LITABA: Re khethile thuto ena hobane ke e 'ngoe ea tsa morao-rao. Ho nkoa ke hore lintho tse peli tse ncha le tse tšosang li etsa hore ho be le mehopolo e matla le ho ithuta.
Tšabo ke tlhaloso e akaretsang ho mahlale. Ha ho tluoa litabeng tsa bootsoa, eng kapa eng e tšosang kapa e hlahisang matšoenyeho e tla phahamisa epinephrine (adrenaline) le norepinephrine (noradrenaline) mme e thuse ho theha lisekete tse ncha tsa memori. Motsoako oa bonohe (dopamine) le "ts'abo" li khothatsa haholo potoloho ea moputso. Motsoako ona o bakile kholo ea mefuta e fapaneng ea litšoantšo tsa bootsoa.
Ho ithuta ka botlalo le litšoantšo
inahaneloang
Stanley O. King II le Cedric L. Williams
Ho ipapisa le maemo a machaba ho hlahisa linaha tse phahameng tsa liphetoho tse tsosang takatso le tsa likokoana-hloko bokong ho matlafatsa mohopolo. Leha ho le joalo, lits'ebetso tse lumellang ho pepeseha habonolo maemong a sa tloaelehang ho phahamisa tlhahiso ea kutloelo-bohloko le ho ntlafatsa mohopolo ha li utloisisoe hantle. Phoso ena e ile ea rarolloa ka ho lekola hore na liphetoho tse hlohlelletsoang ke bocha litleneng le / kapa bohareng li tsosa mohopolo bakeng sa boemo ba tšabo ba Pavlovia. Likhoto tsa banna li ne li pepesetsoa phapusing ea maemo bakeng sa 5-min kapa ha li a ka tsa pepesetsoa 24 h pele li ka ts'oaroa ka lipara tse hlano tsa molumo o nyarosang (0.35 mA). Ho boloka ho ile ha hlahlojoa 48 h hamorao maemong a fapaneng. Liphoofolo tse sa pepesang li bonts'itse leqhoa le leholo haholo nakong ea lipontšo tsa maemo a hlasimollang (CS) ho feta liphoofolo tse pepesitsoeng pele (P <0.05). Ntlafatso ea poloko e hlahisoang ke boqhetseke e ile ea fokotsoa ka ho qhekella thibelo ea ip-adrenergic receptors ka sotalol (6 mg / kg, ip). Phuputso ea 2 e senotse hore keketseho e susumetsoang ke bocha ea tlhahiso ea boipheliso e fetisetsoa bokong ke li-visceral afferents tse lumellanang le li-brainstem neurons ka har'a nucleus tractus solitarius (NTS). Ho thibela ts'ebetso ea AMPA receptor ho NTS le CNQX (1.0 μg) ho fokolitse haholo ho hatsela ho CS ho liphoofolo tse sa pepesang pele (P <0.01). Boithuto ba 3 bo bonts'itse hore ho phahamisa maemo a epinephrine liphoofolong tse tloaetsoeng ho susumetsa ho ithuta ka mekhoa e ts'oanang le e hlahisoang ke ho tsoha ho sa tloaelehang. Liphoofolo tse pepesitsoeng pele tse fuoeng epinephrine (0.1 mg / kg) li ile tsa hatsela haholo ho feta taolo ea letsoai (P <0.01), mme phello ena e ile ea fokotsoa ke ho kenella kahare ho NTS ea CNQX. Liphuputso li bonts'a hore tsoho e entsoeng ka mokhoa o ts'oanang le o mong kapa ho eketsa ts'ebetso e nang le kutloelo-bohloko e nang le epinephrine liphoofolong tse sa hlahisoang pele ho ntlafatsa mohopolo ka mekhoa ea adrenergic e qalileng sebakeng se pota-potileng le ho fetisetsoa ka har'a naha ka tataiso ea vagus / NTS.
Tsa Sehlooho se Ithutoang
Palo e hlahelang ea liphuputso e senola hore mokhoa o sa tloaelehang o amanang le maemo a sa tsejoeng kapa melaetsa e sa tloaelehang ea tšusumetso e simolla ka bobeli liphetoho tsa cellular le tsa 'mele tse lumellanang le likarolo tsa liketsahalo tse ncha ka mohopolo. Ntho e ikamahanyang ea ho itlhahisa ka tsela e ts'oanang le e ncha ho latela mekhoa e laolang mohopolo le synaptic plasticity e hlokomeloa nakong ea 3 wk postnatal (Tang le Reeb 2004) mme e ngotsoe lipotong tse tsofetseng tse lekiloeng ho feta 22 ea lilemo (Sierra-Mercado et al. 2008) . Tšusumetso ea moqoqo oa libuka ho matlafatsa litšoantšiso tse ncha e ka amahanngoa le karolo ea matla a bona a ho qalisa ho fetoha ha liphetoho tsa tlhaho ea likokoana-hloko tse hlokahalang bakeng sa mohopolo oa nako e telele.
Ts'ebeliso ea mekhatlo e mecha e latelang thuto e arolelanoa ke karolo ea phosphorylation ea likarabo tsa likarabo tse kopanyang liprotheine (CREB) le liphatlalatso tsa liphatsa tsa lefutso tse entsoeng ke CRE ho kopanya likarolo tse ling tsa liketsahalo tse ncha molokong oa mohopolo (Alberini 2009). Pōpo ea phosphorylation e phahame ka har'a hippocampus ka mor'a hore sebaka se lule se le teng, 'me mohato ona oa bohlokoa oa ho hopola mohopolo o tsoela pele ka mor'a hora e latelang phihlelo ea libuka empa ha o fetohe litabeng tse hlalositsoeng moelelong o tloaelehileng (Kinney le Routtenberg 1993; Viola et al . 2000; Izquierdo et al. 2001). Ho hlahisa liphoofolo ho e-na le taba e ncha ho etsa hore ho be le mefuta e mengata ea liphatsa tsa lefutso tsa pele tsa tlhaho c-fos le c-jun ka amygdala le hippocampus, empa liphetoho tsena ha li hlokomeloe ka lihlopha tse khutlisetsoang hape, kapa li lumelloa ho hlahloba maemo a tloaelehileng (Papa le al. 1993; Zhu et al. 1997; Sheth et al. 2008). Liphello tse tšoarellang tsa likarolo tse khutšoanyane tsa ho ts'oara lihlahisoa tse tsosang takatso le tse hlokomelehang li boetse li lekaneng ho ntlafatsa ho khutlisetsoa mohopolong o hōle (Izquierdo et al. 2000, 2003) le ho matlafatsa mohopolo tlasa maemo a bonolo a koetliso a atisang ho fana ka ts'ebetso ea boipheliso bo futsanehileng. Moncada le Viola (2007) li bontšitse hore koetliso ea ho qoba ho itšireletsa ka ho ipeha kotsing e fella ka ho hopola lintho tse fokolang kapa tse se nang boikutlo ho laola 24 h hamorao. Leha ho le joalo, batho ba pepesehileng ka taba e sa tloaelehang kapa pele ba koetlisoa le batho ba fokolang ba fokolang ba bontšitse hore ba bolokehile haholoanyane ho latela litaelo ha mohopolo o hlahlojoa 24 h hamorao.
Ho behoa tikolohong e ncha pele ho kenyelletsoa nako e telele (LTP) e nang le bofokoli bo fokolang bo fokolang bo fokolang, bo thusa ho fetela pele ho LTP ho ea morao ho LTP, e leng ho hlokahalang hore ho be le protheine synthesis, le mokhoa ona oa ho lekola nako e telele bakeng sa tlhokomelo ea LTP. ho tloha 8 ho ea 24 h (Li et al. 2003; Straube et al. 2003a, b). Liphello ha li hlokomeloe haeba LTP e qalisoa ka maemo a koetliso a fetoletsoeng a tloaelehileng ka lebaka la tloaelo ea nako e telele. Ho thahasellisang, ho thibela li-receptor noradrenergic tse nang le tšelo-pele ea pirate ea propranolol pele ho kenngoa moetong o sa tloaelehang ho thibela ntlafatso e kenyelletsoeng ea LTP, e bontšang karolo ea norepinephrine ho phelisang liphello tsa morao-rao bokong (Straube et al. 2003a). Ho kenngoa ha neurotransmitter ena ho boetse ho fanoa ka liphuputso tse bontšang li-neurons tsa locus coeruleus (LC) tse fanang ka norepinephrine ho li-forebrain le libopeho tsa maoto li bonts'a tšebetso ea tšebetso ha li qala ho pepesehela tikoloho e ncha, empa ho ts'oloha ho eketsehileng ha ho hlahisoe ke likhoto ho motho ea tloaelehileng moelelo (Vankov et al. 1995). Lintho tse ling tse fumanoeng, ho tlaleha hore libaka tsa norepinephrine tse ka pele ho cortex le hypothalamus li phahame ka ho fetisisa ka mor'a ho pepesehela tikoloho e hlahisitsoeng ke lirafshoa kapa maemo a koetliso a nang le rat e sa tloaelehang (McQuade et al. 1999), e fana ka bopaki bo tobileng ba hore norepinephrine e kopanya liphetoho tse kholo ho hlahella habonolo. Liphuputso tsena tse bonngoeng li bonts'a hore ts'ebetso e ncha e bakoang ke ho ipapisa le maikutlo a sa tloaelehang ho susumetsa liphetoho tse 'maloa tsa methapo ea meriana le ea synaptic tse hlokahalang bakeng sa liphihlelo tse ncha tse kenyelletsoeng ka katleho holellong ea nako e telele.
