Enkosi ngokundwendwela i-Nature.com.Inguqulelo yesikhangeli oyisebenzisayo inenkxaso enyiniweyo yeCSS.Ngowona mava angcono, sicebisa ukuba usebenzise isikhangeli esihlaziyiweyo (okanye uvale iModi yokuThelela kwi-Internet Explorer).Okwangoku, ukuqinisekisa inkxaso eqhubekayo, siya kunika isayithi ngaphandle kwezitayela kunye neJavaScript.
Ingxaki yoxinzelelo lwasemva kokwenzakala (PTSD) luphawulwa ngokuphazamiseka kokukwazi ukuthomalalisa iimpendulo zoloyiko kwiimpawu ezinxulumene nokwenzakala.Uphando lwabantu kunye nezilwanyana lubonisa ukungafani kokubandakanyeka kweendawo ezithile ze-cortex yangaphambili njengabalamli abaphambili abamisela impumelelo yokunciphisa ukwesaba, kodwa ukudibanisa kwee-neural circuits ezimisela ukubandakanyeka kokwahlukana kwezi ndawo akucaci.Ukuqonda ngcono indlela umahluko ngamnye ekukhunjulweni kokutshabalala kubonakaliswe kumahluko kumsebenzi wesekethe ye-neuronal, sibhale iingqikelelo ezivela kwi-rat sublimbic cortex (IL) kunye ne-retrograde tracers kunye nokuthelekisa i-neuronal projection ngaphakathi nangaphandle kwe-IL ye-projection neurons.Sihlalutye ezi datha, sizibeke ngokwamaqela ngokwenqanaba lokugcinwa kwememori ephelayo kwiimpuku.Sifumene ukuba kwiiseli ze-IL-projecting, i-neurons kwi-posterior parathalamus ibonise umsebenzi owandisiweyo kwiigundane, ezibonise ukukhumbula kakuhle ukuphela.Ukongeza kwiiseli ze-IL-projecting, ukwanda komsebenzi we-Fos kwabonwa kwiindawo ezikhethiweyo ze-rat claustrum kunye ne-ventral hippocampus enesisombululo esihle.Iziphumo zethu zibonisa ukuba iyantlukwano ekukhunjulweni kokutshabalala kunxulunyaniswa neepateni ezithile zomsebenzi we-neural ngaphakathi nangaphandle koqikelelo lwe-IL.
Imeko yoloyiko lwenzeka xa i-stimulus engathathi hlangothi ihambelana ne-aversive unconditioned stimulus (UCS), efana ne-original neutral stimulus, ngoku i-conditioned stimulus (CS), ikhupha impendulo yokwesaba okusemgangathweni (CR) ngokungabikho kwe-UCS.Ukuguqulwa kokwesaba okusemgangathweni kuqhutywe kukuhla kwe-CR ukuya kwi-CS ngenxa yokunikezelwa ngokuphindaphindiweyo kwe-CS ngokungabikho kwe-UCS1.Uphando lwangaphambili luye lwabonisa ukuba i-post-traumatic disorder disorder (PTSD) ihambelana nokungakwazi ukukhumbula ukupheliswa kweempendulo zoloyiko olunemeko2.Ilitye lembombo lonyango lokuziphatha kwengqondo kunyango lwe-post-traumatic stress disorder lunyango lwe-exposure therapy esekelwe ekuphelisweni kweempendulo zoloyiko ezifundiweyo3,4.Ke ngoko, izifundo zomahluko ngamnye kuloyiko lokutshabalala kweempuku kunye neendlela ezisezantsi ze-neural zinokunceda ukucacisa umahluko kwiimpendulo zabantu kumothuko kunye nonyango loxinzelelo lwasemva koxinzelelo.Ngaphandle kwenkqubela phambili ekuchongeni iindlela ze-neural ezahlula impumelelo kwiinkumbulo zokutshabalala ezingaphumelelanga, kuninzi okuza kufunyanwa.
Imifuziselo yeempuku iluncedo kulo msebenzi kuba kukho umahluko obalulekileyo womntu ngamnye ekukhunjulweni kokuphela kweempuku7,8,9,10.Umsebenzi wangaphambili ophanda iindlela ze-neural zokuphela koloyiko kwinqanaba labantu ubonise ukuba ukusebenza kwe-infralibic cortex (IL) kuyafuneka ukukhumbula ukuphelelwa (i-refs 11, 12, 13, kodwa bona i-14), kwaye ezinye izifundo zifumene ukuncipha umsebenzi weempuku kwi-IL ebonisa inkumbulo embi malunga nokuphela xa kuthelekiswa neempuku, ezoyikwa kakuhle.Nangona kunjalo, iindlela apho ii-ILs zibandakanyeka ngokwahlukileyo ekuququzeleleni ukupheliswa koloyiko kwiimpuku xa kuthelekiswa nezo zibonisa ukuphela okubuthathaka azicacanga.
Enye into enokwenzeka kukuba iyantlukwano kwinkumbulo yokuphela koloyiko phakathi kwabantu sisiphumo sokusebenza ngokwahlukileyo kwee-ILs ezithile.Izifundo ze-Anatomical18 zibonise ukuba iindawo ezahlukeneyo ze-cortical kunye ne-subcortical yengqondo zithumela iingqikelelo ezixineneyo kwi-IL, ethi ithumele uqikelelo olusebenzayo kwiindawo ezininzi zobuchopho.Uphononongo lwenqanaba labemi lubonise ukuba uqikelelo lwe-IL kwi-amygdala lubalulekile ekufumaneni ukutshatyalaliswa koloyiko20,21,22 kunye negalelo le-IL kwi-amygdala ye-basolateral (BLA) nayo inxulumene nokufunda kokuphela.Kukho uphando oluncinci malunga nokubandakanyeka kweesekethe ezigxile kwi-IL ekukhunjulweni kokutshabalala, nangona umsebenzi wamva nje ucebisa ukuba zombini i-ventral kunye ne-dorsal hippocampus ibandakanyeka kwi-IL yokuqikelela.Uqikelelo olusebenzayo lwe-IL ukuya kwi-nucleus edibeneyo yethalamus, ngokucacileyo, ikwathatha inxaxheba kwinkumbulo yokuphela koloyiko.
Ezi zifundo zangaphambili ziqala ukupeyinta umfanekiso wokusebenzisana kwee-neural circuits ezibandakanyekayo ekukhunjulweni kokutshabalala, kodwa kukho idatha encinane kakhulu malunga nokuba umsebenzi we-IL-centered neural circuits uphembelela ukungafani komntu ngamnye ekukhunjulweni kokuphela.Apha, siye safuna ukufumanisa ukuba ingaba umahluko kwimemori yokuphela koloyiko phakathi kwabantu inxulumene notshintsho kwi-IL yokufaka igalelo kwimimandla ethile yobuchopho.Ngokukodwa, siye savavanya ukusetyenziswa kweeseli ze-IL ezidibeneyo kwi-nucleus ye-paraventricular ye-thalamus (PVT), i-clavicle (CLA), i-BLA, kunye ne-ventral hippocampus (vHPC).Le mimandla yobuchopho ikhethwe zombini ngenxa yokuba ithumela iingqikelelo ezixineneyo kwi-IL kwaye ngenxa yokuba kukho isizathu sokukrokra ukuba banokubandakanyeka ekuboniseni ukwesaba ukuphela kwe-18.Ngokomzekelo, uhlolisiso lwakutshanje lubonise ukuba i-PVT, ummandla owaziwa ngokuba nenxaxheba ekufumaneni uloyiko nokuvelisa kwakhona, iyafuneka ukuze kupheliswe imveliso.Ukongezelela, izifundo zangaphambili zibonise ukunyuka kwe-basal amygdala kunye nomsebenzi we-vHPC kwiigundane ezibonisa imemori yokuphela.Ekugqibeleni, uhlalutyo lwe-claustrum luhlolisise ngakumbi ukuba akukho msebenzi wangaphambili oye wavavanya indima yawo ekuphelisweni.Nangona kunjalo, umsebenzi wamva nje ucebisa ukuba idlala indima kwimeko yoloyiko29.
I-Viral GFP-conjugated retrograde tracers yafakwa kwi-IL yeempuku ngaphambi kovavanyo lokuziphatha, kwaye umsebenzi we-Fos kwi-IL afferents walinganiswa ngexesha lokuphinda kuphele, ukukhunjulwa koloyiko, kunye neempuku ezingafakwanga kuvavanyo lokuziphatha.Iziphumo zethu zibonisa ukuba uqikelelo olusuka kwi-posterior paraventricular thalamus ukuya kwi-IL lubonisa ukwanda komsebenzi kwiigundane ezikhumbula ngempumelelo ukuphela.Ukongeza kwiingqikelelo ze-IL, umsebenzi we-neural kwiindawo ezithile ze-clavicle kunye ne-ventral hippocampus yonyuswa kwiigundane eziye zabuyela kakuhle.Iziphumo zethu zibonisa ukuba iipatheni ze-intrinsic kunye ne-extrinsic neural activity eqikelelweyo kwi-IL inyanyaniswa nomahluko womntu ngamnye kwinkumbulo yokuphela koloyiko.
Amashumi amahlanu anesine angamadoda eempuku ze-Sprague-Dawley (300-325 g ekufikeni) ezifunyenwe kwi-Charles River Laboratories (Raleigh, NC) zisetyenziswe njengezifundo.Iigundane zahlaliswa ngamabini, kunye nokufikelela kwamahhala ekutyeni kunye namanzi, kwi-12 yeyure ukukhanya / umjikelezo omnyama (izibane kwi-7 am).Amaqela amabini eegundane (n = 28 kunye n = 26) asetyenziselwa ezi zilingo.Emva kokukhutshwa kokufa, impazamo yotyando, ukungabikho kokubonakaliswa kwe-GFP kwindawo ekujoliswe kuyo, umgangatho ombi we-tissue, kunye neengxaki zokuziphatha (ezichazwe kwiiNdlela), iqela lokukhumbula ukutshatyalaliswa libandakanya iigundane ze-21 kunye neqela lokukhumbula ukwesaba libandakanya iirathi ze-7, iikheji zendlu.iqela liqulethwe ngamagundane e-7 (iigundane ze-35 zifakwe kuhlalutyo lokugqibela).Zonke iinkqubo zavunywa yiStony Brook University Institutional Animal Care and Use Committee kwaye zihambelana nezikhokelo zokufika (https://arriveguidelines.org) kunye neZikhokelo ze-NIH zokuKhathalela nokuSetyenziswa kweZilwanyana zaseLabhoratri.
Iimpuku zanyangwa kwiintsuku ezimbini phambi kotyando.Amagundane ayenziwe i-anesthetized nge-ketamine (87 mg / kg) kunye ne-xylazine (10 mg / kg), efakwe kwi-stereotaxic apparatus (Stoelting, Woodale, IL) kwaye ifumene i-injection ye-unilateral ye-AAVrg-CAG-GFP (Addgene, 30) kwi-IL.(balance left and right inaliti).I-injection, i-cannula ye-22-gauge yathotywa kwindawo (AP: + 3.00, ML: ± 0.6, DV: - 5.2).Faka i-28G ye-cannula yangaphakathi (edityaniswe kwimpompo yokufakwa nge-PE 20 yetyhubhu) kwi-cannula yesikhokelo ukuhambisa i-0.6 µl yentsholongwane ngesantya se-0.15 µl ngomzuzu kwaye uyishiye endaweni yemizuzu emi-5 emva kokuba ukufakwa kuphelile..Emva kwe-suturing, iigundane zafakwa nge-meloxicam (1 mg / kg) kwaye ngokukhawuleza nje ukuba zikwazi ukuhambisa zabuyiselwa kwiindawo zazo.Iimpuku zagcinwa kwiindlwana zazo malunga neeveki ezisi-7 ukuvumela ukubuyiswa kwentsholongwane kunye nokuthutha ukubuyisela umva.Iigundane ezintathu zafa phantsi kwe-anesthesia, okubangele ukuba iigundane ezingama-51 (94%) zibuyele ngempumelelo kuqhaqho.
