwa ngempumelelo ihlotshaniswa ne-ventral hippocampus.Lokhu kuyisici sama-vHPC "aphakathi" njengoba iphethini efanayo ingazange ibonwe esifundeni esingemuva.Ngokuvumelana nomsebenzi wangaphambilini, asitholanga ushintsho ekwenzeni kusebenze i-Fos kuma-vHPC IL ahlukile.Kunobufakazi obuningi bokuthi i-vHPC28,60,61 iyadingeka ukuze kwenziwe ukwesaba lapho i-CS yenzeka ngaphandle komongo lapho ukufiphala kwenzeka khona, nokuthi lokhu kuncike okungenani ngokwengxenye ekungeneni kwe-vHPC ku-IL13.Ngokusekelwe kule miphumela yangaphambilini, singalindela ukufiphala okubi ukuthi kuhlotshaniswe nokwenyuka komsebenzi we-IL obikezelwe we-vHPC.Nokho, lokhu bekungenjalo njengoba ungekho umehluko kumsebenzi we-Fos kuma-vHPC ahloselwe i-IL ehloselwe ukubuyisela emuva noma amaseli angenalebuli kuma-vHPC.Lokhu kuphakamisa ukuthi ukungakwazi ukukhumbula ukufiphala kumongo wokufiphala kungase kuqalise indlela ehlukile kunokwesaba ukuvuselelwa.
Kubalulekile ukuqaphela eminye yemikhawulo engokwemvelo yokwakheka nokuhlaziya nokuthi izithinta kanjani iziphetho zethu.Okokuqala, sihlukanise izilwane zaba izingxenye ezintathu ezingaphezulu neziphansi, namagundane aba “okuhle” kanye “nokubi” okusekelwe kuzikolo ezikhumbula ukushabalala.Lokhu kwenziwa ukuze kugwenywe izinhlelo zeqembu ezihlukanisa izilwane zibe amaqembu ahlukene kusukela phakathi nokusabalalisa, noma izinhlelo zeqembu ezingafaki izilwane phakathi nendawo yokusabalalisa, njengokuhlukaniswa nge-median noma ukuqhathanisa ingxenye yesithathu yamagundane angaphezulu naphansi. .sifuna ukugwema lesi simo ngoba ukuhlukaniswa okuphakathi akubonisi ukuhlukahluka kwezimpendulo zomuntu ekuhlukumezeni esizama ukukulingisa.Ukwengeza, ngenkathi kuqhathaniswa nengxenye yesithathu yamagundane angaphezulu naphansi kusivumela ukuba siqhathanise amaqembu anobukhulu obufanayo, le ndlela ayizinaki izilwane phakathi nendawo yokusabalalisa futhi ayibonisi ngokunembile ukuhlukahluka ekuphenduleni kokulimala.Nakuba indlela yethu ingase ihlupheke ngenxa yezinkinga ngokuhlukahluka okuhlukahlukene nokuqhathanisa amaqembu anamasampuli angalingani osayizi, ithwebula esizama ukukulingisa kangcono kunezindlela ezihlukile.
Imiphumela evezwe lapha isisiza ukuthi siqonde kangcono ukuthi umehluko ngamunye ekukhunjweni kokushabalala ubonakala kanjani emehlukweni wokujikeleza kwe-neural.Okutholakele kwethu kungase kuhambisane nokuphazamiseka kokucindezeleka kwangemva kokuthuthumela, okwaziwayo ukuthi kuhlotshaniswa nokwesaba okukhulu kanye nokungakwazi ukuqeda izimpendulo zokwesaba.Sibonisa ukuthi umehluko ekukhunjulweni kokushabalala uhlotshaniswa nomehluko emsebenzini wangaphakathi kanye ne-extrinsic neural oklanyelwe ku-IL.Lo mehluko usatshalaliswe ezifundeni ezihlukene eduze kwe-axis ye-anteroposterior, kuqhubeke kugqamisa ukubaluleka kokuhlola ukusebenza kobuchopho ezingeni lesifunda.Ububi bendlela yamanje buhlanganisa ukuhambisana kocwaningo nokugxila kumagundane abesilisa.Ucwaningo lwesikhathi esizayo kufanele lunqume izindlela ze-neurobiological eziyisisekelo zokufunda ngokushabalala kumagundane abesifazane futhi lusebenzise izindlela ukudweba okucatshangwayo okuyimbangela.
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