Liphello tsa ho pepesehela maemong a sa tloaelehang ha li felle feela liphetohong tse nang le liphetoho tse entsoeng bokong. Litlhaloso tse ikemetseng tsa ts'ebetso ea kutloelo-bohloko, tse kenyelletsang letlalo, boitšoaro ba pelo le ho potoloha ha li-hormone tsa correno corticosterone le epinephrine, kaofela li phahame ka ho hlahisa batho kapa liphoofolo tse nang le tšusumetso e ncha kapa ka mor'a ho lumella lipatlisiso tsa mahala sebakeng se sa tloaelehang (De Boer et al. 1990; Bradley et al. 1993; Handa le al. 1994; Gerra le al. 1996; Codispoti et al. 2006). Liphuputso tsena li senola bohlokoa ba pakeng tsa sehlopha sa liphetoho tsa 'mele tse hlahelang ka ho toba ka ho pepesehela tikoloho e ncha le tse hlahisitsoeng ke liketsahalo tse tsosang maikutlong. Le hoja maemo a mabeli a etsa liphetoho tse fokolang tšebetso ea li-visceral le boko bo hlahisoang ke boko ba ho bokella liketsahalo tse ncha mohopolong, mekhoa eo ho eona ho nang le lits'ebeletso tse nkiloeng ka morao-rao le / kapa tsoseletso e bohareng e ka 'nang ea susumetsa mohopolo oa mehopolo ha e utloisisoe ka botlalo.
Lintlha tse 'maloa tsa bopaki li fana ka maikutlo a hore hormone e amanang le tsoalo e epinephrine e bapala karolo ea likarolo tse peli. Ka mohlala, ente ea tsamaiso ea epinephrine ka meriana e mengata e ntlafatsang mohopolo oa likhoto tsa laboratori (E leng Williams le McGaugh 1993; Clayton le Williams 2000; Nordby et al. 2006; Dornelles le al. 2007) ho eketsa lebelo la ho thunngoa ha LC neurons (Holdefer le Jensen 1987) e bontšang ho hlahisa maemo a phahameng ka mor'a hore ho be le maemo a macha (Vankov le al. 1995). Joalo ka mokhoa o hlophisitsoeng, tsamaiso ea epinephrine e thusa ho thusa LTP (Korol le Gold 2008) le ho fokotsa litšenyehelo ho boloka boemo bo tšabehang ba boemo bo bontšitsoeng ke litoeba tse nang le ntho e ngotsoeng ea CREB liphatseng tsa lefutso (Frankland et al. 2004). Tlhaloso ea li-slide tse bonts'itsoeng ho batho e ntlafatsa mohopolo (Fenker et al. 2008) mme e qalisa ho thibela epinephrine ho tswa ho adrenals (Gerra et al. 1996), 'me phetoho ena ea ho tsosoa e lekane ho ntlafatsa ts'ebetso ea morao-rao (Cahill et al. 1994) e tšoanang le e hlahisoang ke tsamaiso e tobileng ea hormone ena (Cahill le Alkire 2003). Tšusumetso e tsosolositsoeng e tsosolositsoeng mohopolong oa batho ka li-slide tse bonts'ang (Strange le Dolan 2004) le ts'ebeliso e entsoeng ka mokhoa o tsitsitseng oa LTP eo ho buuoeng ka eona ka holimo (Li et al. 2003; Straube et al. 2003a, b) li emetsoe ka ho thibela noradrenergic receptor phetiso le li-β-adrenergic receptor mohanyetsi oa propranolol. Mefuta ena ea liphuputso e fana ka motheo oa ho khetholla hore na ho tsosoa ha tlhaho ho tsoaloang le liphetoho tse latelang tsa 'mele tse thusang ho kenyelletsa likarolo tsa liphihlelo tse ncha mohopolong li kopanngoa ka ho sebelisana ho amanang le lisebelisoa tsa hormonal tsa phallo tse susumetsang mosebetsi oa noradrenergic bokong.
Haeba nako e khutšoanyane ea tlhaho ea tlhaho ea tlhaho e etsa hore motho a tsosolose ka mokhoa ona, joale hoa utloahala hore mokhoa o mong oo ho oona ho tsosang takatso ho amang matla a maikutlo a maikutlo ao a a bolokang ho a hopola ke ka ho etsa hore litsela tsa neural li tsamaise liketso tsa sympathomimetic tsa epinephrine tse kopanetsoeng mokokotlong ho ea boko mekhoa e susumetsang norepinephrine e hlahisitsoeng CNS. Makala a sephahla a vagus a phetha karolo ea bohlokoa ts'ebetsong ena ha litebelisoa tse nyolohang tsa vagus li kenngoa haholo le li-ß-adrenergic receptors tse tlama epinephrine (Schreurs et al. 1986; Lawrence et al. 1995), le ho fela ha liphahla tsa vagus litho tsa kutloelo-bohloko tse amohelang haholo ho tsosoa ke kutloelo-bohloko e hlahisoang ke ho lokolloa ha epinephrine kapa mokhoa o sa tloaelehang, ho akarelletsa le pelo, sebete, mala le matšoafo (Shapiro le Miselis 1985, Coupland et al. 1989; Paton 1998a, b). Ho feta moo, ts'ebetso ea motlakase ea likhohlopo tsa vagal e nyoloha ho hlahisa ho phatloha ho hoholo ho LC neurons (Groves et al. 2005; Dorr le Debonnel 2006) mme ho lebisa linthong tse tšoarellang nako e telele ho li-norepinephrine tse bokelloang ho tloha ho amygdala (Hassert et al. 2004) le hippocampus (Miyashita le Williams 2002).
Tlhahisoleseding mabapi le mosebetsi o tsitsitseng o sebetsanang le lits'ebeletso tsa ponahalo ea meriana o fetisetsoa ka ho nyoloha ka lihlopha tse ling tsa lisele tse ling tse tsejoang e le mokokotlo oa pampiri (NTS) (Kalia le Sullivan 1982; Sumal et al. 1983). Ka lebaka la liphetoho tsena, li-NTS neurons li susumetsa tšebetso e ka sehloohong ea li-synapses ka LC neurons (Van Bockstaele et al. 1999) e seng feela e sebetsang ka pel'a botumo ba libuka (Vankov et al. 1995) empa hape e lokolla norepinephrine mehaho e phethang karolo ea bohlokoa ho kenyelletsa liphihlelo tse ncha ho hopola nako e telele tse kang khetla ea prefrontal cortex, hippocampus le amygdala (Ricardo le Koh 1978; Loughlin et al. 1986; Florin-Lechner et al. 1996).
Haeba ts'oaetso e susumelitsoeng ke mokhoa o tsosolositsoeng e eketsa tšireletso ea epinephrine, joale hoa utloahala hore tsela e 'ngoe eo ka eona ho tsosoa ho amanang le matla ao likarolo tsa maikutlo li bolokiloeng ka nako e telele ke ho etsa sena ka tsela ea vagal / NTS. Thuto ea hona joale e leka liteko tsena ka ho sebelisa "tloaelo" ho latela "mokhoa o sa tloaelehang" oa moelelo oa koetliso e le ho qhekella ho eketsa tsosoloso ea mmele pele o ithuta le ho hlahloba hore na ho boloka mehopolo ea lintho tse amanang le maikutlo ho susumelitsoe ke ts'ebetso ea phallo ea adrenergic. Boemo ba tšabo ea li-Pavlovia bo atisa ho sebelisoa ho utloisisa lipotoloho tsa neural tse amehang ho hopola lintho tse tsosang takatso (Kim le Jung 2006), le hoja liphello tsa ho tsosolosa ho tsosoa ha lintho tsa tlhaho nakong ea ho thehoa ha mohopolo oa tšabo li sa hlahlojoe ka ho pharaletseng.