Zonke iinkqubo zenziwa kwi-32 cm × 25 cm × 21 cm amagumbi okulungisa imeko (Clever Systems Inc., Reston, VA) egcinwe kwi-45.7 cm × 43.2 cm × 43.2 cm iibhokisi zokuzihlukanisa ezivakalayo (Clever Sys. . Inc.).).Ngexesha leeseshoni zokufunda nokukhumbula ukuphela kokuphela, imeko yatshintshwa ukuze yahluke kumxholo wemeko yantlandlolo.Imeko A (isizukulwana soyiko) sasiquka i-28-volt incandescent, ii-bulbs zasekhaya (i-Chicago Micro Lighting, e-UK), ngelixa imeko B (uqeqesho lokuphela, uvavanyo lokukhumbula ukutshabalalisa, kunye novavanyo lokukhumbula ukwesaba) luquka izibane ze-infrared ze-LED (I-Univivi IR Illuminator, i-Shenzhen) ., eTshayina;U48R).Ukongeza, ngelixa i-Context A ine-anti-vibration slatted floor with steel stainless and plexiglass wall, Umxholo B uqulethe iintsimbi ezipeyintiweyo ezifakwe emgangathweni nasezindongeni.Imilo yoMxholo B nayo yatshintshwa ngokubeka intsimbi eyi-33.5 cm x 21.3 cm egobileyo kwigumbi lokulungisa eliqhelekileyo.Ukongeza, kumxholo A amagumbi asulwa nge-5% ye-acetic acid, ngelixa kumxholo B amagumbi asulwa nge-5% ye-ammonium hydroxide.Ekugqibeleni, kumxholo B, iimpuku zazingeniswa kwigumbi lovavanyo ngee-emele kunokuba zijikelezwe ngeekheji kwiinqwelo.Iiseshoni zokuziphatha zarekhodwa ngekhamera ephezulu, kwaye isibonakaliso sevidiyo esivela kwikhamera nganye yondliwa kwisofthiwe (FreezeScan 2.00, Clever Sys. Inc., Reston, VA) evavanya ukuziphatha okufiphalayo ngokusekelwe kwiinguqu zepixel.Iiparamitha zikhethwe ukwenzela ukuba ukuziphatha kokuphela koqikelelo yikhompyuter kuhambelane ngokusondeleyo nokuziphatha kwababukeli abaqeqeshiweyo abaqikelelwa ngesandla.Ixabiso elibonisa ipesenti yexesha lomkhenkce elisongwe kwisithuba semizuzwana engama-30.
Zonke iinkqubo zokuziphatha zenziwa ngexesha lenxalenye yokukhanya komjikelezo wokukhanya / omnyama.Iigundane zaphathwa ngeentsuku ze-5 ngaphambi kokuqala kweenkqubo zokuziphatha kwaye zithunyelwa kwigumbi lokuziphatha kwiintsuku ezintathu zokugqibela zonyango.Ngosuku lokuqala lovavanyo lokuziphatha, iqela leempuku zokukhumbula ukutshabalala zafakwa kwi-reflex eyoyikayo, emva koko yabekwa kumxholo A, inikwe i-6 min engavuswanga ixesha lokuqhelanisa, kwaye emva koko inikwe zombini indibaniselwano ye-4 kHz, i-76 dB, i-30 s. .ithoni kunye nokupheliswa ngokubanzi, i-1.0 mA, ukukhaba i-1 s (2 min ITI).Kuzo zonke iziqendu zokuziphatha, iigundane zabuyiselwa kwii-cages 2 min emva kokunikezelwa kokugqibela kwe-stimulus.Ngomso olandelayo, iigundane ezivela kwiqela le-extinction-recall-recall zibekwe kwigumbi lomxholo B kwaye zenze iintetho ze-20 zesandi (i-2 min ITI) njengoqeqesho lokuphela emva kwexesha le-6 min.Ngomso olandelayo, iigundane kwiqela le-extinction reproduction zivezwe kwiitoni ze-4 kumxholo B emva kwexesha le-6-minute acclimatization njengovavanyo lokuphela.Iigundane kwiqela lememori yokucima zixutywe imizuzu engama-60 emva kweseshoni yokuziphatha.Iqela leempuku zokulawula ezikhupha iinkumbulo zoloyiko ziphantsi kwenkqubo efanayo ngosuku lokuqala lwempendulo elungiselelwe uloyiko kumxholo A. Kwiiyure ezingamashumi amane anesibhozo kamva, iigundane zafakwa kwigumbi lomxholo B kwaye zixhomekeke kwiintetho ze-4 zomsindo (2- umzuzu ITI) njengovavanyo lokukhumbula.uloyiko emva kwexesha le-6 lemizuzu yokuqhelanisa.Iigundane zixutywe imizuzu engama-60 emva kweseshoni yokuziphatha.Iqela leempuku zolawulo lwasekhaya lahlala kwiindlwana zabo zasekhaya kulo lonke uvavanyo kwaye zaxutywa kwangolo suku olunye njengeempuku zokulinga.Iqela ngalinye kumaqela amabini eempuku lahlulahlulwe laziintlobo ezimbini, kwaye inani lezilwanyana kwiqela ngalinye lalilinganiswe phakathi koluhlu.Enye impuku kwiqela leenkumbulo zoloyiko yayingabandakanywanga kuhlalutyo ngenxa yokuba ayizange ibonise iimpawu zokoyika imeko (umkhenkce ongaphantsi kwe-15% yexesha ngexesha lovavanyo lweenkumbulo zoloyiko).Jonga uMfanekiso 2A ngomzobo wexesha lokuziphatha.
Iimpuku zaye zagqithiswa ngesisombululo se-Fatal Plus (100 mg / kg), emva koko i-perfused nge-ice-cold 10% ye-PBS ilandelwa yi-10% ye-formalin ye-buffered.Ingqondo isusiwe kwaye igcinwe kwi-30% isisombululo se-sucrose kwi-formalin kwi-4 ° C malunga neveki ye-1.Emva koko ubuchopho buye bakhenkcezwa kwaye basikwa bubunzima obungama-40 µm.Amacandelo agcinwe ngokulandelelana kwi-10% ye-PBS kwi-4 ° C.Emva koko, i-immunofluorescence yenziwa kumacandelo akhululekileyo adadayo aqulethe ummandla wobuchopho onomdla.Amacandelo ahlanjwe ngamaxesha e-3 kwi-10% ye-PBS kwi-5 min nganye.Amacandelo afakwe kwisisombululo se-5% sokuthintela se-serum yebhokhwe eqhelekileyo kwiiyure ezi-2 kwiqondo lokushisa eliqhelekileyo, kwaye emva koko ahlambe amaxesha amathathu angaphezulu kwimizuzu emi-5 nganye kwi-10% ye-PBS.Amacandelo ke afakwe ngobusuku kwi-4 ° C. kwii-antibodies eziphambili (c-Fos, #2250, 1:500) (i-Cell Signaling, i-Danvers, MA) ixutywe kwi-1% ye-BSA kwi-10% ye-PBS.Ngomso olandelayo, amacandelo ahlanjwe kwi-10% ye-PBS ye-30 min kwi-4 ° C, ngoko-3 amaxesha kwi-5 min kwi-10% ye-PBS kwaye ifakwe kwi-antibody yesibini (i-Alexa Fluor 594 ibhokhwe echasene nomvundla, i-conjugate ebomvu, i-1: 500 ).) (I-Invitrogen, iCarlsbad, CA) kwindawo yokushisa kweeyure ze-2.Emva kokuhlamba okongeziweyo kwe-3 kwi-10% ye-PBS kwimizuzu emi-5, amacandelo afakwe kwiislayidi zeglasi kwaye atywinwe ngeFluoromount-G (Invitrogen).Jonga imifanekiso emele ye-immunostaining kwi-Figure 3G.
I-microscope ene-fluorescent isebenzisa i-Infinity3 ikhamera yedijithali (i-Lumenera, i-Ottawa, i-Ontario, i-Canada) kunye ne-injini yokukhanya (i-Lumencor, i-Beaverton, OR) eqhagamshelwe kwi-microscope ye-Zeiss yayisetyenziselwa ukufumana imifanekiso kwingingqi nganye yengqondo yomdla, kubandakanywa namacandelo aqulethe I-IL ngaphandle kwe-immunofluorescence.yenziwe ukuqinisekisa ukubekwa okuchanekileyo kwendawo yokutofa.Imifanekiso esetyenziselwa ukubala iiseli ifunyenwe kwi-20x yokukhulisa.Kwicandelo ngalinye lezicubu, thatha umfanekiso omnye ngesihluzo esivumela ukubonwa kwe-GFP kunye nomfanekiso omnye kunye nesihluzi esivumela ukubonwa kwe-Alexa Fluor ebomvu ye-conjugate kwi-antibody yesibini, kunye nesofthiwe yokucinga (Infinity Analyze, version 3) yayisetyenziselwa umfanekiso. ukwaleka.Fumana yonke imifanekiso yazo zonke iindawo zobuchopho usebenzisa ixesha lokuvezwa okufanayo kwaye ufumane useto.Iigundane ezintandathu zazingabandakanywa kuhlalutyo ngenxa yokuba ukuhanjiswa kwentsholongwane kwenzeka ngaphandle kwe-IL (i-88% yezinga lokubetha).Ezinye iimpuku ezisibhozo azizange zibandakanywe ngenxa yokuba, ngaphandle kokuba intsholongwane ihlasele i-IL, ayizange ibonise ukubonakaliswa okwaneleyo kwe-GFP kuyo yonke imimandla yobuchopho ekujoliswe kuyo enomdla.Ukongeza, impuku enye yayingabandakanywanga ngenxa yomgangatho ophantsi weethishu.
Lungisa ukukhanya kunye nokuchasana nokunciphisa ingxolo yangasemva kumfanekiso J (NIH) usebenzisa inkqubo efanayo kumfanekiso ngamnye.Ubalo lweeseli zeeseli ezibhalwe ngokutsha zizonke, iiseli ezibhalwe ngeFos zizonke, kunye neeseli ezibhalwe kabini zizonke zenziwe ngesandla ngumfuni, ongakhange azichonge izilwanyana, esebenzisa iplug-in ye-Image J cytometer.Ubalo lweeseli luye lwaqhelaniswa neeseli/mm2.Ukuhlalutya imbonakalo ye-Fos kwiiseli ze-IL-projecting, inani leeseli ezibhalwe kabini zaye zenziwa zesiqhelo ukuya kwitotali yenani leeseli ezilebhile ngokutsha.Kwi-mBLA, i-mvHPC kunye nohlalutyo lwe-pvHPC, ukubalwa kweeseli ukusuka kwimifanekiso emininzi ye-20x yashwankathelwa kwaye iqhelekile kwiiseli / mm2.Uhlalutyo lweminye imimandla yobuchopho, umfanekiso we-20x okanye inxalenye yomfanekiso we-20x yahlalutywa kwaye iqhelekile kwiiseli / mm2.Uhlalutyo lwe-vHPC lubandakanya i-CA1, i-CA2 kunye nemimandla engaphantsi kwe-vHPC.Umzobo we-1 ubonisa imimandla yobuchopho ehlalutywe ngemifanekiso echaza umda wangaphambili-ngasemva kwendiza.