Ka lebaka la tlhaloso ena, lipatlisiso tsena li ile tsa hlahloba kamoo liphetoho tsa ponahalo ea 'mele li fetisetsoang ka mokhoa oa vagus / tataiso ea NTS ho senola lits'ebetso tseo ho tsosolosoa ho tsona ho susumetsang mohopolo ka lebaka la tšabo. Sepheo sa liteko 1 e ne e le ho hlahloba monehelo oa poneletso ea adrenergic ho ts'oanela ho tsosolosa ho tsosoa ha lintho tse ncha le liphello tse latelang ho sebetsanang le maikutlo. Phuputsong ena, mokhoa o hlophisitsoeng o ile oa kenngoa ka lihlopha tse fapaneng ka ho tlohela tloaelo le ho emela ho fihlela letsatsing la boemo ba ho hlahisa lihlooho moetong oa koetliso ka lekhetlo la pele. Liphello tsa ho thibela li-receptor tsa piratori pele ho Plolovian conditioning li ne li hlahlojoa ka lihlopha tseo ho tsona moetlo oa koetliso o neng o emela boikutlo ba tlhaho le ho bapisoa le lihlopha tse neng li tloaelane le kamore ea ho tšaba ka mekhoa e fetileng. Thuto 2 e ile ea hlahloba hore na tsela e pakeng tsa phallo ea liphahla-khoeli le likopi tsa kelello ea NTS e fana ka liphello tsa ho eketseha ho bakoang ke mokhoa oa ho etsa lintho ka tsela e nang le kutloelo-bohloko nakong ea boemo ba tšabo. Amino acid glutamate ke moemeli oa motlakase o sebelisanang lipuisano tsa synaptic pakeng tsa vagal afferents le li-neurons tsa NTS ho tloha ha li-terminals tsa vagal li na le glutamate (Sykes et al. 1997) le li-receptor tsa glutamate li sebakeng sa NTS dendrites (Aicher et al. 1999, 2002). Ho phaella moo, ho kenngoa ha intra-NTS ea li-AMPA glutamatergic bahanyetsi ba liphallelo CNQX (6-cyano-7-nitroquinoxaline-2,3-dione) e thibela ho phatloha ho feteletseng ho NTS neurons e susumelitsoeng ke ho susumetsa methapo ea vagus (Granata le Reis 1983a; Andresen le Yang 1990 ) le mefuta e mengata ea maqhubu a eketsang phekolo ea LC (Groves et al. 2005; Dorr le Debonnel 2006) kapa ho khoneha ho lokolloa ha norepinephrine ho amygdala kapa hippocampus (Miyashita le Williams 2002; Hassert et al. 2004). Ho finyella sena, motsoako oa AMPA oa AMPA o ne a sebelisetsoa ho thibela batho ba nang le li-receptor tsa postsynaptic sebakeng sa NTS se amohelang maikutlo ho tloha ho li-terminals tsa vagal. Sebaka sa lisebelisuoa tsa manyolo a li-cannulae le a entsoeng a lebisitsoeng ho NTS e hlalositsoe setšoantšong sa 1.
[Setšoantšo sa 1.]
Thuto 3 e hlokometse hore na mohopolo o futsanehileng o bontšitsoeng ke lihlopha tse laolang tloaelo tse tloaelehileng molemong oa koetliso o ka ntlafatsoa ka ho eketsa ts'ebetsong ea lits'ebeletso ka mor'a hore maphelo a Pavlovian a hlajoe ke epinephrine. Phuputso ena e boetse e reretsoe hore phetisetso ea glutamatergic pakeng tsa vagal afferents le NTS neurons e phetha karolo ea bohlokoa ho thuisa liphetoho tse tobileng mohopolong o hlahisoang ke epinephrine e phahameng. Liphuputso tse hlahang liphuputsong tsena li senola hore ts'ebetso e ts'oanang ea tikoloho kapa ka ho ts'oanela ho etsa mesebetsi e nang le kutloelo-bohloko e nang le epinephrine e ntlafatsa mohopolo oa tšabo oa pelovian ka mekhoa ea adrenergic e qalileng ka thōko le ho fetisetsoa ka har'a naha ka mokhoa oa vagus / tataiso ea NTS.
Results
Leka 1
Koetliso ea boemo bo tšosang
Phuputso ena e ile ea ikemisetsa haeba ntlafatso ea mohopolo e hlahisoang ke tlhaho e hlaheletseng le ho latela koetliso ea tšabo ea tšabo ea Pavlovian e sebelisana le ho etsa lisebelisoa tsa lits'ebeletso tsa pone ea methapo. Ho ne ho tšeptjoa hore sephiri sa epinephrine e ne e tla ba motsoako o hlokahalang bakeng sa tsosumetso e hlahisoang ke batho ho ntlafatsa mohopolo. Tlhahiso ena e ne e hlahlojoa ka ho sebelisa li-β-adrenergic receptor antagonist sotalol ho thibela epinephrine ho tlamella li-receptor tsa li-ß-adrenergic tse ling likokoanyana tse tšoaetsanoang le maemo a boemo ba libuka.
Tloaelo e 'meli ea ANOVA ka palo e boleloang ea ho qhoqhoa, e bonts'itsoe tlhahiso ea ho qetela ea mohopolo (CS; tone) nakong eo e fumanoang ka liphallelo tse hlano tsa CS-tse sa lumellaneng, tse bontšitsoeng phapang pakeng tsa lihlopha tsa phekolo bokhoni ba ho ithuta hore mohala oa CS ke moeletsi ea ka tšeptjoang oa ho qhoqhoa ha US ebile o etsa hore ho be mafifi, F (1,20) = 1.48, P = NS (pele ho pepenene / saline 88.38 ± 7.3, pele ho pepenene / sotalol 90.68 ± 4.0, pele ho pepenene / saline 97.28 ± 2.0, e sa hlahang pele / sotalol 84.16 ± 6.9).
Tlhahlobo ea ho boloka
ANOVA ea litsela tse peli e bonts'itse phello ea bohlokoa ea kalafo ho liperesente tse bolelang tsa ho hatsela ho bontšitsoeng nakong ea lipontšo tse tharo tsa CS nakong ea liteko tsa ho boloka kamoreng e fapaneng ka ho felletseng ea Pavlovian (F (1,20) = 21.26, P <0.01; Fig. 2A). Liteko tsa Post-hoc li senotse hore liphoofolo tse sa pepesetsoang li bonts'a leqhoa haholo nakong ea lipuo tsa CS ho feta liphoofolo tse tloaetseng ho pepesetsoa kamoreng ea maemo 24 h pele ho thupelo (P <0.05). Ntle le moo, liphoofolo tse sa pepesang pele, tse tsamaisang antagonist sotalol ea β-adrenergic receptor, li bonts'a leqhoa le fokolang haholo nakong ea lipuo tse tharo tsa CS mabapi le liphoofolo tse sa pepesang tse fuoeng liente tsa saline (P <0.01). Tlhatlhobo ea molumo oa molumo oa leqhoa le li-ANOVA tsa thuto e bonts'itse hore litaba tse sa pepesoang li bonts'a boemo bo phahameng haholo ba ho hatsela ho polelo e ngoe le e ngoe ea molumo e amanang le lihlopha tse ling tsohle (bona setšoantšo sa 2B). Kahoo, ho tsoha ho tlalehiloeng ho amanang le ho beha lintho tse phelang moleng oa lipale (De Boer et al. 1990; Handa et al. 1994) e kenya letsoho kholisong e ntlafalitsoeng ea ho ithuta ka maikutlo. Ho feta moo, litlamorao tse ntle tsa ho tsosa mohopolo bakeng sa li-pairings tsa CS-US li ipapisitse le ts'ebetsong ea lits'ebetso tsa "hormone" tse amanang le li-receptor tsa β-adrenergic.
[Setšoantšo sa 2.]
(A) Lihlopha: Pheripheral β-adrenergic blockade e nang le sotalol (4 mg / kg) e sitisa ntlafatso ea mohopolo e sa tsoa etsoa. Liphoofolo tse sa pepesang tse fuoeng ente ea letsoai pele e ka lekoa ka phapusing ea nalane li bonts'a liperesente tse kholo tsa ho hatsela (ke hore, 87%) nakong ea liphatlalatso tsa CS ha li bapisoa le lihlopha tsohle tsa liteko (* P <0.05). Ho thibela li-receptors tsa β-adrenergic ka lehlakoreng le sotalol pele ho boemo bo ka kamoreng ea lipale ho fokolitse haholo palo ea leqhoa (ke hore, 49%) e hlahisitsoeng ke tlhahiso ea CS nakong ea liteko tsa ho boloka (** P <0.01). Liphoofolo tse mashome a mabeli a metso e mene li ne li arotsoe ka lihlopha tse latelang tsa kalafo (pre-exposure-saline, n = 6; saline e sa pepesetsoang, n = 5; sotalol e senotsoeng pele, n = 8; le sotalol e pepesitsoeng pele , n = 5). (B) Liteko tsa ho boloka litšebeletso: Kerafo ea mohala e bonts'a leqhoa le lekang ho liteko tsa CS nakong ea liteko tsa ho boloka. Liphoofolo tse sa pepesang tse tšoaroang ka liente tsa ip saline pele ho boemo bo ka kamoreng ea lipale li bonts'a leqhoa le phahameng ho feta lihlopha tse ling tsohle nakong ea tlhahiso ea CS ea pele (* P <0.05). Peresente ea ho hatsela sehlopheng sena nakong ea liphatlalatso tse latelang tsa CS le eona e ne e le kholo haholo ho feta sehlopha ka seng sa kalafo (** P <0.01). Pheripheral β-adrenergic blockade e nang le sotalol (4 mg / kg) e fokotsitse phello ea ponahalo e ncha ea ho pepesetsoa ha serame se ntlafalitsoeng sa molumo oa CS.
Leka 2
Koetliso ea boemo bo tšosang
Phuputso ea bobeli e ile ea hlahloba hore na liphetoho tsa 'mele tse susumelitsoeng sebakeng se itseng ka ho pepesehela tikoloho e ncha li ama mohopolo oa tšabo ka ho etsa li-neurons NTS. Ho ne ho boletsoe esale pele hore tsoho e entsoeng ka tsela e ncha e bontšoang ke ts'ebetso e ikemetseng e eketsa mohopolo ka epinephrine e tlamella li-receptor β-adrenergic hammoho le likhoele tse nyolohang tsa methapo ea vagus. Ho fetisetsa ho eketsehileng ka vagus ho tla etsa hore ho be le methapo e ntle ea NTS e sa hloekisoang ke li-terminals tse hlahisang glutamate. Ho fanoe ka maikutlo ana, ho thibela tšebetso ea amohelehang ea AMPA e amanang le ho lokolloa ha glutamate NTS hang-hang ho phomola ha liphoofolo bakeng sa liphoofolo tse sa tsejoeng pele ho tsona ho lokela ho hlokomoloha ho ntlafatsa mohopolo ho tloha ho tsosolosoa ho susumetsoang ke mocha. Liphuputso tse qalileng tsa thuto ena li bontšitse hore ha ho na phapang pakeng tsa lihlopha tsa phekolo ka matla a bona a ho ithuta mekhatlo ea CS-US nakong ea thupelo. Lihlopha tsohle li ile tsa bontša mahlaseli a tšoanang le a mangata ho fihlela qetellong ea tlhahiso ea CS nakong ea boemo, F (1,25) = 0.670, P = ns (pele ho pepesa / PBS 92.0 ± 5.0, pele ho pepenene / CNQX 86.1 ± 5.2, PBS 96.0 ± 2.2, e sa hlahang pele / CNQX 96.3 ± 1.1).
Tlhahlobo ea ho boloka
ANOVA ea litsela tse peli e senotse litlamorao tse kholo ho liperesente tse bolelang tsa ho hatsela ho li-CS tse tharo tse hlahisitsoeng nakong ea liteko tsa ho boloka, F (1,25) = 9.60, P <0.01. Joalo ka teko ea 1, liphoofolo tse sa pepesetsoang li fuoe liente tsa koloi ho NTS ho bata bakeng sa liperesente tse phahameng haholo tsa nako ha CS e ne e hlahisoa ha e bapisoa le litsamaiso tse pepesitsoeng pele le liphoofolo tse pepesitsoeng tse tsamaisang CNQX ho NTS (P <0.01 ; Setšoantšo sa 3A). Liphetho tsa post-hoc li bonts'itse hore infusion ea linaha tse peli ea CNQX ho NTS e fokolitse haholo liperesente tse phahameng tsa ho hatsela ho bonoang liphoofolong tse sa pepesoang maemong a neng a ka bapisoa le a litsamaiso tse pepesitsoeng pele (P <0.01). Setšoantšo sa 3B se bonts'a palo ea ho hatsela nakong ea e 'ngoe le e' ngoe ea lipontšo tse tharo tsa molumo oa CS. Pontšong ea pele feela ea CS, liphoofolo tse tšoaroang ka letsoai tse sa pepesang li ile tsa hatsela haholo ho feta liphoofolo tse tšoaroang pele tsa CNQX (P <0.02), empa eseng lihlopha tse pepesitsoeng pele. Sehlopha se neng se sa pepesoa se ile sa hoama haholo ho feta lihlopha tsohle nakong ea lipuo tsa bobeli le tsa boraro tsa CS (P <0.01). Liphuputso tsena li bonts'a ntlafatso e susumetsoang ke mohopolo mohopolong oa maemo a tšabo a Pavlovia e fokotsoa ka ho thibela phihlello ea li-postynaptic glutamate receptors ho NTS ka tšoaetso ea linaha tse ling tsa mohanyetsi oa AMPA ea amohelang CNQX.
[Setšoantšo sa 3.]
(A) Lihlopha: CNQX (1.0 μg) thibelo ea phetisetso ea glutamatergic kahara khubu ea pampitšana e le mong (NTS) e thibela ntlafatso ea mohopolo e ncha. Sehlopha se senotsoeng pele se fuoeng koloi NTS se bonts'a liperesente tse phahameng haholo tsa leqhoa ho feta lihlopha tsohle tsa liteko nakong ea lipuo tse tharo tsa CS tse fanoeng nakong ea tlhahlobo ea 48 ea ho boloka (** P <0.01). Ntlafatso ea mohopolo e hlahisitsoeng ke boiphihlelo ka nako ea maemo e ile ea fokotsoa ka ho thibela li-receptor tsa AMPA ho NTS le CNQX. Sehlopha se sa pepesang-CNQX se bonts'itse mohopolo o futsanehileng haholo bakeng sa CS o bonts'itsoeng ka ho fokotsa ho hatsela ho CS mabapi le sehlopha se neng se sa pepesoa se fuoeng PBS ho NTS (* P <0.05). Lihlooho tse mashome a mabeli a metso e robong li arotsoe ka lihlopha tse latelang tsa kalafo (PBS e senotsoeng pele, n = 8; PBS e sa pepesang pele, n = 8; CNQX e senotsoeng pele, n = 6; le CNQX e sa pepesang pele, n = 7). (B) Liteko tsa ho boloka litšebeletso: Kerafo ea mohala e bonts'a leqhoa le lekang ho liteko tsa CS nakong ea liteko tsa ho boloka. Lihlooho tsa sehlopha sa letsoai le neng le sa senoloa pele li bontšitse boemo bo batang haholo ba ho hatsela ho feta lihlopha tse ling tsohle nakong ea tlhahiso ea bobeli le ea boraro ea molumo CS (** P <0.01). Boemo bo phahameng ba leqhoa le bontšitsoeng ke bafo ba seng pepeneneng bo ile ba fokotsoa ka ho thibela li-receptor tsa AMPA ho NTS le CNQX.
Leka 3
Koetliso ea boemo bo tšosang
Phuputso ea ho qetela e ile ea hlahloba hore na likhahla tse ngata tse potolohang tsa epinephrine li ntlafatsa boemo ba Pavlovian ka mekhoa e tšoanang le e hlahisoang ke tsoho e entsoeng ka mokhoa o ncha. Haeba mefuta e 'meli ea litsela e arolelana litsela tse tšoanang, joale liphetoho leha e le life tsa maemo a lumellanang le epinephrine li lokela ho fokotseha ka ho senya mokhoa o tšoanang oa NTS e bontšoang e le oa bohlokoa bakeng sa tsosumetso e susumelitsoeng ke morao ho ama mohopolo. Tlhahiso ena e ile ea hlahlojoa ka ho kenella motseng oa CNPX oa mohanyetsi oa AMPA NTS ~2 min pele ho tsamaiso ea epinephrine (0.1 mg / kg). Liphekolo tse peli li ile tsa fanoa ka mor'a hore li sebetsane le maemo a mahlano a CS-US. Lihlopha tsohle tsa pepese tse pepeneneng li bonts'a likaroloana tse tšoanang tsa ho qhoqhoa ho fihlela tlhahiso ea ho qetela ea CS nakong ea boemo, F (1,26) = 0.057, P = NS (saline / PBS 94.6 ± 3.1, saline / CNQX 92.5 ± 4.0, epinephrine / PBS 97.9 ± 2.1, epinephrine / CNQX 94.3 ± 4.0).
Tlhahlobo ea ho boloka
ANOVA ea litsela tse peli e bonts'itse phello e kholo ea liperesente tse bolelang tsa ho hatsela ho bonts'itsoeng ho linehelano tse tharo tsa CS nakong ea liteko tsa ho boloka ts'ireletso, F (1,26) = 12.13, P <0.01. Liteko tsa Post-hoc li senotse hore liphoofolo tse pepesitsoeng pele li fuoe infra-NTS infusion ea PBS le ente ea systemine ea epinephrine e bonts'a liperesente tse phahameng haholo tsa ho hatsela ho CS ha li bapisoa le lihlopha tse ling tsohle tsa kalafo (P <0.01). Leha ho le joalo, liphoofolo tse pepesitsoeng pele tse fuoeng tekanyetso e tšoanang ea epinephrine kamora ho ts'oaetsoa ha linaha tse peli tsa CNQX ho NTS li ne li sa khethollehe ho tsoa ho taolo e kenngoeng ke PBS (P = NS; Setšoantšo sa 4A). Setšoantšo sa 4B se bonts'a palo ea ho hatsela nakong ea e 'ngoe le e' ngoe ea lipontšo tse tharo tsa molumo oa CS. Sehlopha sa epinephrine (0.1 mg / kg) se ile sa hoama haholo ho feta taolo ea letsoai nakong ea tlhahiso ea pele ea CS (P <0.05), empa liperesente tsa bona tsa ho bata li ne li sa fapane le tsa lihlopha tsa CNQX. Leha ho le joalo, nakong ea tlhahiso ea bobeli le ea boraro ea CS, liphoofolo tse tšoaroang ke epinephrine li ile tsa hatsela haholo ho feta lihlopha tse ling tsohle tsa kalafo (P <0.01). Ntlafatso e hlahisitsoeng ke epinephrine mohopolong oa mekhatlo ea molumo oa molumo e ile ea fokotsoa ke thibelo ea CNQX ea li-AMPA receptors ho NTS kaha maemo a ho bata hoa sehlopha sena a ne a sa fapana haholo le taolo ea letsoai. Liphuputso tsena li fana ka maikutlo a hore sebopeho sa vagus / NTS ke karolo ea bohlokoa ea mekhoa ea ho fetisa maemo a phahameng a ho tsoha ha mmele a hlahisoang ke boiphihlelo bo matla ba maikutlo ho sistimi ea boko e kentsoeng le ho boloka mohopolo bakeng sa maemo a tšabo.