Izifinyezo kunye nendawo yemimandla yobuchopho enomdla.Ingcaciso yezifinyezo kunye neendawo zemimandla yobuchopho enikwe kumbhalo-ngqangi.Imephu yengqondo yedomeyini kawonke-wonke ethathwe kwi-Swanson (2004) Imephu yoBuchopho: Isakhiwo se-Rat Brain, uHlelo lwesi-3, olunikwe ilayisenisi phantsi kwe-Creative Commons Attribution-NonCommercial 4.0 International License (https://creativecommons.org/licenses/by-nc/ 4.0) ./), iyafumaneka ukuze ukhutshelwe apha https://larrywswanson.com.
Ipesenteji yexesha lomkhenkce lilinganiselwa kwi-30 yethowuni yesibini yexesha lokudlala, ngaphandle kwezithuba eziphakathi.Amazinga okukhumbula ukutshabalala aye abalwa ngokuvakalisa ipesenti yexesha lokuphela ngexesha lokukhumbula ukucinywa njengepesenti yokufiphala ngexesha leemvavanyo zoqeqesho zokuphela kokuphela kwe-4 (ukuphela ngexesha lokukhumbula iithowuni ezine zokuphela / ukucima ngexesha leethowuni zoqeqesho ezine zokuphela *100).Amanqaku asezantsi abonisa inkumbulo entle yobumnyama, kwaye amanqaku aphezulu abonisa inkumbulo ebuthathaka.Iimpuku zahlelwa ngenqaku lokukhumbula ukutshabalala, kunye neempuku kwindawo yesithathu ephezulu yokukhumbula amanqaku achazwe "njengeempuku ezimbi zokutshabalala" kunye neempuku kwisibini esithathwini sokugqibela sokukhumbula amanqaku achazwe "njengelungileyo".Iimpuku eziphelayo kwinkumbulo.
Uvavanyo lwe-nonparametric lusetyenziswa ngenxa yokuba idatha ihlala iphula iingcamango malunga nokusabalalisa okuqhelekileyo kunye / okanye i-homogeneity yokuhluka.Ulungelelwaniso lwenqanaba lika-Spearman lusetyenziselwe ukufumanisa ukuba kukho unxulumano olubalulekileyo phakathi kwamanqaku okukhumbula ukutshabalala kunye namanqaku e-Fos kunye nabamakishi abambini kwimimandla yobuchopho enomdla kuzo zonke iimpuku eziphantsi kovavanyo lokukhumbula ukutshabalala.I-Mann-Whitney U-test yasetyenziselwa ukufumanisa ukuba kukho umahluko phakathi kwamaqela amabini azimeleyo.Uvavanyo lwe-Kruskal-Wallis lusetyenziselwa ukufumanisa ukuba amaqela ama-2 okanye ngaphezulu ahluke omnye komnye, kwaye uvavanyo lokuthelekisa oluninzi lwe-Dunn lusetyenziswa xa i-statistic ye-Kruskal-Wallis ibalulekile.Ukufiphaza ngexesha lokufunda kokuphela kwavavanywa kusetyenziswa uhlalutyo lokulinganisa oluphindaphindiweyo lokuhluka kunye neqela njenge-inter-subject factor kunye novavanyo njengento ye-intra-subject factor. Iziphumo zathathwa njengezibalulekileyo xa p <0.05 kuzo zonke iimvavanyo zezibalo. Iziphumo zathathwa njengezibalulekileyo xa p <0.05 kuzo zonke iimvavanyo zezibalo. Результаты считались значимыми при p < 0,05 для всех статистических тестов. Iziphumo zathathwa njengezibalulekileyo kwip <0.05 kuzo zonke iimvavanyo zeenkcukacha-manani.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. Iziphumo zathathwa njengezibalulekileyo kwip <0.05 kuzo zonke iimvavanyo zeenkcukacha-manani.
Umzobo we-2 ubonisa umgca wexesha lokulinga (Umfanekiso 2A) kunye nokusabalalisa rhoqo kuzo zonke iigundane eziphantsi kokuphela (Umfanekiso 2B). Iigundane kumaqela okutshatyalaliswa okulungileyo kunye namahlwempu ahluke kakhulu kula manqaku okukhumbula ukucima okubala (U = 0, p <0.001) (Umfanekiso 2C). Iigundane kumaqela okutshatyalaliswa okulungileyo kunye namahlwempu ahluke kakhulu kula manqaku okukhumbula ukucima okubala (U = 0, p <0.001) (Umfanekiso 2C). Крысы в ​​группах с хорошим и плохим угашением значительно различались потим рассчитаным показателям припоминания уга,00, p. Iigundane kumaqela alungileyo kunye namahlwempu okutshabalala ahluke kakhulu kula mazinga okukhumbula ukutshabalala okubalwayo (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, 1,00. Kula mazinga okukhumbula okutshabalalisa okubalayo, iigundane kumaqela amahle kunye namahlwempu okutshatyalaliswa ahluke kakhulu (U = 0, p <0.001) (Umfanekiso 2C).Kwakungekho nantlukwano ephawulekayo kwixesha lokukhenkceza phakathi kwamaqela anokuphela okulungileyo, ukutshabalala okubi, kunye noloyiko lokukhumbula ngexesha lesiseko seseshoni ye-reflex yoloyiko (X2 (2) = 2.746, p = 0.253) (Umfanekiso 2D).Ukongezelela, ngexesha lokuboniswa kwethoni yokuqala ye-reflex yoloyiko lwe-reflex, kwakungekho mahluko abalulekileyo kwixesha lokukhenkceza phakathi kwamaqela anokutshabalala okulungileyo, ukuphela kakubi kunye noloyiko lokukhumbula (X2 (2) = 1.107, p = 0.575), njengoko kunye nangexesha loloyiko ngexesha leethowuni zesibini.Ngethuba leseshoni yokulungelelanisa, kwakukho umehluko omkhulu kwixesha lokukhenkceza phakathi kwamaqela anokutshatyalaliswa okulungileyo, ukutshatyalaliswa okungahambi kakuhle, kunye nokwesaba ukukhumbula (X2 (2) = 2.214, p = 0.331) (Umfanekiso 2D).Kwakungekho nantlukwano ebalulekileyo kwixesha eliphelileyo phakathi kwamaqela alungileyo kunye namahlwempu okutshatyalaliswa ngexesha loqeqesho olusisiseko lokuphela (U = 45.00, p = 0.799) (Umfanekiso 2D). Emva koko, kwakukho impembelelo ebalulekileyo yebhloko yesilingo (iithoni ze-5 ngebhloko nganye) ngexesha elichithwe iqhwa ngexesha leseshoni yoqeqesho lokucima (F (2.884, 54.80) = 8.331, p <0.001), ebonisa ukuba ukufundwa kokuphela kwenzeka (Fig. 2D ). Emva koko, kwakukho impembelelo ebalulekileyo yebhloko yesilingo (iithoni ze-5 ngebhloko nganye) ngexesha elichithwe iqhwa ngexesha leseshoni yoqeqesho lokucima (F (2.884, 54.80) = 8.331, p <0.001), ebonisa ukuba ukufundwa kokuphela kwenzeka (Fig. 2D ). Затем наблюдался значительный основной эффект пробного блока (5 тонов на блок) на время, затрачиваемое на замирание во время2,8т 0) = 8,331, p <0,001), что указывает на то, что обучение угашению происходило ( umfanekiso 2D). Kwakukho umphumo obalulekileyo obalulekileyo webhloko yesilingo (iithoni ze-5 ngebhloko nganye) ngexesha elithathiweyo ukukhenkceza ngexesha loqeqesho lokuphela (F (2.884, 54.80) = 8.331, p <0.001), ebonisa ukuba ukufunda ukucima kwenzeka (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（4.0803). ，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（4.0803). ，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,8,80,80 (p 01), что указывает на то, что обучение угашению происходило (рис 2D). Emva koko, ngexesha lokufunda ukucima, iibhloko zelingo (iithoni ezi-5 ngebhloko nganye) zinefuthe eliphambili eliphambili kwixesha le-fade (F (2.884, 54.80) = 8.331, p <0.001), ebonisa ukuba ukufunda ukucima kwenzeka (Fig. 2D) .Nangona kunjalo, iqela lokuphela (F (1, 19) = 3.091, p = 0.095) lalingenalo impembelelo ebalulekileyo kwixesha lokucima ngexesha lonke loqeqesho lokuphela, kwaye kwakungekho ukusebenzisana phakathi kwebhloko yesilingo kunye neqela lokucima (F (4) , 19)).76) = 1.890, p = 0.121) (Umfanekiso 2D). Ngexesha leseshoni yovavanyo, bekukho umahluko omkhulu phakathi kokutshabalala okuhle, ukuphela kakubi, kunye noloyiko lokukhumbula amaqela ngexesha elichithwe ngumkhenkce ngexesha lesiseko (X2 (2) = 8.569, p = 0.014) kangangokuba iqela lokukhumbula uloyiko likhenkceke kakhulu. ngaphezu kweqela elilungileyo lokuphela (I-Mean Rank Diff. = 10.57, p = 0.017), kodwa kungekhona iqela elibi lokuphela (Mean Rank Diff. = - 3.714, p > 0.999) (Fig. 2D). Ngexesha leseshoni yovavanyo, bekukho umahluko omkhulu phakathi kokutshabalala okuhle, ukuphela kakubi, kunye noloyiko lokukhumbula amaqela ngexesha elichithwe ngumkhenkce ngexesha lesiseko (X2 (2) = 8.569, p = 0.014) kangangokuba iqela lokukhumbula uloyiko likhenkceke kakhulu. ngaphezu kweqela elilungileyo lokuphela (I-Mean Rank Diff. = 10.57, p = 0.017), kodwa kungekhona iqela elibi lokuphela (Mean Rank Diff. = - 3.714, p > 0.999) (Fig. 2D).Ngexesha leseshoni yovavanyo, bekukho umahluko omkhulu phakathi kokutshabalala okuhle, ukuphela kakubi, kunye noloyiko lokukhumbula amaqela ngexesha elichithwe ngumkhenkce ngexesha lesiseko (X2 (2) = 8.569, p = 0.014), kangangokuba iqela lokukhumbula uloyiko likhenkce. ngokubalulekileyo .больше, чем в группе хорошего вымирания (средняя разница рангов = 10,57, p = 0,017), но не в группе плохого плохого плохого врамирания, 3, 3 0,999) (рис. 2D). mkhulu kuneqela lokuphela elilungileyo (lithetha umahluko wenqanaba = 10.57, p = 0.017) kodwa hayi kwiqela elibi lokuphela (lithetha umahluko wenqanaba = -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安方方面存在显兰方方面存在显着20 (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 方间 方间 方 方 方 方 方 方 方 方 方 方 方 方 方 方 方 方 显 存 显 22 569 , p = 0.014） , 恐惧 回忆组 冻结 显着 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 > 0.999）2） В течение периода тестирования наблюдалась значительная разница между хорошим угашением, группой с плохительная группой с хорошим угашением, группой с плохительная раха с точки зрения времени замирания на исходном уровне (X2 (2) = 8,569, p = 0,014), поэтому припоминание припоминание группа замерзачает , чем л с х с с с с х с соорошимамиеица рагням (н агов = 10,57, P = 0,017), Но Немица ра На Нгов = -3,714, p> 0,999) (рс. 2D). Ngexesha lovavanyo, bekukho umahluko omkhulu phakathi kweqela elilungileyo lokutshabalala, iqela lokutshabalala elibi, kunye neqela lokukhumbula uloyiko ngokwexesha lokukhenkceza kwisiseko (X2 (2) = 8.569, p = 0.014), ngoko ke uloyiko luyakhumbula. iqela liba ngumkhenkce ngokuphawulekayo ngokuphindaphindiweyo ngaphezu kweqela elinokuphela okulungileyo (umlinganiselo werenki uthetha 10.57, p = 0.017) kodwa kungekhona iqela elinokuphela kakubi (umlinganiselo wezinga = -3.714, p> 0.999) (Umfanekiso 2D).Iqela elilungileyo lokutshabalala, iqela elihlwempuzekileyo lokutshabalala, kunye neqela lokukhumbula uloyiko nazo zazinamaxesha ahlukeneyo okufiphala ngexesha lokuboniswa kwethoni yeseshoni yovavanyo (X2(2) = 14.93, p = 0.001), ngoko ke iqela elilungileyo lokutshabalala libe lincinci kakhulu. ixesha.ixesha lokukhenkceza kuneqela lokuphela elibuthathaka (lithetha umahluko wenqanaba = 9.286, p = 0.044) kunye neqela lememori yoloyiko (umlinganiselo wenqanaba = 13.86, p = 0.001) (Umfanekiso 2D).