[Setšoantšo sa 4.]
(A) Lihlopho tse pepesitsoeng pele: Ho hanyetsa li-receptor tsa AMPA ho NTS ho thibela ho nolofalletsa ho etsa epinephrine maemong a tšabo. Peresente ea leqhoa le bonts'itsoeng ke liphoofolo tse senotsoeng esale pele li fuoe ente ea epinephrine (0.1 mg / kg) kamora ho ithuta (71%) e ne e le kholo ho feta taolo e kenngoeng ka letsoai (44%) nakong ea lipuo tse tharo tsa CS ho 48- t tlhahlobo ea ho boloka (** P <0.01). Ntlafatso ea mohopolo e hlahisitsoeng ke systemic epinephrine management e ile ea fokotsoa haholo ha li-receptor tsa AMPA li ne li koetsoe NTS le CNQX (1.0 μg) pele tsoho e eketsoa ka epinephrine (** P <0.01). Ho ne ho se na phapang liperesente tsa ho hatsela ho bonts'itsoeng ke lihlopha life kapa life tse pepesitsoeng pele tse fuoeng CNQX ho NTS ho tsoa liphoofolong tse laoloang tse fuoeng ente ea systemine ea letsoai. Liphoofolo tse mashome a mararo li arotsoe ka lihlopha tse latelang tsa kalafo (saline-PBS, n = 9; saline-CNQX, n = 6; epinephrine-PBS, n = 10; le epinephrine-CNQX, n = 5). (B) Liteko tsa ho boloka litšebeletso: Kerafo ea mohala e bonts'a leqhoa le lekang ho liteko tsa CS nakong ea liteko tsa ho boloka. Sehlopha se fuoeng epinephrine ea morao-rao ea koetliso (0.1 mg / kg) se bonts'itse leqhoa le phahameng haholo ho feta lihlopha tse ling tsohle nakong ea tlhahiso ea bobeli le ea boraro ea molumo CS (** P <0.01). Ntlafatso e hlahisitsoeng ke epinephrine mohopolong oa thuto e amanang le ts'oaetso e ile ea koaloa ke ho hanyetsa li-receptor tsa AMPA ho NTS le CNQX (1.0 μg). * P <0.05.
Puisano
Liteko tsena li ile tsa hlahloba hore na matla a ho tsosoa a susumelitsoeng ke mokhoa o sa tloaelehang oa thuto ea thuto a susumetsa mohopolo oa boemo ba tšabo ea Pavlovian. Liphuputso tse tsoang litsing tse tharo tsa liteko li senola hore mohopolo oa lipapali tse hlahisang molumo o matlafatsoa ka lihlopha tse hlophisitsoeng boemong bo feletseng ba libuka tse amanang le lihlopha tseo pele li neng li pepesoa ke mokhoa oa ho etsa lithuto 24 h pele ho tšoha. Thuto 1 e ile ea boela ea hlahloba hore na liketso tse molemo tsa ho tsosoa ho hlahisoang ke ho pepeseha ho likoti tsa koetliso e ncha li akarelletsa ho ts'ebetswa ha lihomone tse nang le kutloelo-bohloko. Ho finyella sena, sotalol e ne e laoloa ka thata ho thibela li-receptor tsa β-adrenergic tsa meriana tse kopanyang hormone e amanang le hormone epinephrine. Karolo e phahameng ea boitšoaro bo batafatsang bo hlokometsoeng ka lihlopha tse hlophisitsoeng boemong bo sa tloaelehang bo ile ba thibeloa ka ho thibela li-receptors tsena pele li sebetsana le sotalol. Litholoana tsa lithuto tsa 1 li bontša hore liphetoho tse susumelitsoeng ke ts'ebetso tse hlahisoang ke mohlolo oa maemo a boemo bo boima li akarelletsa ho thibela li-hormone tsa adrenal le liketso tse latelang tsa lihomone tsena ho li-receptor tsa β-adrenergic.
Tlhaloso ea ho fumanoa ha sena e ile ea atolosoa ho ithuta 2 ka ho etsa qeto ea hore na liphello tsa monate li kopanngoa joang ka ho ts'ebetsong ha li-brain-neurons tse amohelang ho fetoha ha li-hormone tsa 'meleng le ka kutloelo-bohloko. Hona hormone ea epinephrine e ikamahanya le li-receptor tsa β-adrenergic hammoho le li-vaginal nerve fibers (Lawrence et al. 1995) tse nyolohelang bokong ba bokhelohi le synapse holim'a li-neurons NTS (Kalia le Sullivan 1982). Liphetoho tse bakoang ke ho tsoaloa ha likokoana-hloko tsa adrenal hormone ho nyoloha ha li-vagal afferent fibers (Miyashita le Williams 2006) le tsona li thabisa NTS neurons ka ho lokolla glutamate ho li-terminal (Granata le Reis 1983b; Allchin et al. 1994). Thuto 2 e ile ea hlahloba bohlokoa ba ts'ebetso ea ho tlosoa ha glutamate ho tswa ho vagal afferents ho NTS neurons ho rarolla liphello tsa ts'ebetso ka mohopolo. Liphuputso tse tsoang phuputsong ena li bontšitse hore ho eketseha ha mali ho hlahlojoa nakong ea teko ea ho boloka molumo litabeng tse koetliselitsoeng kamoreng ea boemo ba morao li ile tsa thibeloa ka ho thibela li-receptor tsa AMPA ho NTS le mohanyetsi ea khethiloeng oa glutamate CNQX.
Ke habohlokoa ho hlokomela hore le hoja boemo ba liteko bo sebelisetsoang ho hlahisa mokhoa o tsitsitseng litsing tsa liteko 1 le 2 li thusa ho boloka boitsebiso bo tloaelehileng ba ho kopanya likhallelo tsa morao-rao ka morao-rao, tlhahiso e ncha e nang le tšusumetso e tebileng ea tikoloho e 'nile ea hlokomeloa ho hlahisa liphetoho tse fapaneng ka mokhoa oa ho hlahloba maikutlo. Ho etsa mohlala, ho behoa sebakeng se sa tloaelehang ka ho kopana le thibelo, thibelo le ts'ebetso ea mohatla o kenyelletsoeng, ka pel'a kamo e lokolohileng kapa sethaleng se phahameng se khanyang, ho sitisa ho kenngoa ha LTP, bokhoni ba ho qhoqhoa, le mohopolo oa sebaka ho ithuta (Diamond et al. 1990, 1994; Xu et al. 1997; Akirav le Richter-Levin 1999; Diamond le Park 2000). Phapang pakeng tsa mohopolo le synaptic plastiki e hlokometsoeng liphuputsong tsena ho amana le ba tsebang ho hopola mohopolo ka ho sebelisa nako e khutšoanyane ea tlhaho e sa tsitsang ea tlhaho (Kinney le Routtenberg 1993; Vankov et al. 1995; 2000, 2001; Viola et al 2003; Li-al, X; le; Davis et al. 2000; Moncada le Viola 2003; Sierra-Mercado le al. 2003) e ka amahanngoa le boholo ba tsoalo ea ho tsosoa le khatello ea maikutlo e bakoang ke khatello ea maikutlo ka maemo a fapaneng a koetliso.
Leha ho le joalo, ha ho pepeseha hore ho na le maemo a sa tloaelehang a hlahang ka mokhoa o khutšoanyane ho hlahisa mokhoa o lekanyelitsoeng oa ho tsosa li-hormone tsa adrenal, joale tsamaiso ea epinephrine ho lihlooho tse tloaelehileng e lokela ho eketseha ho isa tekaneng le e hlahisoang ke boemo ba Pavlovian boemong bo neng bo le ncha. Sena se ile sa lekoa thutong ea ho qetela ka ho hlahloba hore na maemong a mangata a matla a ho qhoqeshisoa ke tšabo a bontšoa tlhahlobo ea 48-h ea ho boloka boiphihlelo litabeng tse hlalositsoeng pele ho tlhahlobo ea epinephrine (0.1 mg / kg) mabapi le pele -olaletso e bontšitsoeng feela e bontšang boitšoaro bo bobebe ba ho qhoqhoa ha liteko litsong tsa 1 le 2. Liphuputso tse tsoang ho 3 li hlalositse hore lihlooho tse qalang ho pepesa tse filoeng epinephrine post-conditioning li bontšitse karolo e kholo haholo ea boitšoaro bo hloekisang nakong ea lipontšo feela tsa tlhahiso ea tlhahlobo ea 48-h ho feta ho laola pele ho pelaelo. Ntlafatso ea mohopolo oa epinephrine e bontšitsoeng ka karolo e phahameng ea boitšoaro bo hlephileng e ne e thibisoa ke ho sitisa ho phahama ho potoloha pakeng tsa vagus nerve le bokhoni ba kelello ka ho thibela li-receptors tsa posttnate tsa glutamate NTS. Ho ne ho se na phapang karolong ea palo ea ho qhoqhoa ha CS-lipakeng pakeng tsa litsamaiso le sehlopha se fuoeng epinephrine tsamaiso le mohanyetsi oa litho tsa glutamate CNQX NTS. Liphuputso tse akaretsang li bontša hore ho pepesehela taba ea libuka ho eketsa tsoho ea 'mele,' me liphetoho tsena li ama matla a boemo ba Pavlovia ka ho susumetsa tsamaiso ea lihomone tsa 'mele.
Liphuputso tse fetileng li bontša hore lits'ebetso tse 'maloa tsa ts'oaetso tse kang tsoalo ea pelo le ts'oaetso ea mali li eketseha ka mor'a hore ho pepesehe sekhetho se secha (Carrive 2000). Ka mohlala, ho pepesehela lintho tse sa tloaelehang, tse kang ho qoelisoa metsing, ho sebetsana le maemo kapa ho kenya mohaho o mocha, ho fella ka ts'ebetso e ntseng e eketseha ea mohau-o motlakase o bontšoang ke libaka tse phahameng tsa epinephrine ka lero la mali (De Boer et al. 1990 ). Lipatlisiso li boetse li bontša hore likarabo tsena tse feteletseng tsa li-hormone li fetolelitsoe ka ho tseba lintho tse hlahang mohloling o sa tloaelehang ka ho pheta-pheta kapa nako e telele ho hlahisa maikutlo (De Boer et al. 1988; Konarska et al. 1989, 1990). Ho itšetlehile ka lipatlisiso tsena tsa tlhaho, lipatlisiso tsa hona joale li ile tsa khannoa ho hlahloba mokhoa oo lihlahisoa tse ncha li hlahisitseng tsoho ea 'mele li amang mohopolo.
Lintho tse fumanoang tlhahlobisong 1 e bontša epinephrine e ameha molemong oa tsoho e susumelitsoeng ke morao-rao ho ama matla a macha a macha a kenyelletsoang mohopolong. Boemo ba ho qhoqhoa bo bontšitsoeng ke liphoofolo tse sa tsejoeng pele ho tsona tse fanoeng ka ho pota-potiloe β-adrenergic receptor mohanyetsi oa sotalol li ne li lekana le maemo a ho qhoqhoa a bontšitsoeng ha CS e ne e hlahisoa bakeng sa liphoofolo tse sa hlahisoeng ka letsoai tse sa hlahisoang ka letsoai. Pono ena e boetse e tšehetsoa ke liphuputso tse bontšang hore liphetoho tse bakoang ke ho tsosolosa ha liphallelo tse amanang le liphallelo tsa motlakase, ho eketseha ha likhahla tsa mali, le khatello ea mali li fokotsehile haholo ka ho thibela methapo ea methapo ea β-adrenergic (van den Buuse et al. 2001 ; van den Buuse 2002; Carrive 2006; Miyahsita le Williams 2006). Liphuputso tse tsoang litsong tsa 1 li bontša linaha tse tsosang takatso li ka laoloa ke mokhoa o ts'oanelang oa ts'oaetso, 'me tsoho e susumelitsoeng ka mokhoa o tsosang takatso e ama palo ea mohopolo.
Ke habohlokoa ho hlokomela hore tekanyo ea sotalol e khethiloeng bakeng sa thuto ea hona joale e ne e sa fokotsa mohopolo oa liphoofolo tse laolang pele ho pelaelo, e fana ka maikutlo a hore tekanyo ena ea sotalol e ne e le tlase hoo e ka khotsofatsang li-ß-adrenergic receptors (partielle) le 1989 ). Ho se be le bothata leha e le bofe bo hlokomelehang ho lihlooho tse tšoaroang ka pele ho sotalol li ka amana le liphello tsa fatše tsa ts'ebetso e mongobo e hlahisoang ke koetliso e bonolo. Ka mohlala, 0.35-mA e boima haholo e sebelisitsoeng thuputsong ena e ile ea khetholloa e le boemong bo tlaase ka ho fetisisa ba ho tseba ho ba le lik'hilograma (Phillips le LeDoux 1992; Baldi et al. 2004). Ka hona, matla ana a ne a sebelisetsoa ho hlahisa maemo a bonolo a ho laola lihlopha ho hlahloba habonolo hore na tsoho e hlahisoang ke ts'ebetso e ncha e ntlafatsa thuto ea batho ba nang le ts'oaetso ho lihlopha tseo e seng tsa pele. Hoa utloahala hore litekanyo tsa koetliso tse hlahisang likarolo tse phahameng tsa boitšoaro bo hloekisang ho laola, ha e le hantle, li tla bontša hore ho thibela methapo ea β-adrenergic receptors le sotalol e hlahisa ho fokotsa ho ithuta. Leha ho le joalo, mofuta ona oa koetliso ea koetliso o tla pata liphetoho tsa ho ithuta le ho hopola mohopolo o hlahisoang ke keketseho e susumelitsoeng ke kenyelletso ea ho tsosa.
Liphuputso tse ngata li bontša hore hormone ea khatello ea kelello epinephrine e etsa hore ho be le mohopolo oa liketsahalo tse amang batho kapa liphoofolo. Liphello tsena li bakoa ke epinephrine e sebetsang ka ho toba metseng ea methapo ea β-adrenergic (Sternberg et al. 1986; Introini-Collison et al. 1992) le ka tsela e sa tobang ka li-NTS le LC neurons tse ka etsang hore ho se ke ha e-ba le ho ts'oaroa ha amygdala le hippocampus (Williams et al. 1998, 2000; Miyashita le Williams 2004). Tlhahlobo 2 e hlahlobile hore na keketseho e entsoeng ka tsela e ncha ea pone ea motlakase le ea hormone e ama phapang ea methapo e kholo ka ho eketseha ha motlakase oa li-vagal le li-neurons tseo li kopanelang ho tsona NTS. Tšebeliso ea vagus e ne e nkoa e le tsela ea putative ho tloha ha phetoho ea pheko ea vagus e na le litho tse ngata tse utloisang bohloko tse bontšang tšebetso e matla ka lebaka la ho thibela epinephrine (Shapiro le Miselis 1985; Coupland et al. 1989; Paton 1998a, b), 'me tsamaiso ea systemine ea epinephrine e eketsa menahano ea neural e hasanngoa le maqhubu a vagus hammoho le litefiso tsa ho thunngoa ka NTS neurons (Papas et al. 1990; Miyashita le Williams 2006). Liphuputso tsena li bontša hore methapo ea vagus e khona ho fetisetsa liphetoho tsa phekolo ea meriana ka mor'a hore sephiri sa epinephrine se bolokehe bokong ka lebaka la liphihlelo tse tsosang haholo.
Amino acid glutamate ke motsoako oa mantlha oa li-neurotransmitter bakeng sa ho buisana le puisano ea synaptic pakeng tsa li-vagal afferents le neurons tseo li lumellanang ka tsona kahare ea boko. Ka mohlala, ho tsosolosa ka ho toba ha likhohlopo tsa vagal ho nyoloha ho etsa hore ho be le keketseho e kholo ea likhahla tsa glutamate tse nkiloeng ho NTS (Granata le Reis 1983b; Allchin et al. 1994). Tumellanong le liphuputso tsena, liteko 2 li bontšitse hore ho thibela litho tsa glutamate NTS le mohanyetsi CNQX ho thibela ntlafatso ea mehopolo e hlokometsoeng ka ho beha liphoofolo maemo a sa tloaelehang. Motsoako oa CNQX o neng o sebelisetsoa ho thibela li-receptors tsa AMPA ho NTS o khethiloe ka ho khetheha ho ba neng ba bontšitsoe pele ho thibela NTS neuronal ho thunya ho arabela ka ho tsosoa ha methapo ea vagus (Granata le Reis 1983a; Andresen le Yang 1990). Liphuputso tse tsoang litsing tsena li bontšitse hore ho hopola ho tsosolosa ho hlahisitsoeng ha tlhaho ho hlokomolohuoa ha synaptic puisano pakeng tsa vagal afferents le brainstem neurons NTS e sitisoa.