Umahluko ngamnye ekukhunjulweni kokutshabalala.(A) Inkcazo yeenkqubo zotyando kunye nokuziphatha.(B) Ukuhanjiswa rhoqo okubonisa ukungafani komntu ngamnye kumanqaku ememori alahlekileyo.(C) Ubungqina bokuba amaqela ayilwe ngokusekwe kumanqaku okubalwa okukhumbula ukutshabalala amele iiphenotypes ezimbini ezahlukeneyo.(D) Ipesenti yeepesenti zexesha leempuku zikhenkceza ukutshabalala okubi, ukuphela okulungileyo, kunye nokukhumbula ukwesaba kwiiseli ze-30 zeseshini ye-conditioned reflex session, kwiitoni ze-20, i-30 s, yawela kwiibhloko ezi-5 ngexesha leseshoni yokufunda yokuphela (iithoni ezi-4) .nganye), kunye neetoni ezine kwiiseshoni zeenkumbulo zokuphela kunye neenkumbulo zoloyiko.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo. *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.
Isalathisi sokubuyisela sifakwe kwi-IL (Umfanekiso 3A) kunye nenani leeseli ze-GFP + kunye ne-anterior-posterior axis yommandla womdla unqunywe (Umfanekiso 3B-F).Kwakukho umehluko omkhulu kwinani leeseli ze-GFP + phakathi kwe-PVT yangaphambili, ephakathi kunye ne-posterior (X2 (2) = 8.200, p = 0.017), ngoko i-mPVT ibonise ngokuphawulekayo iiseli ze-GFP + kune-aPVT (inqanaba eliqhelekileyo) Diff.= 18.37, p = 0.035) kunye ne-pPVT (inqanaba lithetha i-Diff. = 17.71, p = 0.045) (Umfanekiso 3C).Nangona izilwanyana ezininzi azizange zibone naziphi na iiseli ze-GFP + kwi-pCLA kwaye ngenxa yoko azikwazanga ukwenza imephu kulo mmandla, akukho mahluko ubalulekileyo phakathi kwe-CLA yangaphambili, ephakathi, nasemva (X2 (2) = 5.596, p = 0.061).Inani leeseli zeGFP + (Figure 3D).Emva koko, ekubeni ezinye iiseli ze-GFP + zifunyenwe kwi-aBLA okanye i-avHPC kwiigundane ezininzi, kuphela umbindi kunye nomva wale mimandla yahlalutywa.I-BLA ephakathi kunye ne-posterior (U=393, p = 0.009) yahluke kakhulu kwinani leeseli ze-GFP +, ngoko ke i-pBLA ibonise iiprojekthi ze-IL ezininzi kune-mBLA (Umfanekiso 3E).Ngokufanayo, bekukho umahluko omkhulu phakathi kwe-vHPCs ephakathi nasemva, ngoko ke ii-pvHPCs zibonise iingqikelelo ze-IL ezininzi kune-mvHPCs (U = 403.5, p = 0.014) (Umfanekiso 3F).Umfanekiso we-3G ngumfanekiso ongumzekelo obonisa i-Fos, i-aavRG-GFP, kunye neeseli ezibhalwe kabini.
Ukulinganisa i-IL afferents kuwo wonke ummandla wobuchopho onomdla.(A) Umelo olucwangcisiweyo losasazo lwe-aavRG-CAG-GFP kuyo yonke irat IL.(B) Imifanekiso emeleyo yabamakishi be-retrograde kwiindawo ezahlukeneyo ze-anteroposterior kwingingqi yengqondo yomdla.Ubungakanani belebhile yokubuyisela umva ecaleni kwe-anteroposterior axis (C) paraventricular thalamus, (D) clavicle, (E) i-basolateral tonsil, kunye (F) ne-ventral hippocampus.(G) Imifanekiso emeleyo ebonisa ukuleyibhelishwa kwakhona kwe-aavRG, ukuleyibheli kwe-Fos, kunye ne-aavRG ephindwe kabini kunye ne-Fos yokuleyibheli kwi-APVT.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo. *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.Ibar yesikali 100 µm.Imephu yengqondo yedomeyini yoluntu kwiphaneli A yenziwe kwakhona ukusuka e-Swanson (2004) Imephu yoBuchule: I-Rat Brain Structure, uHlelo lwesi-3, olunikwe ilayisenisi phantsi kwe-Creative Commons Attribution-NonCommons 4.0 International License (https://creativecommons.org/licenses/by-nc )./4.0/) ekhoyo ukuze ukhutshelwe apha https://larrywswanson.com.
Umsebenzi we-Fos wehlabathi jikelele kunye ne-IL wahlalutywa kwi-APVT, i-mPVT kunye ne-pPVT kuzo zonke iimpuku.Kwakungekho mahluko ubalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya kwintetho yeFos kwi-APVT (X2(3) = 3.888, p = 0.274) (Fig. 4A), kwaye kwakungekho nxulumano lubalulekileyo phakathi kweFos. kwi-aPVT phakathi kwentetho kunye nokuphela kokukhumbula (rs = 0.092, p = 0.691) (Umfanekiso 4B) okanye phakathi kwe-Fos ibinzana kwi-APVT IL afferents kunye nokukhumbula ukuphelelwa (rs = 0.143, p = 0.537) (Umfanekiso 4D).Nangona kunjalo, kwi-APVT IL afferents, i-Fos expression yahluke kakhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya (X2(3) = 15.05, p = 0.002), ngoko ke iqela lokukhumbula uloyiko libonise ukuphela okulungileyo.ukuhlehla (umlinganiselo wenqanaba lomlinganiselo = 11.54, p = 0.003), ukuhlehla okungahambi kakuhle (umlinganiselo wenqanaba = 10.57, p = 0.034), kunye neseli yasekhaya (umlinganiselo wenqanaba = 12.79, p = 0.005) amaqela (Fig. 4C).Ukuqhubela phambili, kwakungekho mahluko ubalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya ze-Fos expression kwi-mPVT (X2(3) = 2.272, p = 0.518) (Fig. 4E) kunye ne-Fos expression kwi-mPVT..Ulungelelwaniso olubalulekileyo kunye nokukhumbula ukuphelelwa (rs = 0.168 p = 0.468) (Umfanekiso 4F).Nangona bekukho umahluko omkhulu phakathi kokulungileyo, okubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya kwintetho ye-Fos kwi-IL afferent mPVT iiseli (X2(3) = 9.252, p = 0.026), uthelekiso lwe-post hoc alukhange luveze nanye okanye ezimbini.Umahluko omkhulu phakathi kwamaqela (Figure 4G).Ngaphaya koko, kwakungekho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kwiiseli ze-IL-afferent mPVT kunye nokukhumbula ukuphela (rs = 0.174, p = 0.450) (Umfanekiso 4H). Okulandelayo, kwakukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula ukwesaba, kunye namaqela ekheji yasekhaya kwintetho ye-Fos kwi-pPVT (X2 (3) = 13.89, p = 0.003), kangangokuba iqela lokuphela okulungileyo Umahluko = 14.96, p = 0.010), kodwa hayi ukuphelelwa kakubi (Mean Rank Diff. = 12.86, p = 0.113) okanye iqela lokukhumbula uloyiko (Mean Rank Diff. = 2.571, p > 0.999), ebonisa imbonakalo yeFos engaphezulu kuneyo iqela lekheji yasekhaya (Umfanekiso 4I). Okulandelayo, kwakukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula ukwesaba, kunye namaqela ekheji yasekhaya kwintetho ye-Fos kwi-pPVT (X2 (3) = 13.89, p = 0.003), kangangokuba iqela lokuphela okulungileyo Umahluko = 14.96, p = 0.010), kodwa hayi ukuphelelwa kakubi (Mean Rank Diff. = 12.86, p = 0.113) okanye iqela lokukhumbula uloyiko (Mean Rank Diff. = 2.571, p > 0.999), ebonisa imbonakalo yeFos engaphezulu kuneyo iqela lekheji yasekhaya (Umfanekiso 4I). Далее, наблюдалась значительная разница между группами с хорошим, плохим угашением, отзывом страхай домашение в домашение (X2 (3) = 13,89, p = 0,003), так что группа с хорошим угашением (средний ранг Ukwahluka = ​​14,96, p = 0,010), но не в группе плохого угашения (средняя ранговая разница = 12,86, p = 0,113) или групппе плохого угашения. разница = 2,571, p > 0,999), демонстрировалась более выраженная экспрессия Fos, чем в группе группа домашних клеток (рис. 4I). Ngaphaya koko, bekukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya kwinkcazo ye-Fos kwi-pPVT (X2(3)=13.89, p=0.003), ukuze iqela elilungileyo lokutshabalala (lithetha inqanaba leDiff. = 14.96, p = 0.010), kodwa hayi kwiqela elibi lokubhangisa (umlinganiselo wenqanaba lomahluko = 12.86, p = 0.113) okanye iqela lenkumbulo yoloyiko (umlinganiselo wobumehluko = 2.571, p > 0.999), ubonise ukuchazwa kweFos evakale ngakumbi kunakwi iqela leseli yasekhaya (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 9） 3x2 ） 3 x 9 = 0.使得 消组 （(intsingiselo yeRenki Diff.= 14.96, p = 0.010)，但不是较差的消退(UMahluko oLingeneyo weNqanaba. = 12.86, p = 0.113) 或恐惧回忆组(Isikhundla esiLingeneyo 4I ）.= 14.96, p = 0.010) , 但不是较差的消退(Mean Rank Diff. = 12.86, p = 0.113) ）.Okwesibini, bekukho ukungafani okubonakalayo kwintetho ye-Fos kwi-pPVT phakathi kokulungileyo, okubi, ukukhumbula ukwesaba, kunye namaqela eeseli zasekhaya (X2 (3) = 13.89, p = 0.003), okwenza iqela elihle lokuthatha (lithetha umahluko wezinga = 14.96)., p = 0,010), но не хуже по угашению (средняя разница рангов = 12,86, p = 0,113) okanye группе отзыва страха (средня разница, 9,9, 9), в группе домашней клетки (рис. 4I) . , p = 0.010), kodwa akukho mbi ngakumbi ekuphelisweni (umlinganiselo werenki ulinganisa = 12.86, p = 0.113) okanye iqela lokukhumbula uloyiko (umlinganiselo wenqanaba = 2.571, p > 0.999) kuneqela leseli yasekhaya (Umfanekiso 4I)..Nangona kunjalo, akukho nxu lumano lubalulekileyo phakathi kwe-pPVT Fos inkcazo kunye nokukhumbula ukuphela (rs = 0.051, p = 0.825) (Umfanekiso 4J).Okokugqibela, bekukho umahluko omkhulu kwintetho ye-Fos kwi-pPVT IL afferents phakathi kwamaqela anokuphela kakuhle, ukuphela kakubi, iinkumbulo zoloyiko, kunye nakwiiseli zasekhaya (X2 (3) = 12.34 p = 0.006), imbonakalo entle yeFos kwi-IL- kubi kakhulu kunamaqela okuphela (umlinganiselo wenqanaba = 12.54, p = 0.014) nakwiseli yasekhaya (umlinganiselo wenqanaba = 12.89, p = 0.049) (Umfanekiso we-4K) kwaye uhambelana kakhulu ne-IL afferents ngaphakathi kwe-pPVT phakathi kokusebenza kunye ukurhoxiswa kokutshabalala, ukukhumbula ukupheliswa okungcono kwakunxulunyaniswa nokusebenza okukhulu kwezi zixhobo ze-IL (rs = -0.438, p = 0.047) (Umfanekiso 4L).
Umsebenzi we-Fos unyuswe kwii-IL afferents ze-posterior paraventricular thalamus (PVT) kwiigundane, ezibonise ukuhlehla okulungileyo.(A) Akukho mahluko ubalulekileyo phakathi kwamaqela ekubonakalisweni kwe-Fos kwi-APVT.(B) Akukho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kunye nokukhumbula ukuphela kwe-APVT.(C) Iqela lokukhumbula uloyiko libonise ukwanda kokubonakaliswa kwe-Fos kwii-IL afferents xa kuthelekiswa nawo onke amanye amaqela.(D) Bekungekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-IL afferents kunye nokuphela kokukhumbula kwi-APVT.(E) Akukho mahluko ubalulekileyo weqela eliphakathi kwintetho yeFos kwi-mPVT.(F) Akukho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kunye nenkumbulo yokuphela kwe-mPVT.(G) Inkcazo ye-Fos kwiiseli ze-IL ezihlukeneyo kwi-mPVT ayizange ihluke kakhulu phakathi kwamaqela.(H) Khange kubekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-IL afferents kunye nokuphela kwenkumbulo kwi-mPVT.(I) Iqela elatshabalalayo, kodwa alikho elinye iqela, elibonise ukwanda komsebenzi we-Fos kwi-pPVT xa kuthelekiswa neqela lekheji yasekhaya.(J) Akukho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kunye nokukhumbula ukuphela kwe-pPVT.(K) Iqela elilungileyo lokutshabalala libonise ukwanda kokubonakaliswa kwe-Fos kwiiseli ezihambelanayo ze-IL xa kuthelekiswa neqela elibuthathaka lokuphela kunye neqela leseli yasekhaya.(L) Kukho unxulumano olubalulekileyo phakathi kwenkcazo ye-Fos kwi-IL afferents kunye nokukhumbula ukuphela, ke ukukhunjulwa okulungileyo kokutshabalala kunxulunyaniswa nenkcazo enkulu ye-Fos kwii-IL afferents.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo. *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 wehlabathi jikelele kunye ne-IL woqikelelo oluthe ngqo kwi-aCLA kunye ne-mCLA yeempuku yahlalutywa kuwo onke amaqela. Bekukho umahluko obonakalayo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwimbonakalo yeFos kwi-aCLA (X2 (3) = 8.455, p = 0.036) kangangokuba iqela lokukhumbula uloyiko (Mean Rank Diff. = 14.50, p = 0.049), kodwa hayi amahlwempu (Mean Rank Diff. = 10.21, p = 0.373) okanye ukuphela okulungileyo (Mean Rank Diff. = 4.607, p > 0.999) amaqela, abonise ukubonakaliswa kweFos ngaphezulu kuneqela lekheji yasekhaya ( Umzobo 5A). Bekukho umahluko obonakalayo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwimbonakalo yeFos kwi-aCLA (X2 (3) = 8.455, p = 0.036) kangangokuba iqela lokukhumbula uloyiko (Mean Rank Diff. = 14.50, p = 0.049), kodwa hayi amahlwempu (Mean Rank Diff. = 10.21, p = 0.373) okanye ukuphela okulungileyo (Mean Rank Diff. = 4.607, p > 0.999) amaqela, abonise ukubonakaliswa kweFos ngaphezulu kuneqela lekheji yasekhaya ( Umzobo 5A). Между группами с хорошим угашением, плохим угашением, припоминанием страха и домашними клетками наблюдалась значительский ) = 8,455, p = 0,036), так что 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 ранговая раз,90,973, 9,999, p ровали большей экспрессии Fos, чем группа в домашней клетке ( Рис 5A). Bekukho umahluko omkhulu kwintetho ye-aCLA Fos phakathi kokuphela okulungileyo, ukuphela kakubi, ukukhumbula uloyiko kunye namaqela eeseli zasekhaya (X2(3) = 8.455, p = 0.036), ukuze iqela lokukhumbula uloyiko (lithetha umahluko wenqanaba = 14.50, p. = 0.049), kodwa ingengabo abahlwempuzekileyo (umlinganiselo womahluko werenki = 10.21, p = 0.373) okanye iqela elilungileyo lokuphela (lithetha umahluko wenqanaba = 4.607, p > 0.999) libonise ukubonakaliswa kweFos okuninzi kuneqela 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) = 惧回忆、恐惧回忆和家庭笼组之间存在显着差异(X2 (3) = 槐 = 8.45,槐 = 8.45忆组(UMahluko oLingeneyo weNqanaba. = 14.50,p = 0.049），但无论是差（平均秩差= 10.21，p = 0.373)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 2 差异 差异 差异 2. 455, p = 0.036) , 因此 恐惧 回忆组 回忆组 回忆组 回忆组 (ithetha ukuba Isikhundla Diff = 14.50 , P = P = P = P = P = 14.差 秩差 = 4.607 , p> 0.999） 组 都 出 比 家庭 笼组 更 多 的 表达 图 图 5a) . Была значительная разница между группами с хорошим угашением, плохим угашением, отзывом страха и домашней клеппами клет5 клеткой, 4 клеткой, 5 клеткой, 4 5 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) kwaye с хорошим вымиранием (средня разница рангов, 9, 9, 9, p сокую экспрессию Fos, чем группа с домашней клеткой (рис. 5A). Bekukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko kunye namaqela eeseli zasekhaya kwinkcazo ye-aCLA Fos (X2(3) = 8.455, p = 0.036), ngoko ke iqela lokukhumbula uloyiko (lithetha umahluko wenqanaba = 14.50), p = 0.049 ), kodwa amaqela ahluphekileyo (umlinganiselo wezinga elilinganisiweyo = 10.21, p = 0.373) kunye nokuphela okulungileyo (umlinganiselo wenqanaba = 4.607, p > 0.999) ubonise ukubonakaliswa kwe-Fos ephezulu kuneqela leseli yasekhaya (Umfanekiso 5A). .Bekungekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos yehlabathi kwi-(rs = 0.036, p = 0.876) (Figure 5B) okanye i-Fos expression in IL aCLA afferent cells (rs = -0.282, p = 0.215) kunye nokukhumbula ukuphela (Figure 5B)..5D), bekungekho mahluko ubalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya kwi-Fos expression kwi-aCLA IL afferents (X2(3) = 6.722, p = 0.081) (Figure 5C)..). Okulandelayo, bekukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwinkcazo ye-Fos kwi-mCLA (X2 (3) = 10.12, p = 0.018) kangangokuba iqela elilungileyo lokuphela (Lithetha iRanga Diff . = 12.93, p = 0.038), kodwa akukho kutshabalala okulambathayo (Mean Rank Diff. = 5.143, p > 0.999) okanye amaqela oloyiko lokukhumbula (Mean Rank Diff. = 14.00, p = 0.063) eboniswe ngokuphawulekayo ngakumbi iFos yokubonakalisa kwi-mCLA ngokunxulumene neqela lekheji yasekhaya (Umfanekiso 5E). Okulandelayo, bekukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwinkcazo ye-Fos kwi-mCLA (X2 (3) = 10.12, p = 0.018) kangangokuba iqela elilungileyo lokuphela (Lithetha iRanga Diff . = 12.93, p = 0.038), kodwa akukho kutshabalala okulambathayo (Mean Rank Diff. = 5.143, p > 0.999) okanye amaqela oloyiko lokukhumbula (Mean Rank Diff. = 14.00, p = 0.063) eboniswe ngokuphawulekayo ngakumbi iFos yokubonakalisa kwi-mCLA ngokunxulumene neqela lekheji yasekhaya (Umfanekiso 5E). Затем наблюдалась значительная разница между группами с хорошим угашением, воспоминаниями о спорошим, плохим угашением, воспоминаниями о спорошением mCLA (X2 (3) = 10,12, p = 0,018), так что группа с хорошим угашением (средняя разница рангов = 12,93, p = 0,038), но ни группы плохого угашения (средня ранговая разница = 5,143, p > 0,999), ни групппы плохого угашения . а = 14,00, p = 0,063) не показали значительно большей экспрессии Fos в mCLA. Kwakukho umahluko obalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, iinkumbulo zoloyiko kunye namaqela eeseli zasekhaya kwi-mCLA Fos expression (X2(3) = 10.12, p = 0.018), ukuze iqela elilungileyo lotshabalalo (lithetha umahluko wenqanaba = 12.93, p. = 0.038), kodwa ingengawo amaqela ahlwempuzekileyo okuphela (othetha umahluko werenki = 5.143, p > 0.999) okanye amaqela oloyiko lokukhumbula (umlinganiselo werenki umahluko = 14.00, p = 0.063) ubonise ngokuphawulekayo ukubonakaliswa kweFos kwi-mCLA.xa kuthelekiswa neqela lekheji yasekhaya (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 ((((((((()) ((((((iMean 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=0p=0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 = 5.143 , p> 0.999） = 0.999） = 4.999） ＆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 в 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 (средня разность рангов = 12,93, p = 0,038), но в mCLA ни слабое угасание (средня разница рангов = 5,143, pыpы999, p няя разница рангов = 14,00, p = 0,999) = 0,063) показали лучшую Много Много Ngaphaya koko, bekukho umahluko obalulekileyo ekubonakalisweni kwe-Fos kwi-mCLA phakathi kweqela elilungileyo lokuphela, iqela elilambathayo, iqela lengxelo yoloyiko, kunye neqela leeseli zasekhaya (X2(3) = 10.12, p = 0.018), ngoko ke, okulungileyo. ukuphela (umlinganiselo wenqanaba lomahluko = 12.93, p = 0.038), kodwa kwi-mCLA akukho kutshabalala olubuthathaka (umlinganiselo womahluko werenki = 5.143, p > 0.999) okanye iqela loloyiko lokukhumbula (umlinganiselo wenqanaba = 14.00, p = 0.999) = 0.063) ubonise ngcono Ukuchazwa kwe-Multi Fos xa kuthelekiswa neqela leseli yasekhaya (Umfanekiso 5E).Nangona kunjalo, ukubonakaliswa kwe-Fos yehlabathi kwi-mCLA (rs = 0.321, p = 0.156) (Fig. 5F) okanye kwiiseli ezidibeneyo ze-IL mCLA (rs = -0.121, p = 0.602) kunye nokukhumbula ukuphela (Fig. 5H), akukho mahluko ubalulekileyo phakathi amaqela anokuphela okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye neseli yasekhaya ye-Fos expression kwi-IL mCLA afferent cells (X2(3)=4.923, p=0.178) (Figure 5G).
Umsebenzi we-Fos uphakanyiswe phakathi kweclaustrum kwiimpuku ezinenkumbulo entle yokutshabalala.(A) Iqela lokukhumbula uloyiko, kodwa hayi amanye amaqela, libonise ukwanda komsebenzi we-Fos xa kuthelekiswa neqela leseli yasekhaya e-CLA.(B) Khange kubekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-aCLA kunye nokukhumbula ukuphela.(C) Ukuchazwa kwe-Fos kwi-IL afferent aCLA seli azihlukanga kakhulu phakathi kwamaqela.(D) Khange kubekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-IL afferents kunye nokukhumbula ukuphela kwe-aCLA.(E) Iqela elatshabalalayo, kodwa hayi amanye amaqela, libonise ukwanda komsebenzi we-Fos kwi-mCLA xa kuthelekiswa neqela leseli yasekhaya.(F) Bekungekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kunye nokukhumbula ukuphela kwe-mCLA.(G) Ukuchazwa kwe-Fos kwi-IL mCLA afferent cells akwahlukanga kakhulu phakathi kwamaqela.(H) Khange kubekho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-IL afferents kunye nokuphela kwenkumbulo kwi-mCLA.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo. *p <0.05. *p <0.05. * р < 0,05. *p <0.05. *p <0.05. *p <0.05. * р < 0,05. *p <0.05.