Ho eketseha ho tsosang takatso ea liketso tsa boipheliso tse fetisetsoang bokong ke meriana ea vagus e phetha karolo ea bohlokoa ho hlahisang liphetoho tse sebetsang le tse hlophisitsoeng liphuthehong tse nang le bokooa ba ho ithuta. Ka mohlala, liphetoho tsa synaptic li eketsa bokhoni ba ho bonahatsa matšoao a liphatsa tsa lefutso ka NTS, tse kang ho eketseha ha poleloana ea AMPA e nang le litlhoko tsa mohopolo le liphetoho tsa moralo ka synapse, ho hlaha ka matšoao a holimo a holimo le a ntseng a nyoloha a kang ho hatelloa ke khatello ea mali le ho tsosoa ha methapo ea matla (bakeng sa ho hlahloba, bona Kline 2008). Ho feta moo, mefuta e khethiloeng ea liketso tse ikemetseng tse ikemetseng, tse kang litoeba tse nang le khatello ea mali, li bontša liphetoho tse ngata tsa NTS tse kang likhahla tsa dendritic, keketseho ea karolo ea likhahla tse nang le GluR1 seboka sa AMPA, le ho eketseha ha polelo ea AMPA ea receptor mRNA ka hare ho NTS ha e bapisoa le litoe tse tloaelehileng (Aicher et al. 2003; Saha et al. 2004; Hermes et al. 2008). Ka mokhoa o ts'oanang, liphetoho tse khutšoanyane tsa khatello ea mali li tšoana le lihlopha tse matla tsa ho tsosoa ha maikutlo ho tloha ketsahalong ea maikutlo ho tlisa liphetoho tse hlophisitsoeng li-neurons tse bontšang ho eketseha ho ngoloa ha li-synapses tsa glutamaterg ka NTS. Ka lebaka leo, liphuputso tse kopanetsoeng li bontša hore liphihlelo tse tsotehang tse hlahisoang ke li-NTS ka ho lokolloa ha glutamate li ka emela mokhoa o mong oo liketsahalo tsa maikutlo li qalileng ho kenngoa ka oona 'me hamorao li sebetsanoe ke mehaho e meng ea limilimi ho ea ka nako e telele ea mohopolo.
Liphuputso tse 'maloa tsa boits'oaro li bontša phetoho e eketsehileng ea glutamate mehopolong ea NTS e ntlafatsang bakeng sa liphihlelo tse tsosang maikutlo. Ka mohlala, microinjecting glutamate ho NTS moo li-neurons tsa eona li lumellanang le li-vagal afferents li ntlafatsa mohopolo oa taba eo liphoofolo tsa laboratori li ileng tsa qetella li tsosoa ka eona mosebetsing oa ho qoba ho thibela metsi (Miyashita le Williams 2002; Kerfoot et al. 2008). Phuputso ea morao tjena e bontšitse hore phetisetso ea glutamatergic e thibelang matšoao a NTS le motsoako oa khethiloeng oa AMPA oa CNPA e thibela ho hopola ho tsosolosoa ho holimo ho se nang pele ho kamoreng ea maemo. Thuto ena e fetisa kutloisiso ea rona ea liphello tsa ho tsosa litsebo tsa ts'oaetso ka ho senola hore li-receptor tsa AMPA tse fanang ka nako e telele ka har'a NTS li fetisa liphetoho tsa 'mele ho tloha ho tsosoa ho ts'oaetso e ncha e ntlafatsang mohopolo o tšabehang oa ho tšoha.
Ka kakaretso, liphuputso tse tsoang litsing tsa 1 le 2 li fana ka tsosumetso e hlahisoang ke mochine e amanang le ts'oaetso e amanang le ts'ebetso ea bonyane ka ho susumetsa phetoho ea hormone ea meriana le ts'ebetso e ncha ea vagal / NTS e rarahaneng. Tlhahlobo ea ho qetela e ile ea etsoa ho lebisa tlhokomelo e tobileng pakeng tsa li-hormone tsa phallo tse lokollotsoeng ka mor'a ho tsosoa ha tlhaho e hlahisoang ke tlhaho le liphello tsa tsona ho li-NTS neurons tse bakoang ke boko bo tsebang ho feto-fetoha ha ponahalo ea boipheliso ea ponahalo. Ho finyella sena, liphoofolo tse sa tsejoeng pele li hlahile (li-nonaroused) li ile tsa koetlisoa mosebetsing oa tšabo oa tšabo ea Pavlovian ka mekhoa e ts'oanang le e sebelisitsoeng litsing tsa 1 le 2 ntle le hore lihlopha tse fapaneng li fumane li-injection post-conditioning tsa saline kapa epinephrine. Lintho tse fumanoang tlhahlobo 3 e fana ka maikutlo a hore ntlafatso ea mohopolo e hlokometsoeng ho arabela ho tsosoa ha motho e ncha e ka 'na ea akarelletsa ho boloka li-hormone tsa phallo. Phuputso ena e bonts'itse hore ho eketseha ha lisele tse nang le kutloelo-bohloko e nang le liente tsa epinephrine ho matlafatsa haholo maemo a tšabehang a tšabehang a hlokometsoeng ka lihlopha tse e-s'o pepese ka kamoreng ea boemo bo botle ka tloaelo. Liphetoho tse tsositseng moeeng tse hlahisoang ke epinephrine lihlahisoa tse e-s'o pepese li bile le litekanyetso tse phahameng tsa ho qhoqhoa ho CS tse neng li tšoana haholo le tse fumanoang liphoofolong tse sa tsejoeng pele li lekoa litsong tsa 1 le 2. Ho phaella moo, ha tsoho ea 'mele e eketseha ka ho sebetsana le boemo bo tšabehang bo tšosang (Hui et al. 2006), ho tsamaisa epinephrine kapa corticosterone ka mor'a ho ithuta mesebetsi e kang ho amoheloa ka ntho (Roozendaal et al. 2006; Dornelles et al. 2007), ho shebella letoto tsa Cahill le Alkire 2003), kapa ho fumana mohoete ka mokhoa o fapaneng (Introini-Collison le McGaugh 1988), mohopolo oa CS, sebaka sa lintho, li-slides tse nkiloeng, kapa moelelo oo ho fanoang ka oona ho ntlafatsoa ho ntlafatsoa. Liphuputso tse tsoang litsong tsa 3 li boetse li fana ka maikutlo a linaha tse ntseng li eketseha tsa mohopolo oa ho tsosa mofuthu. Ho feta moo, phuputso ena e bontša hore pherefine epinephrine e kenyelelitsoe ho ntlafatsa mohopolo oa ho hopola mohopolo ka mokhoa o ts'oanang le mocha ka hore o hloka mokhoa o tšoanang oa glutamatergic NTS. Ka lebaka la bopaki bo ntseng bo eketseha ba hore mekhoa e metle le ea poneletso ea adrenergic e sebetsa ho kopana ho tiisa maqhama a synaptic, liphuputso tsa morao-rao li totobatsa bohlokoa ba ho saena pakeng tsa vagus le tataiso ea NTS ho rarolla liphello tse molemo tsa ho tsosoa maikutlo maikutlong.
Lisebelisoa le mekhoa
Lihlooho
Likhoto tse mashome a robeli a metso e meraro tsa banna tsa Sprague-Dawley (275-300 g) tse fumanoeng ho Charles River Laboratories (Wilmington, MA) li sebelisitsoe litekong tsa 1 (n = 24), 2 (n = 29), le 3 (n = 30). Likhoto li ne li beoa ka bonngoe ka masakeng a polasetiki 'me li bolokiloe ka potoloho e tloaelehileng ea 12: 12-h khanya e lefifi e nang le mabone ho 7: 00am Lijo le metsi li ne li fumaneha ad libitum nakong ea phetoho e sa sitisoeng ea 7-d ho vivarium. Liteko tsohle li entsoe ho latela melaoana le litataiso tsa Komiti ea Ts'ebeliso ea Liphoofolo le Ts'ebeliso ea Univesithi ea Virginia.
Phekolo
Kampo e 'ngoe le e' ngoe e ile ea fumana ente ea atropine sulfate (0.1 mg / kg, ip, American Pharmaceutical Partners, Inc.) e lateloa ke 10 min hamorao ka sejo sa manonyeletso a sodium pentobarbital (50 mg / kg, ip, Abbot Laboratories). Ho ile ha etsoa sekhahla sa midline ea scalp, 'me 15-mm-length, lehlakoreng le nang le lehlakoreng le nang le lehlakoreng le lekaneng la tšepe ea cannulae (25.0 gauge, Likaroloana tse Nyenyane) li ile tsa kenngoa lipakeng tsa 2 mm ka holim'a NTS (AP: -13.3; ML: ± 1.0 ho tloha bregma ; DV: -5.6 ho tloha holim'a lehata) ho latela li-coordinate tse nkiloeng ho tloha atlas ea Paxinos le Watson (1986). Mokhoa oa cannulae le lithako tsa lehata li ne li khomaretsoe lehata le samente ea meno, 'me sekoti se koetsoe ka sutures. Stylets (15 mm, 00 likokoana-hloko tsa likokoanyana) li kenngoa ka cannulae e entsoeng ka jenete ho boloka patency ea kankere. Penicillin (0.1 mL, im, Fort Dodge Animal Health) e ile ea fanoa hang-hang ka mor'a ho buuoa hammoho le litsebi tsa li-analgesic (0.05 mL sc, Hospira, Inc.) ho fokotsa khatello ea lipapali. Likokoanyana li ne li lula ka kamoreng e laoloang ke mocheso bakeng sa bonyane 1 h ka mor'a ho buuoa 'me ba fuoa 7 d hore ba hlaphoheloe pele ho qala thuto.