Emva koko, umsebenzi we-Fos wehlabathi jikelele kunye ne-IL ethile kwi-mBLA kunye ne-pBLA yahlalutywa kuwo onke amaqela eempuku.Kwakungekho mahluko ubalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya kwi-Fos expression kwi-mBLA (X2(3)=0.944, p=0.815) (Figure 6A).Kwakungekho mahluko ubalulekileyo phakathi kwamaqela anokuhlehla okuhle, ukuhlehla okungalunganga, ukukhumbula uloyiko, kunye nenkcazo yeseli ye-Fos yasekhaya kwi-IL mBLA iiseli afferent (X2(3)=0.518, p=0.915) (Figure 6C).Ukongeza, kwakungekho lunxulumano lubalulekileyo phakathi kwentetho yehlabathi ye-Fos kwi-mBLA (rs = 0.126, p = 0.588) (Umfanekiso 6B) kunye nokubonakaliswa kwe-Fos kwiiseli ezihambelanayo ze-IL mBLA (rs = 0.200, p = 0.385) (rs = 0.200, p. = 0.385).p = 0.385).Umzobo 6D) kunye nokukhumbula ukuphela.Kwakungekho mahluko ubalulekileyo ekuphelisweni kakuhle, ukuphela kakubi, inkumbulo yoloyiko, kunye namaqela eeseli zasekhaya kwi-Fos expression kwi-pBLA (X2 (3) = 4.246, p = 0.236) (Fig. 6E), kwaye kwakungekho mahluko ubalulekileyo. kwi pBLA Good.ukuphela, ukuphela kakubi, uloyiko lokukhumbula, kunye namaqela eeseli zasekhaya kwi-Fos expression kwiiseli ezidibeneyo ze-IL (X2(3)=1.954, p=0.582) (Figure 6G).Ekugqibeleni, i-global Fos expression in pBLA (rs = 0.070, p = 0.762) (Fig. 6F) kunye ne-Fos expression in pBLA IL afferent cells (rs = 0.122, p = 0.597) kunye nokuphela kokukhumbula (Fig. 6H).
Umahluko womntu ngamnye ekuveliseni kwakhona ukutshabalala awuzange uboniswe umahluko kwintetho ye-Fos kwi-basolateral amygdala.(A) Akukho mahluko ubalulekileyo weqela eliphakathi kwintetho yeFos kwi-mBLA.(B) Kwakungekho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kunye nokukhumbula ukuphela kwe-mBLA.(C) Ukubonakaliswa kwe-Fos kwi-IL mBLA afferent cells awahlukanga kakhulu phakathi kwamaqela.(D) Kwakungekho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kwiiseli ezihambelanayo ze-IL kunye nokukhumbula ukuphela kwe-mBLA.(E) Akukho mahluko ubalulekileyo weqela eliphakathi kwintetho yeFos kwipBLA.(F) Akukho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kunye nokukhumbula ukuphela kwe-pBLA.(G) Ukuchazwa kwe-Fos kwiiseli ezidibeneyo ze-IL pBLA azihlukanga kakhulu phakathi kwamaqela.(H) Akukho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwiiseli ezihambelanayo ze-IL kunye nokukhumbula ukuphela kwe-pBLA.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo.
Okokugqibela, umsebenzi we-Fos wehlabathi jikelele kunye ne-IL wahlalutywa kwi-mvHPC kunye ne-pvHPC kuzo zonke iimpuku. Kwakukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwinkcazo ye-Fos kwi-mvHPC (X2 (3) = 8.056, p = 0.045) kangangokuba ukuphela okulungileyo (Kuthetha iRanga Diff. = 13.29 , p = 0.031), kodwa akukho kuphelelwa kakuhle (Mean Rank Diff. = 6.857, p > 0.999) okanye uloyiko lokukhumbula (Mean Rank Diff. = 8.000, p = 0.864) amaqela abonise ukubonakaliswa kweFos ngaphezulu kuneqela lekheji yasekhaya (Fig. 7A). Kwakukho umahluko omkhulu phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye namaqela ekheji yasekhaya kwinkcazo ye-Fos kwi-mvHPC (X2 (3) = 8.056, p = 0.045) kangangokuba ukuphela okulungileyo (Kuthetha iRanga Diff. = 13.29 , p = 0.031), kodwa akukho kuphelelwa kakuhle (Mean Rank Diff. = 6.857, p > 0.999) okanye uloyiko lokukhumbula (Mean Rank Diff. = 8.000, p = 0.864) amaqela abonise ukubonakaliswa kweFos ngaphezulu kuneqela lekheji yasekhaya (Fig. 7A). Между группами с хорошим угасанием, плохим угашением, отзывом страха и домашними клетками наблюдалась значительась значительась значительасы = 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 знраням зрезавос зред, 0, p = 0,864) экспрессия Fos была выше, чем в группе с домашней клеткой (рис. 7А). Kwakukho umahluko omkhulu kwi-mvHPC Fos yokubonakaliswa phakathi kwamaqela anokuphela okulungileyo, ukuphela kakubi, ukukhumbula uloyiko, kunye neeseli zasekhaya (X2 (3) = 8.056, p = 0.045), ngoko ke ukupheliswa okulungileyo (umlinganiselo wenqanaba = 13.29), p = 0.031), kodwa hayi kwiqela elibi lokubhanga (umlinganiselo womahluko werenki = 6.857, p > 0.999) okanye kwiqela lokukhumbula uloyiko (umlinganiselo womahluko wenqanaba = 8.000, p = 0.864) yayiyimbonakalo yeFos ephezulu kuneqela lasekhaya.iseli (Fig. 7A). mvHPC 中Fos 表达的良好消退、不良消退、恐惧回忆和家庭笼组之间存在显着差异（消退（X2.05）（X2.05）（X2Ｍ3）（X2））（8（3)好消退（平均秩差= 13.29） ，p = 0.031 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 （ 30 = 5 （ 30 = 5 （ 30 = 5 （ 30 = 5). ） , 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 60 = 60 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 экспрессии3PC =(v60, p80, p,80,60,60,60,5,5,5,8,5,5,5,5,5,5,5,5,5,5,5,5,5,5,20; и, 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 = 0,80 = 0,857, p ,864) показали более высокую экспрессию Fos, чем группа в домашней клетке (рис. 2). Bekukho umahluko omkhulu phakathi kokulungileyo, okubi, ukukhumbula uloyiko, kunye namaqela eeseli zasekhaya zokubonakaliswa kwe-Fos kwi-mvHPC (X2(3) = 8.056, p = 0.045) kwaye ke ukuhlehla okuhle (uthetha umahluko wenqanaba = 13.29), p = 0.031), kodwa amaqela anokutshabalala okubuthathaka (umlinganiselo werenki ethetha i-6.857, p> 0.999) kunye neenkumbulo zoloyiko (umlinganiselo wenqanaba = 8.000, p = 0.864) ubonise ukubonakaliswa kwe-Fos ephezulu kuneqela kwikheji yasekhaya (Umfanekiso 2).7A).Nangona kunjalo, kwakungekho mmahluko obalulekileyo (X2 (3) = 4.893, p = 0.180) (Umfanekiso 7C).Ukongeza, akukho nxulumano lubalulekileyo phakathi kwentetho ye-Fos yehlabathi jikelele kwi-mvHPC (rs = -0.233, p = 0.309) (Umfanekiso 7B) kunye nokubonakaliswa kwe-Fos kwiiseli ze-IL ze-mvHPC (rs = 0.056, p = 0.810) (Umfanekiso 7D).kunye nengxelo ngokulahleka.Ukuqhubela phambili, akukho mahluko ubalulekileyo phakathi kokutshabalala okulungileyo, ukuphela kakubi, ukukhumbula ukwesaba, kunye namaqela eeseli zasekhaya kwi-Fos expression kwi-pvHPC (X2 (3) = 3.623, p = 0.353) (Umfanekiso 7E), kwaye akukho mahluko ubalulekileyo.Umahluko ekubuyiseleni okuhle kwentetho ye-Fos kwiiseli ze-IL ze-pvHPC, ukuhlehla okungahambi kakuhle, inkumbulo yoloyiko, kunye namaqela eeseli zasekhaya (X2(3)=3.871, p=0.276) (Fig. 7G).Okokugqibela, kwakungekho lunxulumano lubalulekileyo phakathi kwe-pvHPC yelizwe jikelele intetho ye-Fos (rs = -0.127, p = 0.584) (Umfanekiso 7F) kunye nokubonakaliswa kwe-Fos kwiiseli ze-IL-afferent pvHPC (rs = 0.176, p = 0.447) kunye nokukhumbula ukuphela (Umfanekiso 7F ).7H).
Ukuchazwa kwe-Fos kuphakanyiswe kwi-ventral hippocampus yeempuku, ebonisa ukuphela kwenkumbulo elungileyo.(A) Iqela elaphelayo, kodwa hayi amanye amaqela, libonise ukwanda kokubonakaliswa kweFos kwi-mvHPC xa kuthelekiswa neqela leseli yasekhaya.(B) Bekungekho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kunye nokukhumbula ukuphela kwe-mvPHC.(C) Ukubonakaliswa kwe-Fos kwi-mvHPC afferent IL iiseli azihlukanga kakhulu phakathi kwamaqela.(D) Bekungekho lunxulumano lubalulekileyo phakathi kwenkcazo ye-Fos kwi-IL afferents kunye nokuphela kokukhumbula kwi-mvHPC.(E) Akukho mahluko ubalulekileyo phakathi kwamaqela kwintetho ye-Fos kwi-pvHPC.(F) Akukho lunxulumano lubalulekileyo phakathi kwentetho ye-Fos kunye nokukhumbula ukuphela kwe-pvHPC.(G) Inkcazo ye-Fos kwiiseli ze-IL ze-pVHPC azihlukanga kakhulu phakathi kwamaqela.(H) Akukho lunxulumano lubalulekileyo phakathi kwenkcazelo ye-Fos kwi-IL afferents kunye nokuphela kwenkumbulo kwi-pvHPC.Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo. *p <0.05. *p <0.05. * р < 0,05. *p <0.05. *p <0.05. *p <0.05. * р < 0,05. *p <0.05.Uhlalutyo lwethu oluphambili kuyo yonke imimandla lubonise ukuthelekisa kumanqanaba amathathu kunye ne-anteroposterior axis, nangona siphinde sihlalutye ummandla ngamnye oye wawa kunye ne-anteroposterior axis.Iziphumo zolu hlalutyo zinikwe kwitheyibhile yoku-1.
Apha siye savavanya ukuba ngaba iyantlukwano yomntu ekukhunjulweni kokutshabalala ingabonakaliswa kwiipatheni ezahlukeneyo zomsebenzi ochaseneyo kwi-limbic cortex esezantsi.Ukuza kuthi ga ngoku, siye savavanya umsebenzi we-Fos kuqikelelo lwe-IL ukusuka kwi-paraventricular thalamus, iclaustrum, i-basolateral tonsil, kunye ne-ventral hippocampus emva kokuphinda kuphele.Kwiiseli ze-IL-projecting, sifumene umsebenzi ophezulu kwingingqi yangasemva ye-PVT kwiigundane ezibonise ukukhumbula ukutshabalala okulungileyo xa kuthelekiswa neempuku ezinokuphela kakubi.Kwakungekho nantlukwano kwi-IL afferents kwi-nucleus ye-clavicular, i-ventral hippocampus, okanye i-tonsil ye-basolateral.Ukongeza kwiiseli ze-IL-projecting, ukwanda komsebenzi we-neural kwabonwa kwiindawo ezikhethiweyo ze-rat claustrum kunye ne-ventral hippocampus enesisombululo esihle.Iziphumo zethu zibonisa ukuba inkumbulo yokuphela kwempumelelo icwangciswe ngoqikelelo oluthile lwe-PVT ukuya kwi-IL kunye neeseli ezingajoliyo kwi-IL kwi-claustrum kunye ne-ventral hippocampus.
Sifumene ukuba i-PVT IL yokubikezela yayisebenza kwiigundane ezibonisa ukukhumbula ukutshatyalaliswa okulungileyo, okuhambelana nophononongo lwakutsha nje olubonisa ukuba i-PVT iyadingeka ekukhunjulweni kokuphela.Olu phononongo aluzange lusebenzise ukuguqulwa kwe-subregion-specific manipulations, kodwa lubonise ukuba zombini iingqikelelo ze-PVT kwi-lateral central amygdala kunye noqikelelo lwe-IL kwi-PVT zazifuneka ukuvelisa kwakhona ukuphela.Iziphumo zethu zibonisa ukuba, ngaphezu kwe-IL-PVT-CeL chain, ukungena kwe-post-PVT kwi-IL kunokufuneka kwakhona ukucima ukukhumbula.Ngaloo ndlela, kuvela ukuba zombini i-efferent kunye ne-afferent uxhumano lwe-IL lubandakanyeka ekwenziweni kwakhona kokuphela.Isinyathelo esilandelayo esibalulekileyo kukufumanisa ukuba yintoni ebangela ukuba i-pPVT ibonise ukutshatyalaliswa kokuveliswa kwakhona kwinqanaba le-neural circuit.Ukongeza kubudlelwane kunye ne-IL, izifundo ze-duct-tracking zangaphambili31,32 zibonise ukuba i-pPVT ifumana igalelo kwi-ventral periaqueductal grey (vPAG), ehambelana nokufunda kokuphela33,34,35,36.Nangona indima ye-vPAG ekukhunjulweni kokutshabalala ayizange isungulwe, ukuqikelelwa kwe-pPVT yi-vPAG ngumgqatswa onomtsalane ngenxa yobuninzi babo kunye nokubandakanyeka kwayo yomibini imimandla ekuveliseni ubungqina bangaphambili bokuphela koloyiko.
Omnye umba obalulekileyo weziphumo zethu zePVT kukuba zikhethekileyo ecaleni kwe-anterior-posterior axis.Okumangalisayo kukuba, umsebenzi we-neuronal wentelekelelo ye-PVT kwi-IL ihambelana nemeko yokuziphatha echaseneyo, efana nokuba umsebenzi woqikelelo wangaphambili we-PVT kwi-IL unxulunyaniswa nokukhumbula uloyiko, ngelixa uqikelelo lwe-pPVT lusebenza emva kokuphela kwenkumbulo eyimpumelelo (okt, uloyiko).Le heterogeneity esebenzayo ngaphakathi kwe-PVT ayimangalisi inikwe umsebenzi wangaphambili [kuxoxiwe kwi-37].Umzekelo ophawulekayo wokusasazwa okusebenzayo kwi-PVT usanda kuvela kwisifundo esiye sabonakalisa iimpawu zeentlobo ezithile zeeseli kwi-PVT.Olu phononongo lubonisa ukuba iiseli ze-dopamine ezibonisa i-DRD2 zibonakaliswa ikakhulu kwi-pPVT, zingena ngaphakathi kwi-cortex yangaphambili, kwaye ziphendule kwi-aversive stimu.Isibalo sesibini seeseli sibonakaliswa kakhulu kwi-APVT kwaye iphawula utshintsho oluya kwimeko ephantsi yokuvuselela umzimba kwaye ingenasiphelo i-cortex yamalungu angaphantsi.Iziphumo zethu azifane zilingane kule pateni, njengoko iiseli ze-IL-projecting aPVT zisebenza ngexesha lokukhumbula uloyiko, ngelixa uqikelelo lwe-pPVT lusebenza kwaye izilwanyana zibonisa amanqanaba asezantsi oloyiko.Ubuncinci zimbini iinkcazo ezinokubakho zokungangqinelani okubonakalayo.Okokuqala, iintlobo zeeseli ezichongiweyo azifumanekanga kuphela kwindawo enye yangaphambili-ngasemva yeTVV.Ke ngoko, iiseli ze-IL-projecting pPVT ezisebenzayo kwiimpuku ezinenkumbulo yokuphela kakuhle zinokuba zezodidi lweeseli ezinokuthi zibhaqwe kwi-aPVT kwaye zibonise inguqu ukuya kwimeko ephantsi yokuvuka.Okufanayo kunokuba yinyani kwiiseli ze-IL-projecting kwi-PVT evuselelwe emva kwenkumbulo yoloyiko.Okwesibini, izifundo zangaphambili zokulandela umkhondo zichonge ubukho be-IL3-projecting pPVTs, nangona ezinye zibonakala ziphuma kwiiseli ezine-DRD2, ezinye iintlobo zeeseli zinokuprojekthi kwi-IL kwaye zisebenze ekuveliseni ngempumelelo ukucima.
Nangona injongo yolu phononongo yayikukubona umahluko phakathi kweempuku ezibonisa i-phenotypes ezahlukeneyo zokutshabalala, ezi zilingo ziveze idatha entsha enxulumene neendlela zokukhumbula uloyiko.Okubangela umdla, sifumene ukonyuka komsebenzi we-Fos kwi-CLA yangaphambili kwiimpuku ezinenkumbulo yoloyiko.
I-clavicle ibekwe njengeziko lonxibelelwano lwe-cortical kwaye ibandakanyeka kwiinkqubo ezivela ekudibaneni kweengqondo ukuya kwingqalelo kunye nokulala40,41,42,43.Kukho ubungqina obulinganiselweyo malunga nendlela i-claustrum ebandakanyeka ngayo kwisimo soloyiko okanye ukubonakaliswa koloyiko, nangona kunjalo, uphando lwangaphambili lubonise ukuba ukubonakaliswa koloyiko lomxholo kubandakanyeka kumsebenzi we-Fos kwi-claustrum.Kutshanje kuye kwaxelwa ukuba inhibition ye-atresia projections kwi-entorhinal cortex ngexesha loloyiko lomxholo luphazamisa ukubunjwa kwememori yexesha elide, nangona imfuno yabo yokoyika ukubonakaliswa ingakhange ivavanywe.Kuphononongo olufanayo, ukonyuka kokwenziwa kusebenze kwe-Fos kwabonwa xa izilwanyana zisesichengeni sendawo entsha xa kuthelekiswa neempuku ezivezwe kwindawo eqhelekileyo.Unale nto engqondweni, ukusebenza kwe-CLA esiyinika ingxelo apha kusenokuba kungenxa yokuvezwa kwikhamera entsha ngexesha lovavanyo, endaweni yokoyika ukuzikhumbula.Ukubonakalisa ngokuchanekileyo ngakumbi umsebenzi wezitshixo kuloyiko kunye nokulungiswa kwemeko, izifundo zexesha elizayo kufuneka zisebenzise ukuqhatha okujoliswe kuko.
Nangona umsebenzi wangaphambili ubonise ukuba i-PVT inxulunyaniswa nokuchazwa kwenkumbulo yoloyiko, i-45,46,47 asikhange siqaphele naluphi na utshintsho kwinkcazo ye-Fos kwiimpuku xa bekhumbula uloyiko kwiiyure ezingama-48 emva kokumiswa.Lo mahluko unokuchazwa zizinto ezininzi, kubandakanywa uvavanyo lomsebenzi wangaphambili uloyiko lweempawu ezicacileyo kwimeko efanayo apho ukulungiswa kwemeko kwenzeka khona, ngelixa kuvavanyo lwethu, uvavanyo lwenziwa kwigumbi elitsha.Ukongeza, sisebenzise izilwanyana zethu imizuzu engama-60 emva kovavanyo, ngelixa umsebenzi wangaphambili wasebenzisa ixesha lemizuzu engama-90.Ekugqibeleni, kwizifundo zangaphambili, uvavanyo lwenziwa kwigumbi apho izilwanyana zingaphendula ngokutya, kanti emsebenzini wethu, iigundane zavavanywa ngaphandle kwempendulo yesidlo.Ngelixa oku kuvumela inqanaba elithile lothintelo olumiselweyo, kukho ubungqina bokuvumela izilwanyana ukuba zithintele uxinzelelo lokufumana ukutya ngelixa zivavanya ukuba ziyazoyika na izalathiso kudala ungquzulwano lwenkuthazo (okt, uloyiko ngokuchasene nomvuzo), eyona nto iphambili evuselelayo.inxaxheba PVT48, 49. .
I-amygdala ye-basolateral iyaziwa ngokubandakanyeka ekufumaneni ukutshatyalaliswa koloyiko50,51 kwaye kukho ubungqina bokuba uqikelelo lwe-BLA kwi-IL lubandakanyeka kule nkqubo23.Nangona kunjalo, akucaci ukuba i-BLA kunye nokudibanisa kwayo kubandakanyeka kwimbuyekezo yokuphela.Izifundo ze-imaging23,28 zibonise ukwanda komsebenzi we-BLA kwizilwanyana ezikhumbula iinkumbulo eziphelileyo.Ngelixa umsebenzi wethu wangaphambili ubonisa ukuba akukho mahluko ekusebenzeni kwe-BLA phakathi kweegundane zokutshabalala ezilungileyo nezimbi, iziphumo zethu apha zibonisa ukuba ukukhumbula ukutshabalala ngokuqhelekileyo akuchaphazeli i-BLA okanye kusebenze ukuqikelela i-BLA IL.Ngokuhambelana neziphumo zethu, nangona uphando lwe-circuit manipulation lubonisa ukuba i-IL inputs kwi-BLA ibalulekile kwi-extinction learning, ayiyimfuneko ekukhunjulweni kokuphela.Nangona kunjalo, indima ye-BLA ayinakuhoywa ngokupheleleyo njengoko ubungqina bamva nje bubonisa ukuba iintlobo ezithile zeeseli kwi-BLA ziyafuneka ukuba zivelise kwakhona ukuphela.
Okuqaphelekayo, ukukhumbula uloyiko akuzange kubangele ukuba kusebenze i-Fos kwi-BLA, njengoko isilonda sangaphambili, ichiza, kunye nezifundo zokucinga ziye zachaphazela le ndawo ekubonakalisweni koloyiko kunye / okanye ukudibanisa uloyiko kwakhona emva kokufunyanwa54,55,56,57.Idatha eboniswe apha idibanisa i-basal kunye ne-lateral subnucleus ye-amygdala, kunye nedatha yangaphambili ibonisa ukuba ukubonakaliswa koloyiko kuqhuba umsebenzi we-Fos kwicala le-dorsal nucleus esecaleni.Sihlalutye idatha engaphantsi kunye necala ngokwahlukileyo, kodwa akukho mahluko kuyo nayiphi na imeko (idatha engaboniswa) kwaye zombini iindawo ziye zadilika kwiidatha esizibonisa apha.Asizange sihlalutye imimandla engaphantsi kwe-amygdala esecaleni, ngoko ke utshintsho oluthile kule ndawo lunokuthi lufakwe.Enye into enokwenzeka ukuba kungabikho tshintsho kumsebenzi we-Fos kwi-BLA kungenxa yexesha leenkumbulo zoloyiko xa kuthelekiswa nemeko.Umsebenzi othile wangaphambili ubonise ukuba igalelo le-BLA ekubonakalisweni koloyiko liyancipha kunye nexesha emva kokulungiswa, oko kuthetha ukuba i-BLA ixhomekeke kwiiyure ze-24 ze-post conditioning kodwa ezizimeleyo kwiintsuku ze-7 (ref. 45 kodwa bona i-58).kwenzeke kwiiyure ezingama-48 emva koqeqesho, okwenza ukuba ukungabikho kotshintsho kumsebenzi we-Fos ngeli xesha kubonise utshintsho oluxhomekeke kwixesha lokuthatha inxaxheba kwe-BLA ekubonakalisweni koloyiko.