Tsamaiso ea Microinjection
Rat e 'ngoe le e' ngoe e ne e thibiloe ka letsoho liropeng tsa liteko, li-stylets li ile tsa tlosoa, 'me tsa hlatsoa li-sindano tse bolelele ba limilimithara tse 17, tse 30 tsa gauge ka hare ho cannulae ea NTS. Ntlha ea nale ea ente e atolositse 2 mm ho feta botlaaseng ba "cannulae" ea tataiso. Lisebelisoa li ne li hokahane le li-syringe tsa 10-μL Hamilton ka peipi ea PE-20 (polyethylene). Pompo ea seringe e ikemetseng (Sage-Orion) e tlisitse 0.5 μL ea PBS kapa mohanyetsi oa AMPA receptor CNQX (1.0 μg; Sigma Aldrich) ho NTS ka nako ea metsotso e 60. Tekanyo ea CNQX e sebelisitsoeng phuputsong ena e khethiloe ho tsoa litekanyetsong tse fapaneng tse fokotsang ts'ebetso ea methapo ea kutlo ea NTS (Andresen le Yang 1990). Linaliti tsa ente li ne li bolokiloe ka har'a li-cannulae bakeng sa metsotso e meng e 60 kamora ho ts'oaetsoa ho netefatsa ho tlisoa ha lithethefatsi. Li-style li ile tsa kenngoa hape ka har'a li-cannulae, 'me rat e' ngoe le e 'ngoe ea fuoa ente ea ip ea saline kapa epinephrine (0.1 mg / kg).
Meriana e tloaelehileng
Lihlooho ts'ebetsong ea pele li fumane li-injection tsa systemic (ip) tsa saline kapa sotalol (4 mg / kg), 5 min pele ho kenngoa likamoreng tsa boemo ba maemo.
Lisebelisoa tsa boitšoaro
Lisebelisoa tse sebelisoang bakeng sa tšabo ea Pavlovian li ne li e-na le kamore ea boitšoaro ea Coulbourn (12 inches width × 10 bophara ba lisenthimithara × 12 lisenthimithara tse phahameng, mohlala oa H13-16) e neng e kenngoa lebokoseng le leholo le nang le molumo (28 inches width × 16 inches botebo × 16 bolelele ba lisenthimithara). Mabota a ka pele le a ka morao a kamoreng a ne a entsoe ka polasetiki e hlakileng ka mahlakoreng a tšepe a sa hloekiseng le mokatong oa tšepe o sa khaotsoeng oa tšepe. Boitšoaro bo hlephileng bo ngotsoe nakong ea tlhahlobo ea boitšoaro ka mochine oa lisebelisoa tsa thepa ea mochini (mohlala oa H24-61) oo sampuli e tsamaeang ka 'ngoe le 400 msec. Likamore tse neng li sebelisetsoa ho hlahloba ho boloka likhahla tse peli tsa molumo li ne li tšoana hantle le lisebelisoa tsa koetliso empa li fetotsoe hore li fapane ka ho feletseng le likamore tsa maemo a maholo 'me li ne li le ka kamoreng e fapaneng e arohaneng le laboratori. Likamore tsa maemo a ho hloekisa li ne li hloekisoa ka tharollo ea 10% ea joala ka mor'a hore li koetlisetsoe le ho hlahlojoa. Lisebelisoa tsohle tsa lisebelisoa tsa litekanyetso tsa boitšoaro li fumanoe ho Coulbourn Instruments.
Mekhoa ea boitšoaro
Boemo ba tšabo
Li-rats li ne li tsamaisoa ho tloha ho vivarium ho ea laboratoreng 1 h pele ho hlahloba boitšoaro. Ka letsatsi le leng pele ho boemo bo botle, likhoto li ne li tloaetse ho ea ka kamoreng e nang le 5 min ea tlhahlobo ea mahala. Liphoofolo tse abetsoeng boemo bo sa e-s'o hlahise le tsona li ne li isoa lekalateng empa li lula malapeng a tsona nakong eo sehlopha se neng se sa tsejoe se tloaelehile ka kamoreng ea boemo bo botle. Lihora tse mashome a mabeli a metso e mene hamorao liphoofolo lihlopheng tse qalileng ho pepesa kapa tse sa tsejoeng pele li kenngoa ka kamoreng ea boemo ba boemo bo botle. Metsotso e meraro ka mor'a hore likhoto li se li le teng, molumo oa sekhahla oa 30 (5 kHz, 75 db) CS o ne o hlahisoa mme o kenngoa ka lekhetlo le le leng ka 1-sec, 0.35-mA e tsitsitseng US. Karohano ea 60-sec intertrial e arohane ho tloha ho hlahisa molumo o latelang. Boemo bo ne bo e-na le lihlopha tse hlano tse hlabang.
Tlhahlobo ea ho lula
Liphoofolo li ne li tsamaisoa ka bobeli ho ea kamoreng ea ho hlahloba ka ho fapaneng le kamoreng ea boitšoaro ho hlahloba mohopolo oa mohala oa CS 48 h ho latela maemo. Phoofolo e 'ngoe le e' ngoe e ile ea fuoa nako ea pele ea 3 ea ho hlahloba kamoreng e ncha. Ka mor'a moo, mohala oa CS (5 kHz, 75 db) o ile oa hlahisoa ka metsotsoana ea 30 ha ho se na basebetsi ba Amerika. Karolo ea 30-sec intertrial e arohane ho fela ha molumo o le mong le ho hlahisa se latelang. Litlhahiso tse tharo tsa tonopo tsa CS li fanoe nakong ea teko ea ho boloka. Palo ea lihlooho tsa nako e bontšitse ho arabela ka lekhahla nakong ea tlhahiso ea mohala oa CS oo pele o neng o e-na le likoti tse neng o sebelisoa e ne e sebelisoa e le letšoao la ts'ireletso.
Tlhahlobo ea statistical
Mehato ea boits'oaro e tsoang mosebetsing oa maemo a tšabo e hlahisoa e le liperesente tse bolelang tsa nako ± SE likhoto tse sebelisitsoeng li sa sisinyehe nakong ea tlhahiso ea molumo. Papiso ea lipakeng tsa sehlopha bakeng sa boits'oaro bo batang bo lekantsoeng nakong ea liteko tsa ho boloka e entsoe ka ANOVA ea litsela tse peli e lateloang ke liteko tsa Fisher's post-hoc Liphapang tse ka tlase ho P <0.05 li ne li nkuoa e le tsa bohlokoa.
Histology
Ho netefatsa hore ho na le lisebelisuoa tse nepahetseng tsa lisebelisoa tsa lintlheng le sesebelisoa sa cannulae ho NTS kamora ho phethoa ha liteko, phoofolo e 'ngoe le e' ngoe e ne e sa ts'oaroe ke tharollo ea euthanasia Euthasol (0.5 mL, Virbac Corporation) le ho sebelisoa hampe ka 0.9% saline e lateloa ke 10% formalin. Boko bo ne bo bolokoa ka 10% formalin ho fihlela ho aroloa vibitome. Likarolo li ile tsa khethoa 60 μm tse teteaneng, tse khabisitsoeng ka li-slide tsa khalase, tse nang le chromium-aluminium, tse nang le li-cresyl violet. Libaka tsa lisebelisuoa tsa lisebelisoa tsa cannulae le lisebelisoa li ile tsa netefatsoa ka ho hlahloba lisebelisoa tse atolositsoeng tsa li-slide (Setšoantšo sa 1). Boitsebiso bo tsoang ho liphoofolo tse hlano bo ne bo sa kenyelletsoa tlhahlobo ea lintlha ka lebaka la ho behoa ha molumo o sa nepahalang.
Karolo e fetileng Karolong e latelang
lumela hore baa fokola
Re leboha Mokhatlo oa American Psychological Association Diversity Program ka Neuroscience bakeng sa ts'ehetso ea bona ea pele. Ho phaella moo, re leboha Erica J. Young, Erin C. Kerfoot le Sumi Park ka menehelo ea bona ea bohlokoa. Lipatlisiso li ne li tšehetsoa ke National Science Foundation (NSF-0720170 ho CLW).
Karolo e fetileng Karolong e latelang
Mongolo o botlaaseng ba leqephe
*
↵1 Mongoli ea tšoanang.
E-mail [imeile e sirelelitsoe];; fax (434) 982-4785.
*
Sehlooho se fumaneha Inthaneteng ho http://www.learnmem.org/cgi/doi/10.1101/lm.1513109.
*
o Fuoe June 16, 2009.
o O amohetse July 31, 2009.
* Copyright © 2009 ke Press Cold Spring Harbor Laboratory
Karolo e fetileng
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