Okokugqibela, sifumana ubungqina bokuba inkumbulo yokuphela kwempumelelo inxulunyaniswa ne-ventral hippocampus.Olu luphawu lwe-vHPC "ephakathi" njengoko iphethini efanayo ayizange ibonwe kummandla ongasemva.Ngokuhambelana nomsebenzi wangaphambili, asifumananga tshintsho kusetyenziso lweFos kwii-vHPC ILs ezingafaniyo.Kukho ubungqina obuninzi bokuba i-vHPC28,60,61 iyadingeka ngenxa yoloyiko lokwenene xa i-CS isenzeka ngaphandle komxholo apho ubumnyama buvela khona, kwaye oku kuxhomekeke ubuncinane kwinxalenye yokungena kwe-vHPC kwi-IL13.Ngokusekwe kwezi ziphumo zangaphambili, sinokulindela ukungafihli kakuhle ukuba kudityaniswe nokonyuka komsebenzi we-IL oqikelelweyo we-vHPC.Nangona kunjalo, oku bekungenjalo njengoko bekungekho mahluko kumsebenzi we-Fos kwi-IL-eqikelelweyo yokubuyisela ebhalwe phantsi ii-vHPCs okanye iiseli ezingabhalwanga kwii-vHPCs.Oku kuphakamisa ukuba ukungakwazi ukukhumbula ukuphelelwa kumxholo wokuphelelwa kunokubangela indlela eyahlukileyo kunoloyiko lokuhlaziya.
Kubalulekile ukuqaphela ezinye zemida yoyilo kunye nohlalutyo kunye nendlela ezichaphazela ngayo izigqibo zethu.Okokuqala, sahlulahlula izilwanyana zibe zisithathu eziphezulu nezisezantsi, kwaye iimpuku zibe "zilungileyo" kunye "ezimbi" ngokusekelwe kumanqaku okukhumbula ukutshabalala.Oku kwenziwa ukuze kuthintelwe iinkqubo zeqela ezahlula izilwanyana zibe ngamaqela ahlukeneyo ukusuka embindini wosasazo, okanye amacebo eqela angazibandakanyi izilwanyana phakathi kosasazo, njengokwahlulwa ngokwemidi okanye ukuthelekisa isithathu esingaphezulu nesisezantsi seempuku. .sifuna ukuphepha le meko kuba ulwahlulo oluphakathi alubonakalisi ukuhlukahluka kweempendulo zabantu kwintlungu esizama ukuyixelisa.Ukongezelela, ngelixa ukuthelekisa iigundane eziphezulu kunye nezantsi ezithathu zivumela ukuba siqhathanise amaqela obungakanani obufanayo, le ndlela iyayihoxisa izilwanyana kwiziko lokusabalalisa kwaye ayibonakali ngokuchanekileyo ukuguquguquka kweendlela zokulimala.Ngelixa indlela yethu inokubandezeleka kwiingxaki kunye nokwahluka okungafaniyo kunye nokuthelekisa amaqela anesampulu ezingalinganiyo, ibamba into esizama ukuyilinganisa ngcono kunezinye iindlela.
Iziphumo ezithiwe thaca apha zisinceda ukuba siqonde ngcono ukuba umahluko womntu ngamnye ekukhunjulweni kokutshabalala ubonakaliswa kumahluko kumsebenzi wesekethe ye-neural.Iziphumo zethu zinokuthi zihambelane nokuphazamiseka kwengxaki emva kokuphazamiseka, eyaziwayo ukuba idibene noloyiko olugqithisileyo kunye nokungakwazi ukuphelisa iimpendulo zokwesaba.Sibonisa ukuba iyantlukwano ekukhunjulweni kokutshabalala kunxulunyaniswa nomahluko kwi-intrinsic kunye ne-extrinsic neural activity eqikelelweyo kwi-IL.Lo mahluko wasasazwa kwimimandla eyahlukeneyo kunye ne-anteroposterior axis, egxininisa ngakumbi ukubaluleka kokuvavanya ukusebenza kwengqondo kwinqanaba elingaphantsi.Ukungalungi kwendlela yangoku kubandakanya ukufaneleka kophononongo kunye nokugxila kwiimpuku zamadoda.Uphando lwexesha elizayo kufuneka lumisele iindlela ze-neurobiological ezisisiseko sokuphela kwentonga kwiimpuku zabasetyhini kwaye zisebenzise iindlela zokuzoba incasa ye-causal.
Iiseti zedatha ezisetyenzisiweyo kunye/okanye ezihlalutyiweyo kuphononongo lwangoku ziyafumaneka kubabhali abachaphazelekayo ngesicelo esinengqiqo.
I-Pavlov IP Conditioning: isifundo somsebenzi we-physiological we-cortex ye-cerebral.(Oxford University Press, 1927).
Rothbaum, BO, & Davis, M. Ukusebenzisa imigaqo yokufunda kunyango lokuphendula emva kokuphazamiseka. Rothbaum, BO, & Davis, M. Ukusebenzisa imigaqo yokufunda kunyango lokuphendula emva kokuphazamiseka.I-Rotbaum BO kunye noDavis M. Ukusebenzisa imigaqo yokufunda kunyango lwe-post-traumatic reactions.I-Rotbaum BO kunye noDavis M. Ukusetyenziswa kwemigaqo yokufunda kunyango lwe-post-traumatic reactions.faka.IKholeji yaseNew York.inzululwazi.1008 (1), 112-121 (2003).
Rauch, SA, Eftekhari, A. & Ruzek, JI Uphononongo lwe-exposure therapy: umgangatho wegolide wonyango lwe-PTSD. Rauch, SA, Eftekhari, A. & Ruzek, JI Uphononongo lwe-exposure therapy: umgangatho wegolide wonyango lwe-PTSD.I-Rauch SA, i-Eftekhari A. kunye no-Ruzek DI Uphononongo lonyango lokuvezwa: unyango olusemgangathweni lwegolide loxinzelelo lwasemva koxinzelelo. Rauch, SA, Eftekhari, A. & Ruzek, JI 暴露疗法回顾：PTSD 治疗的黄金标准。 Rauch, SA, Eftekhari, A. & Ruzek, JI Uphononongo lwe-exposure therapy: Umgangatho wegolide wonyango lwe-PTSD.I-Rauch, i-SA, i-Eftekhari, i-A. kunye ne-Ruzek, i-DI Uphononongo lwe-exposure therapy: unyango olusemgangathweni lwegolide lwe-posttraumatic stress disorder.J. Ukubuyisela kwisimo sangaphambili.uphuhliso lwedama 49, 679–687.https://doi.org/10.1682/jrrd.2011.08.0152 (2012).
Foa, EB Unyango lokuvezwa ixesha elide: elidlulileyo, langoku kunye nekamva.Ukunyanzeliswa koxinzelelo lwe-28, 1043-1047.https://doi.org/10.1002/da.20907 (2011).
UMnu. Millard et al.Izinto ezikhoyo kunye nezifunyenweyo zokulahleka kwememori kwingxaki yoxinzelelo lwasemva koxinzelelo: iziphumo ezivela kuphononongo lwamawele.J. Ugqirha weZigulo zengqondo.itanki yokugcina.42 (7), 515-520 (2008).
UMnu. Millard et al.Isiseko se-neurobiological sokungakwazi ukukhumbula iinkumbulo eziphelayo kuxinzelelo lwasemva koxinzelelo.ibhayoloji.IPsychology.66 (12), 1075-1082 (2009).
Bush, DEA, Sotres-Bayon, F. & LeDoux, JE Ukwahluka komntu ngamnye ekoyikeni: Ukuhlukanisa ukwesaba ukuphinda kusebenze kunye noloyiko lokubuyisela i-phenotypes. Bush, DEA, Sotres-Bayon, F. & LeDoux, JE Ukwahluka komntu ngamnye ekoyikeni: Ukuhlukanisa ukwesaba ukuphinda kusebenze kunye noloyiko lokubuyisela i-phenotypes.Bush, i-DEA, i-Sautre-Baillon, i-F. kunye ne-LeDoux, i-JE Ukwahluka komntu ngamnye ekoyikeni: ukwahlula i-phenotypes ye-reactivity yoloyiko kunye nokubuyiselwa koloyiko. Bush, DEA, Sotres-Bayon, F. & LeDoux, JE 恐惧的个体差异：隔离恐惧反应和恐惧恢复表型。 Bush, DEA, Sotres-Bayon, F. & LeDoux, JE Ukwahluka komntu ngamnye ekoyikeni: ukuhlukaniswa kwempendulo yoloyiko kunye netafile yokubuyisela uloyiko.Bush, i-DEA, i-Sautre-Baillon, i-F. kunye ne-LeDoux, i-JE Ukwahluka komntu ngamnye ngoloyiko: ukuhlukaniswa kweempendulo zoloyiko kunye ne-phenotype yokubuyisela ukwesaba.J. Ukwenzakala.uxinzelelo 20 (4), 413-422 (2007).
I-Russo, i-AS & Parsons, i-RG I-Acoustic impendulo eyothusayo kwiimpuku iqikelela umahluko phakathi kwabantu ngabanye kuloyiko lokutshabalala. I-Russo, i-AS & Parsons, i-RG I-Acoustic impendulo eyothusayo kwiimpuku iqikelela umahluko phakathi kwabantu ngabanye kuloyiko lokutshabalala.I-Russo, i-AS kunye ne-Parsons, i-RG I-Acoustic impendulo eyothusayo kwiimpuku iqikelela ukwahlukana komntu ngamnye kuloyiko lokuphela. Russo, AS & Parsons, RG 大鼠的声学惊吓反应预测恐惧消退的个体差异. Russo, AS & Parsons, RGI-Russo, i-AS kunye ne-Parsons, i-RG I-Acoustic impendulo eyothusayo kwiimpuku iqikelela ukwahlukana komntu ngamnye kuloyiko lokuphela.I-Neurobiology.funda.Inkumbulo.139, 157-164 (2017).
URusso, AS, Lee, J. & Parsons, RG Ukuhluka komntu ngamnye ekukhunjuleni ukutshatyalaliswa koloyiko kuhambelana ne-phosphorylation ye-mitogen-activated protein kinase kwi-cortex ye-infralimbic. URusso, AS, Lee, J. & Parsons, RG Ukuhluka komntu ngamnye ekukhunjuleni ukutshatyalaliswa koloyiko kuhambelana ne-phosphorylation ye-mitogen-activated protein kinase kwi-cortex ye-infralimbic.URusso, AS, Lee, J. kunye noParsons, RG Ukuhluka komntu ngamnye ekuphelisweni koloyiko lokukhumbula ukutshatyalaliswa kuhambelana ne-mitogen-activated protein kinase phosphorylation kwi-cortex ye-infralimbic. Russo, AS, Lee, J. & Parsons RG Russo, AS, Lee, J. & Parsons, RG Inkumbulo yoloyiko oluphelayo inxulumene nomahluko ngamnye kwiperipheral 美裯中丝裂原活化筒能激酶的phosphorification.URusso, AS, Lee, J. kunye noParsons, RG Ukwahluka komntu ngamnye ekuphelisweni koloyiko ngexesha lokukhumbula kuhambelana ne-phosphorylation ye-mitogen-activated protein kinases kwi-cortex yomlenze ophantsi.I-Psychopharmacology 236(7), 2039-2048 (2019).